Catalog of Hymenoptera in America North of Mexico

Carl F. W. Muesebeck

This catalog is dedicated to our cherished colleague with affectionate regard
for his kindliness and with admiration for his distinguished scholarly contributions to

our knowledge of North American Hymenoptera for more than half a century.

Catalog of
Hymenoptera in America
North of Mexico

Prepared cooperatively by
specialists on the various groups of Hymenoptera
under the direction of
Karl V. Krombein and Paul D. Hurd, Jr.
Smithsonian Institution
and
David R. Smith and B. D. Burks
Systematic Entomology Laboratory,
Insect Identification and Beneficial Insect (eeeedincne Institute
Science and Education Administration,

United States Department of Agriculture

VOLUME 2
Apocrita (Aculeata)
SMITHSONIAN INSTITUTION PRESS

Washington, D.C.
1979

Library of Congress Cataloging in Publication Data
Main entry under title:

Catalog of Hymenoptera in American north of
Mexico.

“An outgrowth of... Hymenoptera of America
north of Mexico, synoptic catalog (1951) in-
cluding the first and second supplements (1958,
1967)”

Includes index.

CONTENTS: v.1. Symphyta and Apocrita
(Parasitica). —v. 2. Apocrita (Aculeata)

1. Hymenoptera— North America. 2. Insects —
North America. I. Krombein, Karl V.

QL567.1.A1C37 595.7'9'097 78-606008

Contents

Page

Introduction by Karl V. Krombein, Paul D. Hurd, and David R. Smith.................ccee Vv
Hymenoptera by, KarliVe Krombeinjand) Paul Dy Eirdyelirs cere sesccrerecess eres ceceenenittee cece eeencees 1
Symphyta’by, Davids Srmithsccce seers sects cceneseseeccercste cece eoecaeeceas tee nee cee te eee eeees eee 3
IMegallodontoide@ arc. -. scictc.eceveceuessc cu ckcne caaeuattceudien escuuns -¢ tesco ese tenses cuuctesvscuecaucauecaeeceeececeeces a

DRG VC fests ca ccs vase cvate sus Coc euep oi vac toes seo: cov ekin dios Sesame atest Dean deeoncuutenev assets nleeade aetes ranseesees 7
Pam phn tlii d a@ 5 osc ccsveseascesoste ce <tecssreceune sons esasteadcnc side Gy tence d desu ee occ dase a uceethacetbncersetereats 10

Men threGin OL CAs. cc. ccs soscicscesvcseesecctsessse woe sta loceettace san tre seek cere htc N Me Nas teee se RRS See eee 19

Or 1 Ae eco oc. wcxascesnsnansssistcecniasacacosunseeschuna cease ts ctetaceen Sacet tenedt acectcesetes seentaete sa veacess 19
TAT OT Ceara secre on ae eae caer ae eee coe R REE n oat cee eae e ME Goce a eee RE eS 20
(Gian oie Esterase ane aeece i eiee eea pe er cae area eee acece rc anencn oad ncys ete oan rse ode eaacossae 26

LDS) oy aKa) s10 F2\ erentecedaaccoctescosaocee pe cece Ocee Cos eecee onic cas c-ec crete suaLER po seeno a eeer sO seeee scuencuaseee 29

Mem thie inna @ eee oss cae cess ve conte sc oeel ocev ee cn carseat ace ee dee SON ace see oe 39

SUFI COI C8e ns scccsencsckesssassess ssaudens cctvasedcavedsteceinaiceensu ceeuptsostesectse Ml: cctae Sazcae So. lode sate nseonumecoase: 125
Sybex de Cee ocenc cs sescane occas sc swasvs nce nsaseaesssss Sucvun atin cae eee Soe count Cnet coe eae Men een cas cncees 125

IS irigi(es fs FY \esesceopeee ser esec pe cee pce eee ebec a cea eeR eRe e Eero er cee sceereroi Le Roce eC sas meas onedttia sense Lene eencee 125
Maphyrildac vic... tees scasscScessvascassuccoeaue~ssnessaelesinscatsasiassevsescssut gun dem eben canta teare ooeeeose 130
OYUSS1GE Cec cess ceacscese cca ce rseesaues as doe taut Sa aesaaes stacks ceaneesette sient bade Set mUM IEDM RET deianacaee ose se seen ces 131

( Gf} 0) oVos Wo ret aa aeecenenaceccaaaucececbecoeecaaaaeBeeuceArecea doc a Suonictooa cr eccae eso sa ccaoaancsc0es sag eo sseecooacocddvOacecn 133
Cephi dae seciescese science scess une seueceasssssncs stvesc Vesa ss ckecee nie cenest Pala tet anaes wate ee coe es 133
Apocrita by KarllV = Korom beim’. jscscdee5)scese ses once sobcsevageetceessncese tea eh cece Me Caoek waco actnceae os ee eaee 139
Parasiticalby,rauliMaMarshvand Robert We Carl somtee-.sssce-cc-ce-cesecacssetcesateeieteessceees ee 141
Ichneumonoidea by Paul M. Marsh and Robert W. Carlson. .............:::ccssceesceeseereeeees 143
Braconidae by/Pauli Ma Marshtrrcce.cccccorceresccsrcecveceecaecestten cee seeene cen eee ie snes coro oo 144
AphidiidaeibysPauliMe Marsh yecscesscccteccecsoccsess-csce sos con cereneecoeetrencee ooten aacemteseackeestest= 295

Hy brizontidae bys aullMe Marshes .rscceccece-ssc-cecsecsreesocccresoree coset ee nce csesecttercascestaee 313
Ichneumonidae bysRobert Wa Carlson cescccceecesessscee neces ce ese eee ncenennee here eerenee 315
Stephanidaeiby;RobertiWer Carlsomis.-c.cc--cscctesccccttesccecesseactssesceeec ctence oueeatebaecane seen 740

Chal cidoideatbysGordoml Gord lipeceee cores cess ence eee 743
Torymidae (except Agaoninae) by E. E. Grissell...................::scccsesecceeeecceeeensceeeenees 748
Pteromalidae by JB sD ry Burks eyccscec ces cas accasceecerees soe cee sSees cee ccea soe ea eer nap ewan aCe eee es 768
Hurytomidae bys DB ui kShecvecsscstcscsscnccocencesesec coves cteesurees ons censoaecnMesceneredinescsvecensss 835

Chal cididae by sB SD RB rkspercsecesceecccccssee coerce eee area e rear eee coe oc ae cae Settee coeeeeee ee eteees 860

Meucospididae by iB. Do Burks: sic. os ccenscesnectes ceca teerecsenseesecessceess cance steeceen eee 874

Mucharitidde by, Be MBUrks etree cose ae oee eee oan een 875
Eupelimidae bys BaD i Burkis\sscrcccwsters rons teres secrete au ste coe Sasa cade ate noe ee 878
Emeyrtidae bya Gord omni Gord hersesssteccr es ceceere eect ete rece eter reee earn eee 890
Hulophidaciby By DAIBuUrks i..201.01..zsccceecaesdewen coccicst aces as ot cece uscestees ssaceuns Coot sesso eae enen eee 967
Mymaridae byiB cD WB Ur ks sessesee coe aca san nc oot eee root aes cers da eac en cose eeet eee 1022
Mrichogrammatidae) bys Ss UT Kstaeeee tee sececccecescceccer nee cescecnesstaees con nteesesrencetee tore: 1033
WnplacedMlaxaaniChalcidordealbysB Ds Burks seceeeseesetereece reste ere eee 1042
Cynipoidea'lb yaB WDA UKs ies eraser cacceeseacee cacnenee cea an ees ees eee an een oe 1045
Mb ea Nb Gh ieee ne eee ca sce cae ee ers eee eae eee SoS SELES oo RETO ee 1045
MGTOP COT EAC ose ow sees cesee ccc ceca eee eae seco ae cc sae Se see esau cunt SSeS wa R CLES Tae eRe ae 1046

1 Fea (0 E22 acy Re Ne EES PTA Re COPE RATE Ee ey eee eer ee mt ee Sart 1047

1 DAD CELOS Yb fc Fs Seer a Cera ec rene EC PE ere Ser cca Oe cane eer acorteaans ieee eee eos ere ntecacacieo 1052
WOR SUG AG ee sce cote s seca c ace seus eee s Se steno ace Setar Tene maces eso escent e ea eSe Ne NTE Ee 1058
(Gaia oF 10 FO escnaacecneessaceeeecaetenca oacsoe sacra cca a enon CSCU concen aSesotod anasaadeaceaebeor teense Sctcaeace sada uie pence 1060
Hvanioid carbyzhobent) Wal Carl sombesssncssesesseeeses reese eee ee 1109
SW UTADA oss os ec cee ca ace ce suice obese ee Sales ee SSR Hse nase See Se Sc gwaten SHEE reece REE ao EE ORE EESGETS 1109
NAN S CYC 8 Olaerce.catesewcsnceactiocunsedaatcrcersen tae sawscseosaastee succes wager eee sdaccs vse Mes tout cua lela ceeaaeeceeeene tela
(Gras temp bid a Ces reccertee vase Sevasccester saeco te - Saeco ee ees to aera cs RS See ae 1115
PelecinoideaibyiCarllkn Wa Muesebeck: -vccccccecc-cecccccescccsesccoceeecnscetsoerer commento eee 1119
IPGL OCIA GC esc veesce. Ssctace coeees a see cactis ven Woven vane Saeees te Soca tadee soe cvuue donc avvudsivewon cosa eee ete
Eroctotrupoides byg@ ar lil Wr MuUleseb ec kgeceesesseeeree eases ects aeeee eee rece senator es arerereeee 1121
WaniOrnii dae tres co s.crrsccc ree cece ccd aoa ate oer OT ee ts hE, Se 1122
FROPYONII CRC virsrsoeceee ns ck ieon eect cots leet cates cee TI EE Fs as eee a ES 1122
Fleloridae's:..1.. ee ee ee ee 1122
Proctotrupidaey.. v4.0 nee ee a eee eee iene so Re ieecees 1123
Diapriida ey. ce oes wets: 1127
Scelioni dae seca eres o eects dessecteors de Cont areca 2209 inn se, eR Ren ERS Sorc putreet 1150
Platypastrid a tet ice ern Weitere AEE ee a Ee ie Seta Iau
Ceraphronoidealbya@ aruba WelMiulesebeckeeesscesseseercetcese setae eee eee ee ee eee eee 1187
GCeraphronida ciated. coe tees ener ee eto 1187
Mecaspilidae:. erotik ree sere ee tes eee non nr pennes RIT TIE es ak 1191
Trigonaloideailby Robert*wWe Carlsoncrsescetcccccsscsecotsrceececsec oases eee eee ase aceneeeene 1197
Wriporvalli dae eee ie eee ers acess ae B eae ean ea eee Tee TE A sie c ae 1197
Aculeata by Karli Ve Kar ombeimn tr crrcettccecerceseee eee cccts coors tose dead eaten ee cece 19S
Bethyloideaiby-Karliv= Kr omnb ein ers ove cceeecececcescsoevcosescossccevece terrace erceeees 1203
Bethy lid aes. cesses cree eres PN ha Bees cians BAL ISR ER8 91288 2629 55 <8 SUR RR RR 5S SSS 1203
Scleropibbidae v1 n0veeel acess eh ne tee hea eaten ore as ds NO oe aces 1219
Chrysidi dae sere aed SS LETTE SESE AT Tes Soe 1220
DD VATA AC see Ree TES IDOI se SEs cerns Sen se SNEED 595 ere 1240
Erm bole mid a er .eeee ee ae eee a naine a ar kn hs ta ge ace 1251
Scolioidea’ by: Karl Vs Kirombeimeeececereavesctrrcctetecceon crete oe oe nT ee tee 1253
Dap dae vce eevee eee eae Te oe ND LE AE Soy 1253
Siérol omorphidae eee. sec ee ene 1276
Mirtillid aes .ovrewte scene ee ee TE NT OIE on eee 1276
Scoliida@yvess sree cr eeeaie reese, Te, Oe A ES OE ooo ovace ese 1314
Sapy cide ee ee as ee ee ee eee eee 1319
Formicordealbys David RSimit heres ssceeceeres cere cee sot eaee ences eee ee eee ee ceee cee 1323
OTIC AC Siero es ae cea ee Reece ee ee SR one ee ee 1323
Vespoidéa' by: Karl' Vs: Kromibeimn: sss s2re se ee cascode 1469
MiaSari da Gn niiiseicasec ee eee eee TO LE esc ctet ewes 1469
Bane mide reso Seer ET etree 1472
Boo) 0) (0 £2 Rene RP ERE CRE EE SCHEELE CEE RCO EECA EEC creche LE race Cacocosacctach Seco scosnecr Recess: 1510

vi

ompiloideaby Wan Vesktrombell tere ee 1523

HZOMMP UIA ae tes. erccrewraersea ner ence. cates eeeesieees cede eeat sas cenetrer crete shooter sists oe eee 1523
Ritopalosomaticae mers ccesrcctem tr ccstrccte steers oe cc nee rt ee: 1570
Sphecoidedibyaarlay., Kaombelm -.recseccttecerces eee ee 1573
PATNI ULI CLA aCe sewers eet tak ce hte AO BT oc eM ke ee, sO Ua 1574
Sphecidae..24-2e hhh OR: Oe ee eS SRS ee ee ee ee 1575
Bem phred onidaes. £2... tecverescctestessec ste tosceoe cn teteat Soacsaew ets ene se ee 1595
Berri dae: 20,- AI ee aN AS, SNE SCC AE. SR ee Re 1617
Crabronidaet:... tee) er. Ake RO A ng A PO nT eee 1650
Mellimid Aer Ries, Se TRO Ee, Pee NSM 2 ocosssxarst Hepiees e 1683
INivSSOMIG AC SMa. acee tte eee steers teat ss ee eM ern oe RCE at ae ee Die PENS 1684
Bhilanthidaetis- Gat peewee, We 8 Receet s B eeD .kee AEae 1720
ApoideatbysbauliD urd sort. 2. 3 tee oc ee ee Se 1741
Colletidae®.... £8 :santien ee SAE, BAO ee ae eee Sa ee 1748
Oxacidae: cue" OF. Ae Ee. eee A A ae) Pelee te 1770
PSVGOTI al Cire Satree 08 te | Pete ben, iene 2 25a sec fetsaleasPaeasnt ca vacvne Seca eetene eee 1772
lalla cibid de). eee satetes witch Meeert ss PEs PRIMI oo occ Gc sb Perea satus kes touostesceeeasnruoneee 1932
MGIC tia em. Ft Shae o ieeorersie eos BN 2 eR POMIROO 5occc5s ch ap chcvass a ssochceuce deci sees eutoee 1978
Mepachili dacs. tte Asia Ni, A Pere: Pee esc seed ios fos owas soe eeu 1981
PAM GH OPHOTIGA Cs. <teee$ taser tt eos Aenen saat ees: SB coos d os asseaacsok Gh Ae oe eee 2081
PNT LAG sense Bs ness raade so Se Naat, cae sae sae ae cec ee AAD Fase cass Seo sncdessavesdeaicsasede oo Ssusadee cook 2188

Institutional Affiliation of Contributing Authors

Smithsonian Institution: Paul D. Hurd, Jr., Karl V. Krombein, Carl F. Muesebeck

U.S. Department of Agriculture: B. D. Burks, Robert W. Carlson, E. E. Grissell,
Paul M. Marsh, David R. Smith

University of California, Riverside: Gordon Gordh

vil

Introduction

By

Karl V. Krombein,
Paul D. Hurd, Jr.,

and

David R. Smith

This catalog, which is an outgrowth of the
highly successful Hymenoptera of America
North of Mexico Synoptic Catalog (1951),
including the first and second supplements
(1958, 1967), provides simultaneously, by
use of computer technology (Krombein, Mello
and Crockett, 1974. Ent. Soc. Amer., Ann.
20 :24-29), a printed version as well as a
computer-queriable data base of the basic
systematic, biological, and morphological in-
formation on the order Hymenoptera as it
occurs in America north of Mexico. While the
format of the present catalog closely parallels
that of the original catalog, every effort has
been made to increase the information con-
tent to fulfill better the needs of the biological
community. This has been accomplished chief-
ly by the introduction of textbook prose at
most higher category levels, by the presenta-
tion of explanatory or descriptive comments
as appropriate, and by the inclusion of more
complete listings of citations to taxonomic,
biological, and morphological literature at all
hierarchical levels together with parentheti-
cal annotations concerning the content for
many of these citations. Similarly, the data
about hosts, parasites, prey, predators, and
pollen sources are covered more fully than in
the original catalog. Since these data have
been reported in various ways and under dif-
fering names in the primary literature, an

viii

attempt has been made to organize this in-
formation and present it, usually, alphabeti-
cally even though sometimes such data have
been recorded in the literature with scien-
tific or vernacular names or both. Although
many of these names have been checked for
accuracy, no consistent attempt has been
made to verify that the names agree with
current usage or status. While those authors
of zoological names are usually abbreviated
in the citation of hosts, parasites, and the
like, the authors of the scientific botanical
names are not included. Elsewhere in the
catalog, the last name of an author is cited
in full, but without initials. Since the tech-
nology employed in the preparation of this
catalog precluded the use of diacritical marks,
these have been omitted and consequently,
as in the case of the umlaut, a variant spell-
ing has been substituted as appropriate.

The International Code of Zoological No-
menclature (1961) including the intent and
preamble of the Code and of any pertinent
opinions usually has been followed. Thus a
name in current usage as deterimned by the
code is employed, but mention is made, as
appropriate, that an application is pending
before the International Commission of
Zoological Nomenclature.

An attempt has been made to record all the
recent taxa, with their synonyms, described

from, or known to occur in, the political di-
visions of the conterminous United States,
Canada, Alaska, and Greenland. Insofar as
known, all species introduced from other
countries and liberated in America north of
Mexico for biological control purposes have
been listed. Some of these have never been
recovered and, apparently, were unsuccessful
in establishing themselves. For each such
species a statement has been included, follow-
ing the distribution, that the species was
liberated but did not become established.

As in the original catalog, the arrangement
is systematic for species-groups and higher
categories insofar as our present knowledge
and the limitations of a linear arrangement
permit. The generic and subgeneric concepts
represented in this arrangement are based
upon what are believed to be the correct type-
species. In each instance the type-species is
cited together with the authority for the
selection. Where designations of type-species
have been found to be invalid under the Inter-
national Code of Zoological Nomenclature,
new type-species designations, believed to be
valid, are given in the catalog. Generic syn-
onymy is included under the generic head-
ings except where subgenera are recognized,
in which cases such synonymy is given under
the subgeneric names, and references to re-
visional or other papers are listed under the
appropriate higher category.

The arrangement of species within genera,
subgenera, and species groups is alphabetical.
Where subspecies are recognized, the sub-
specific names are placed in alphabetical or-
der under the species to which they belong;
and varieties are listed under the particular
species or subspecies in which they were
described. In each case the specific, subspeci-
fic, or infrasubspecific, name is followed by
an indication of the known distribution, and
by brief statements, as appropriate, of pre-
ferred habitats or the like, hosts, parasites,
prey, predators, or pollen sources. Much of

this information on synonymy, distribution,
ecology, hosts, parasites, prey, and so forth
has not been published previously. The type
localities are usually recorded for those forms
that are known only from the localities where
the type specimens were obtained. Otherwise
the distribution is usually shown by states
and provinces, or by other means such as
life zones.

Since unquestionably a catalog is indispen-
sable in the support of systematic and other
biological research, no effort has been spared
toward making this catalog as useful as pos-
sible to all of the scientific community inter-
ested in these fascinating insects.

LITERATURE COVERAGE

All authors have attempted to include all

pertinent references to synonyms, revisions,

taxonomy, biology, and morphology begin-

ning with 1758, the publication date of the

10th edition of “Systema Naturae” by Lin-

naeus. The cut-off dates vary for the several

sections of the catalog and are as follows:

Symphyta through 1974;

Ichneumonoidea through 1976;

Chalcidoidea—Torymidae (except Agaoninae)
and Encyrtidae through 1976; other fam-
ilies and Agaoninae through 1972;

Cynipoidea through 1972;

Evanioidea through 1976;

Pelecinoidea, Proctotrupoidea, and Ceraphro-
noidea through 1972;

Trigonaloidea through 1976;

Bethyloidea and Scolioidea through 1975;

Formicoidea through mid-1975;

Vespoidea, Pompiloidea, and Sphecoidea
through 1975; and

Apoidea through 1976.

All authors have included some references

subsequent to the dates listed above.

Abbreviations and Symbols

LITERATURE CITATIONS

The source for journal abbreviations is Whit-
lock, C., 1939, Abbreviations used in the De-
partment of Agriculture for titles of publica-
tions, United States Department of Agricul-
ture Miscellaneous Publication No. 337, 278
pages. Abbreviations for other titles and
journals not found in Whitlock essentially
follow the same format and abbreviations
that she recommends. Book titles are usually
shortened to omit irrelevant adjectives and
include abbreviations, e.g.: Wheeler and
Wheeler, 1963. The Ants of North Dakota,
p. —, is cited as Wheeler and Wheeler, 1963.
Ants of N. Dak., p. —. Certain lengthy non-
serial titles are also abbreviated, e.g.: Say,
1824. In Keating, Narr. Long’s 2nd Exped.,
v. 2 (App.), p. —, rather than Say, 1824. In
Keating, Narrative of an Expedition to the
Source of St. Peter’s River, Lake Winnepeek
., etc. The titles are intended to be uni-
form throughout the catalog, but, in a work
of this magnitude, there will naturally be
some deviations. The abbreviations should be
adequate to find the cited publication.

SYMBOLS AND ABBREVIATIONS

Certain symbols and abbreviations are fre-

quently used in this catalog. Though there

may be slight variations in some, they are

generally as follows:

(!)—lapsus or misspelling of a scientific
name.

“9” = 8 or “3” = 9—incorrect sex deter-
mination.

? (é misdet.) or ¢ (2 misdet.)—only one
of the sexes described belongs to the spe-
cies cataloged.

9 —female.
é —male.

%—worker.
U4 —soldier.

ab.—aberration.
app.—appendix.
cent.—central.

changed status—used after a species-group
name to indicate a rank different from that
previously accorded to it; not necessarily
the same as new status.

desig.—designated; e.g., in type-species de-
signation, “Desig. by Rohwer, 1911.”

e., east.—east, eastern.

emend.—emendation.

fasc.—fascicle.

fig., figs.—figure, figures.

h.—heft.

n. comb.—new combination; used after a
species-group name to indicate a new ge-
neric assignment.

n. name—new name; used after a genus-
group or species-group name to indicate
a substitute name for a homonym.

N. name—New name; used after a biblio-
graphic citation to indicate a previously
proposed name.

Nom. nud.—Nomen nudum.

n. S.—new series.

n. status—new status; used where a taxon is
here accorded a rank different from that
which it had previously.

N. syn.—New synonymy; used to indicate a
synonym newly proposed in this catalog.

n., no., north.—north, northern.

n.e., northeast.—northeast, northeastern.

n.w., northwest.—northwest, northwestern.

orig. desig.—original designation; used to
indicate type-species designation.

p., pp.—page, pages.

pl., pls.—plate, plates.

preocc.—preoccupied; used after a genus-
group or species-group name to indicate a
homonym.

pt.—part.

revised status—revised status; used to de-
note a taxon that has been removed from
synonymy.

ser.—series.

s., so., south.—south, southern.

s.e., southeast.—southeast, southeastern.

s.w., southwest.—southwest, southwestern.

sp., spp.—species.

ssp., Sspp.—subspecies.

subg.—subgenus.
transcont.—transcontinental.
v., vol.—volume.
var.—variety.

w., west.—west, western.

FAUNAL ZONES

Bor. Boreal

Canad. Canadian
Huds. Hudsonian
Transit. Transition
Austr. Austral

U. Austr. Upper Austral
L. Austr. Lower Austral
Alleghan. Alleghanian
Austrorip. Austroriparian
Carol. Carolinian
Sonor. Sonoran

U. Sonor. Upper Sonoran
L. Sonor. Lower Sonoran

GEOGRAPHICAL NAMES

Abbreviation Political Unit
Ala. Alabama
Alaska Alaska

Alta. Alberta

Ariz. Arizona

Ark. Arkansas
B.C. British Columbia
Calif. California
Canada Canada

Colo. Colorado
Conn. Connecticut
Del. Delaware
D.C. District of Columbia
Fla. Florida

Ga. Georgia
Greenland Greenland
Idaho Idaho

Ill. Illinois

Ind. Indiana

Iowa Iowa

Kans. Kansas

xi

Ky.
La.
Labrador

Maine
Man.
Mass.
Md.
Mexico
Mich.
Minn.
Miss.
Mo.
Mont.
N. B.
Nee:
N. Dak.
Nebr.
Nev.
Newfoundland

Que.
Fee le
Sask.
Ss. C.
S. Dak.
Tenn.
Tex.
Us
Utah
Vt.
Va.
Wash.
W. Va.
Wis.
Wyo.
Yukon

Kentucky

Louisiana

Labrador, Newfoundland
(Labrador)

Maine

Manitoba

Massachusetts

Maryland

Mexico

Michigan

Minnesota

Mississippi

Missouri

Montana

New Brunswick

North Carolina

North Dakota

Nebraska

Nevada

Newfoundland,
Newfoundland (insular)

New Hampshire

New Jersey

New Mexico

Nova Scotia

Northwest Territories

New York

Ohio

Oklahoma

Ontario

Oregon

Pennsylvania

Prince Edward Island

Quebec

Rhode Island

Saskatchewan

South Carolina

South Dakota

Tennessee

Texas

United States

Utah

Vermont

Virginia

Washington (state)

West Virginia

Wisconsin

Wyoming

Yukon Territory

TAXONOMIC AND NOMENCLATURAL CHANGES

The catalog contains one undiagnosed new genus also in the Ichneumonidae, Woldsted-
species in the Ichneumonidae, Pterocormus tius Carlson, type-species Bassus biguttatus
clasma Carlson, p. 521, proposed for the Gravenhorst, p. 719, proposed for Syrphoc-
taxon misidentified as Jchnewmon canadensis tonus Foerster sensu Dasch (1964).
Cresson by Heinrich (1961). The following new names are proposed to
The catalog contains one undiagnosed new replace preoccupied names:
Tenthredinidae
Pachynematus gamus Smith for Pachynematus graminis Marlatt (1896) ........ p. 58
Nematus attus Smith for Amauronematus dyari Marlatt (1896) ................ p. 68
Amauronematus peralus Smith for Nematus pectoralis Cresson (1880) .......... p. 80
Ichneumonidae
Oedomopsis davisi Carlson for Trophon ? nasutus Cresson (1868) .............. p. 366
Gelis cushmani Carlson for Hemiteles apantelis Cushman (1927) ................ p. 405
Oresbius shumaginensis Carlson for Stiboscopus ferrugineus Ashmead (1902) ....p. 438
Pterocormus dionymus Carlson for Ichnewmon anonymus Heinrich (1961) ...... p. 522
Casinaria affinisima Carlson for Casinaria affinis Walley (1947) ................ p. 635
Pteromalidae
Mesopolobus fuscipedes Burks for Platyterma fuscipes Ashmead (1896) ........ p. 816
Eurytomidae
Harmolita ovatella Burks for Harmolita ovata Phillips and Emery (1919) ...... p. 840
Eulophidae
Syntomosphyrum orgyiazele Burks for Tetrastichomyia orgyiae Girault (1916) ..p.1005
Diapriidae
Trichopria kiefferi Muesebeck for Diapria montana Kieffer (1906) .............. p.1147
Scelionidae
Trimorus contractus Muesebeck for Gryon flavipes Ashmead (1893) ............ p.1162
Ceraphronidae
Aphanogmus harringtont Muesebeck for Aphanogmus salicicola Harrington (1899) p.1190
Megaspilidae
Dendrocerus obscurellus Muesebeck for Atritomus californicus Kieffer (1906) ..p.1194
Masaridae
Euparagia richardsi Bohart for Psiloglossa simplicipes Rohwer (1909) .......... p.1470
Philanthidae
Cerceris bolingeriana Krombein for Cerceris bolingeri Scullen (1972) .......... p.1730
Anthophoridae
Triepeolus mitchelli Hurd for Triepeolus sublunatus Mitchell (1962) ........... p.2094
There are a number of other nomenclatural valid genera and species for each family and
and taxonomic changes. These are considered higher category. These nomenclatural and
of lesser bibliographic importance than the taxonomic changes are as follows:
new names, so tabulations of them are de- A number of generic transfers are made.

ferred to Volume 3 which will also contain They are usually cited in the text as xanti-
the indexes and a table of the number of anum (Saussure), n. comb. The authority

Xil

responsible for the transfer is the author
of that section unless the name of another
specialist is included.

There are also a number of instances where
a taxon formerly considered to be a species is
treated here as a subspecies of another taxon,
or where a taxon. formerly considered to be a
subspecies is now raised to specific rank. The
authority responsible for the change is the
author of that section unless the name of
another specialist is included. These are usu-
ally cited in the text as clavatum johannis
(Richards), n. status or alba Rohwer, n.
status.

The words—changed status—occasionally
follow the author of a species-group name.
This indicates that the taxon has a rank

different from that accorded it elsewhere. It
is not the same as new status for it reflects a
change which has already been published.

In a few taxa, the words—revised status—
follow the author of a species-group name.
This denotes a taxon which has been removed
from synonymy.

There are a few new synonyms at the
genus-group level and numerous new syn-

. onyms at the species-group level. These are

indicated by the abbreviation N. syn. follow-
ing the bibliographic citation of the new syn-
onym. As noted above, the synonymy is to be
attributed to the author of the section unless
the name of another specialist appears in
parentheses following the abbreviation N. syn.

Volume 3

It is intended that Volume 3 will contain
separate indexes to the taxa of Hymenoptera,
and to their hosts, parasites, prey, predators,
and pollen and nectar sources. Preparation of
the indexes has already begun, and we antici-
pate that the tapes for Linotron production
will be sent to the Government Printing Office

during 1978. We will also include in Volume
3 a tabulation of the number of valid genera
and species for each family and higher cate-
gory, and lists of the nomenclatural and
taxonomic changes other than the new names
which are listed above.

Acknowledgments

Preparation of the catalog and funding for its
publication have had the enthusiastic support
of Porter M. Kier, Director, National Museum
of Natural History (NMNH), Smithsonian
Institution (SI) and of Lloyd V. Knutson,
Chairman, Insect Identification and Beneficial
Insect Introduction Institute, U.S. Depart-

xiii

ment of Agriculture. The catalog in its printed
form could not have been achieved without
their help, and we are most grateful that
their assistance was available whenever we
required it.

We are indebted to a host of cooperating
hymenopterists for generous assistance which

has greatly enhanced the content and quality
of the catalog. The aid furnished has involved
such diverse activities as reviewing prelimi-
nary drafts of various sections, and providing
new information on taxonomy, synonymy,
distribution, and biology.

In the Symphyta, H. E. Milliron, formerly
of the Biosystematics Research Institute
(BRI), Agriculture Canada, Ottawa, re-
viewed parts of the manuscript and provided
information, as did H. R. Wong, Northern
Forest Research Centre, Edmonton, Alberta,
for Pristiphora Latr. H. Greenbaum, Uni-
versity of Florida, Gainesville, furnished
data on Florida sawflies.

The accuracy of host names in the Para-
sitica was checked by the following special-
ists: Smithsonian Institution—J. F. G.
Clarke, D. R. Davis, W. D. Duckworth, T. L.
Erwin, W. D. Field, and R. C. Froeschner;
Systematic Entomology Laboratory (SEL),
U. S. Department of Agriculture—D. C.
Ferguson, R. J. Gagné, R. D. Gordon, A. B.
Gurney, J. L. Herring, R. W. Hodges, J. M.
Kingsolver, J. P. Kramer, A. S. Menke, D. M.
Miller, L. M. Russell, C. W. Sabrosky, D. R.
Smith, T. J. Spilman, M. B. Stoetzel, E. L.
Todd, R. E. Warner, and R. E. White. D. M.
Weisman (SEL) identified the remains of
some lepidopterous larvae which served as
hosts.

Specialists who were helpful in the Ichneu-
monoidea were: L. E. Caltagirone, University
of California, Albany, and C. C. Loan (BRI),
who reviewed parts of the manuscript on
Braconidae; C. van Achterberg, Waarder, The
Netherlands, W. R. M. Mason (BRI), R. D.
Shenefelt, University of Wisconsin, Madison,
and R. Wharton, University of California,
Berkeley, who provided advice and informa-
tion on Braconidae; H. K. and M. C. Townes,
American Entomological Institute, Ann Ar-
bor, Michigan, who provided information and
advice on Ichneumonidae; and P. M. Marsh
(SEL), who proofed the first-phase computer
printouts for Mesochorinae, Diplazontinae,
Oxytorinae and Orthocentrinae, and the third-
phase edit for Ichneumoninae.

Z. Boucek, Commonwealth Institute of En-
tomology, London, and M. Graham, Oxford
University, provided much information on
Chalcidoidea, and D. P. Annecke, Plant Pro-

Xiv

tection Research Institute, Pretoria, South
Africa, advised on the placement of some
species assigned erroneously to Aphycus
Mayr. A special debt of gratitude is due C. F.
W. Muesebeck who painstakingly proofed all
edit phases of the computer printouts for
Encyrtidae and Torymidae and all but the
first-phase edits of all other families of
Chalcidoidea.

D. B. Krombein provided welcome assist-
ance by helping to proof the manuscripts and
printouts for all superfamilies of aculeate
wasps. F. D. Parker, Bee Biology and Syste-
matics Laboratory, Utah State University,
Logan, furnished biological data for a number
of Utah wasps.

We are grateful to R. M. Bohart, Univer-
sity of California, Davis, for reviewing the
manuscript on Chrysididae and for consider-
able other assistance which included informa-
tion on new synonyms and distribution in the
Elampinae and Chrysididinae, and the assign-
ment of taxa to species groups in Chrysis
L. H. E. Evans, Colorado State University,
Fort Collins, reviewed the section on Bethyli-
dae.

In the Scolioidea, the late J. C. Bradley,
Cornell University, Ithaca, New York, and
J. G. Betrem, Deventer, The Netherlands,
reviewed the manuscript on Scoliidae, and
H. W. Allen, Moorestown, New Jersey, that
on Tiphiinae. W. E. Ferguson, San Jose State
University, California, and C. E. Mickel, Uni-
versity of Arizona, Tucson, furnished infor-
mation on synonymy and taxonomy of some
Mutillidae.

R. R. Snelling, Los Angeles County Museum,
California, and M. R. Smith, Arlington,
Virginia, reviewed the section on Formicoi-
dea. J. F. Watkins II, Baylor University,
Waco, Texas, reviewed the section on Dory-
linae, and A. C. Cole, University of Tennessee,
Knoxville, that on Pogonomyrmex Mayr.
A. Francoeur, University of Québec, Chicou-
timi, provided information on the fusca
group of Formica L.

O. W. Richards, British Museum (Natural
History), London, and R. M. Bohart reviewed
the entire manuscript for Vespoidea. J. van
der Vecht, Putten, The Netherlands, recom-
mended the systematic sequence adopted in
the Eumenidae and reviewed the manuscript.

M. J. West-Eberhard, Universidad del Valle,
Cali, Colombia, and R. R. Snelling reviewed
the manuscript on Vespidae and supplied
data on taxonomy and biology. J. E. Gillaspy,
Texas A & I University, Kingsville, provided
information on Polistes Latr.

H. E. Evans reviewed the manuscript on
Pompilidae, and F. E. Kurezewski, University
of Syracuse, N. Y., contributed some prey
records.

R. M. Bohart and A. S. Menke generously
made available a copy of their manuscript,
“Sphecid Wasps of the World,’ which was
most helpful in assembling the section on
Sphecoidea; Menke, in addition, reviewed the
catalog manuscript for this superfamily.
Other specialists contributed data on the
groups mentioned after their names: R. E.
Coville, University of California, Berkeley
(taxonomy and distribution of Trypoxyloni-
nae); J. E. Gillaspy (Bembicinae); F. E.
Kurczewski (biology of Larridae and Cra-
bronidae) ; R. C. Miller, Cornell University
(taxonomy, distribution, and biology of
Crabronidae); W. J. Pulawski, Wroclaw,
Poland (taxonomy and distribution of Tachy-
sphex Kohl) ; and D. Vincent, University of
Maryland, College Park (taxonomy and dis-
tribution of Passaloecus Shuck.).

E. G. Linsley, University of California,
Berkeley, reviewed the entire manuscript on
Apoidea and offered valuable suggestions.
C. D. Michener, University of Kansas, Law-
rence, discussed with the author of that sec-
tion the biology and systematics of bees;
many of his suggestions have been incorpo-
rated in the classification adopted. The fol-

lowing specialists have cooperated by con--

tributing data on the groups specified after
their names: G. E. Bohart, Bee Biology and
Systematics Laboratory, Utah State Univer-
sity, Logan (taxonomy and biology of
Apoidea); W. E. LaBerge, Illinois Natural
History Survey, Urbana (taxonomy of An-
drenidae and Anthophoridae) ; U. N. Lanham,
University of Colorado, Boulder (taxonomy
of Apoidea) ; M. A. Lieftinck, Rhenen, The
Netherlands (taxonomy of Anthophoridae) ;
A. Loken, University of Bergen, Norway
(taxonomy of Apidae) ; T. B. Mitchell, North
Carolina State University, Raleigh (taxon-
omy of Apoidea) ; J. S. Moure, Universidade

XV

Federal do Parana, Curitiba, Brazil (taxon-
omy of Apoidea); F. D. Parker (taxonomy
and biology of Apoidea); J. G. Rozen, Jr.,
American Museum of Natural History, New
York, New York (taxonomy and biology of
Apoidea); R. R. Snelling (taxonomy of
Hylaeus F.); R. W. Thorp, University of
California, Davis (taxonomy and biology of
Apidae) ; P. H. Timberlake, University of
California, Riverside (taxonomy of Apoidea);
and T. J. Zavortink, University of San
Francisco, California (taxonomy of Antho-
phoridae).

C. W. Sabrosky (SEL) has been helpful to
all of us in the discussion of abstruse nomen-
clatural problems. J. F. Gates Clarke (SI)
participated patiently in philosophical dis-
cussions of a wide variety of subjects per-
taining to the catalog. G. C. Steyskal (SEL)
was the resource person for the grammar of
scientific names and other linguistic matters.

Computerization of the catalog and its pro-
duction by the computer-driven Linotron re-
quired the highly skilled technical expertise
of specialists in computer storage and appli-
cations. James F. Mello, formerly Chief of the
Data Processing (ADP) Program at NMNH,
carefully analyzed the 1951 Hymenoptera
Catalog and developed the data analysis
matrix which governed entry of information
into the computer. R. Creighton, Manager,
Information Retrieval and Indexing Division,
Office of Computer Services (OCS), SI, de-
vised the programs for editing, arranging,
querying, and displaying data from the man-
uscripts. J. J. Crockett, Manager, Software
Systems and Program Maintenance (OCS),
developed the program for conversion of the
computerized data to special magnetic tapes
capable of driving the Photo Typesetting
Unit, the Mergenthaler Linotron 1010, in the
Government Printing Office. T. G. Gautier,
Chief (ADP), and D. Bridge, Operations
Manager (ADP), maintained daily collabora-
tion with the Editorial Board in assignment
of clerical assistance, and production of the
edit phases, merge files, and SELGPO print-
outs.

Finally, we are most grateful for the care-
ful, accurate typing of the manuscripts for
computer entry by the corps of dedicated
clerk typists. R. M. Garlick served with the

program as principal typist from its inception
until his reassignment in mid-1976 as a com-
puter technician; he also trained the other
assigned typists. P. R. Brown is currently the
only typist assigned full time. Other typists
who worked for varying lengths of time

Xvi

during the six years of manuscript produc-
tion and computer entry were: L. E. Back,
L. M. Bybell, R. Cloyed, L. E. Hatton, M.
Monahan, L. G. Oliver, J. Peabody, P. A.
Sunkel, and M. F. Ward.

1199

Division ACULEATA

By KARL V. KROMBEIN

This divisional name is retained because of the substantial biological literature published on
the groups of Hymenoptera popularly called ants, wasps and bees. No clear-cut unambiguous
criteria exist by which one can separate Aculeata from Parasitica for there are annectent forms
in both divisions.

In North America we recognize about equal numbers of valid species-level taxa in the Acu-
leata and Parasitica (or Terebrantia). However, there are comparatively few undescribed Acu-
leata, and subsequent revisionary studies probably will synonymize nearly as many taxa now
considered to be valid as there will be new taxa described. Undoubtedly there are numerous un-
described small Parasitica.

Aculeata occur in all major zoogeographic regions and on many of the oceanic islands; they are
absent from Antarctica. Brothers (1975) recognizes 38 families of Aculeata. The majority occur
in America north of Mexico except the Plumariidae, Scolebythidae, Loboscelidiidae and Fide-
liidae (sometimes placed in Megachilidae), all small families with very few species. The Cleptidae
are here considered to be a subfamily of Chrysididae. The exotic Loboscelidiidae are best con-
sidered as an extremely aberrant subfamily of Chrysididae allied to the Amiseginae. Brothers
considered the Crabronidae, here treated as a family, to be a subfamily of Larridae.

Brothers recognized only three superfamilies of Aculeata, placing the scolioid, pompiloid and
vespoid families in the Vespoidea, and consolidating the Sphecoidea and Apoidea under the
former name. Further discussion of Brothers’ arrangement will be found under appropriate su-
perfamily headings.

In general the Hymenoptera included in the Aculeata are characterized by conversion of the
ovipositor to a stinging function only. The eggs are no longer exserted through the ovipositor as
in most Parasitica but through an orifice anterior to it. The ovipositor with associated poison
glands now serves several purposes, the temporary or permanent paralysis of the prey of wasps,
as a defensive mechanism in bees and some ants, and as an offensive mechanism in some ants.
However, annectent forms occur in some Proctotrupoidea of the Parasitica, and Bethyloidea and
Scolioidea of the Aculeata. In many higher Parasitica the wing venation and thorax are much
more reduced than in the Aculeata.

Biologically the majority of Aculeata may be distinguished by their non-parasitic habits and
the construction of nests for their young. Most higher wasps belonging to the Vespoidea, Pompi-
loidea and Sphecoidea are predaceous upon other arthropods and build nests which vary from
simple to quite elaborate. A few species of Pompiloidea and Sphecoidea behave as parasitoids,
paralyzing the prey, laying an egg upon it, and making no nest; the prey later recovers and leads
a normal life until killed by the growing larva. Most of the vespoid Masaridae and all of the
free-living bees have converted to a larval diet of pollen and nectar. Cleptoparasites or
brood-parasites, whose larvae develop in the nests of other wasps or bees, have evolved inde-

1200 Hymenoptera in America North of Mexico

pendently a number of times in all aculeate superfamilies. However, these biological distinctions
break down in most of the more primitive wasps belonging to the Bethyloidea and Scolioidea.
Many of these behave as true parasitoids in that the larval prey may be only temporarily para-
lyzed, occasionally several eggs may be laid on a single prey larva, and frequently no nest what-
ever is made, the prey being left in situ, or at most a crude cell may be constructed around the
subterranean prey as in most Tiphiidae and Scoliidae. Parasitism of the egg stage of the host is
known only among the Amiseginae (Chrysididae). Polyembryony is unknown, but parthenogene-
sis occurs in some Aculeata. Usually this is of the facultative kind as is found in social insects
such as some ants, vespid wasps and honeybees. It may be obligate in some aculeates, such as
the tiphiid wasp Methocha and some species of the bee genus Ceratina, where males are rare or
unknown.

The simplest kind of nest among the aculeates is made by the wasp dragging the paralyzed
prey into a crevice in or above ground or back into the prey’s burrow; the opening is usually
sealed off by particles of the substrate to make a crude cell. A second type of nest is also made
in a pre-existing cavity, such as borings of beetle larvae in wood or twigs, or in old insect galls or
abandoned mud cells. The nest in this second type may be unicellular as in the first kind of nest,
or it may consist of a linear series of cells, each cell separated from its neighbor by a partition of
mud, wood chips, resin, masticated plant leaves, or other substances. A third kind of nest is ex-
cavated by the wasp or bee in the ground, in rotten wood, or in the soft pith of such shrubs as
sumac and elderberry. The subterranean nests are frequently unicellular but multicellular nests
in the ground, rotten wood or pith may have the individual cells arranged in a linear series or in
clusters with the individual cells sealed by a partition or closing plug of the substrate. Occa-
sionally a mud turret may be constructed over the entrance of subterranean nests. Next, there
are the nests constructed entirely from foreign materials. Usually these are above ground
although some Vespidae and Bombinae have subterranean nests. The nests of solitary species
may be made of mud, or a mixture of resin and pebbles, with the cells arranged in parallel tubes,
or in clusters or with separate but adjacent cells; some exotic social Vespidae make a mud en-
velope around combs of hexagonal cells. A number of social species make paper or carton nests
in which the nesting material consists of masticated wood fibers, bark or rotten wood. Finally,
there is the complex nest of the honeybees constructed from wax secreted from glands in the
abdomen of workers.

Aculeate larvae are normally cannibalistic if they come in contact accidentally. This tendency
is prevented in multicellular nests by the existence of partitions separating adjacent larvae.
However, some species make brood cells in which several larvae develop amicably without the
occurrence of cannibalism. Such nests have been reported for a few North American Isodontia
(Sphecidae) and Megachile (Megachilidae), and for many exotic Allodapini (Anthophoridae).

True sociality (eusociality) has arisen independently several times in the higher aculeates, the
Formicoidea, Vespoidea, Sphecoidea and Apoidea. All of the ants are eusocial or are social
parasites of other ants, but the majority of wasps and bees are solitary species. Wilson (1971)
considers that eusocial insects must possess three traits: “individuals of the same species
cooperate in caring for the young; there is a reproductive division of labor, with more or less
sterile individuals working on behalf of fecund individuals; and there is an overlap of at least
two generations in life stages capable of contributing to colony labor, so that offspring assist
parents during some period of their life.” Presocial insects exhibit one or two of the above traits.
Solitary species have none of these traits.

Most solitary aculeates practice mass provisioning, that is, the egg is laid and a store of food is
placed in the cell with it, then the cell is closed; in many species the store of food is provided be-
fore oviposition. Some sphecoid wasps, such as many Bembicinae (Nyssonidae), have taken the
first step toward subsocial status by adopting progressive provisioning. This behavior is charac-
terized by placing the egg on a single prey specimen and not furnishing additional prey until the
egg has hatched or by the hatching of the egg before any food is provided; after hatching the
larva is fed daily or at intervals as required. Other aculeates, e.g., Moniaecera (Crabronidae),
some Andrena (Andrenidae) and Exomalopsis (Anthophoridae), have achieved the higher level
of communal status, in which several females use a common burrow entrance but presumably
maintain separate cells. A higher level of presocial behavior (quasisocial) is found rarely in some
exotic bees where two or more gravid females of the same generation cooperatively construct
and provision the cells. Some of our Halictidae have attained the semisocial stage which is
similar to the quasisocial except that unmated females of the same generation associate with a

Division ACULEATA 1201

gravid female or females and care for the larvae of the latter. A semisocial colony may evolve
into a primitive eusocial colony as happens later in the season in some Au gochlorella nests when
the colony becomes monogynous and only the workers forage for pollen. Another stage toward
the eusocial is the subsocial in which one female cares for her own larvae as in many Allodapini
(Anthophoridae) and young nests of Bombus (Apidae); such a colony may later become truly
eusocial as happens when the first brood of Bombus workers ecloses and takes over the foraging
activities previously performed by the queen.

Taxonomy: Lanham, 1960. Ent. News 71: 85-86 (significance of hind tibial strigil in
classification). —Richards, 1972 (1971). Ent. Essays to Commemorate Retirement of Prof.
K. Yasumatsu, pp. 1-18, 10 figs. (thoracic spiracles in classification). —Brothers, 1975.
Kans. Univ. Sci. Bul. 50: 483-648, 101 figs., 7 tabs. (phylogeny, especially Mutillidae).

Biology: Walsh and Riley, 1869. Amer. Ent. 1: 122-143, figs. 96-112 (habits of wasps).
—Ashmead, 1894. Psyche 7: 19-26, 39-46, 59-66, 75-79 (habits of wasps). —Peckham and
Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 1-245, 14 pls. (instincts and habits of
solitary wasps in Wis.). —Hartman, 1905. Tex. Acad. Sci., Trans. 7: 15-85, 24 figs., also
published as Univ. Tex., Bul. 65, Sci. Series 6: 3-73, 24 figs. (habits of some Texan solitary
wasps). —Peckham and Peckham, 1905. Wasps social and solitary, 311 pp. (prey, nests, life
history in Wis.). —Rau and Rau, 1918. Wasp studies afield, 372 pp., 68 figs. (nests, prey,
life history in Mo.). —Wheeler, 1919. Amer. Phil. Soc., Proc. 58: 1-40 (evolution of parasitic
Aculeata). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 1-44 (prey, nests, ecology of Mo.
wasps, bees, ants). —Wheeler, 1923. Social Life Among the Insects, 375 pp., 113 figs.
—Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 157-277, 8 pls. (ecology of wasps and bees
nesting in clay bank in Mo.). — Williams, 1928. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Bul. Ent. Ser. 19: 30-60, 112-174 (tropical wasps and bees). —Wheeler, 1928. Social Insects,
378 pp., 79 figs. —Rau, 1928. Acad. Sci. St. Louis, Trans. 35: 325-489, 68 figs. (behavior
non-social wasps in Mo.). —Reinhard, 1929. The witchery of wasps, 291 pp., 14 pls., 10 text
figs. (prey, nests, life history in Md.). —Rau, 1933. Jungle bees and wasps of Barro
Colorado Island, 324 pp. (nests, prey, life history). —Iwata, 1942. Tenthredo 4: 1-46, 5 pls.,
1 fig., 2 pp. unnumbered figs. (compar. studies behavior of solitary wasps). —Hurd, 1955.
Century of Progress in Natural Sciences, pp. 573-575, Calif. Acad. Sci. (history of wasp
taxonomy). —Cooper, 1957. Jour. Expt. Zool. 134: 469-514, 26 figs. (functions of cell
partitions in preventing parasitism, predation and cannibalism, and in orienting larva for
pupation). —Evans, 1958 (1956). Tenth Internatl. Congr. Ent., Proc. 2: 449-457 (evolution
of social life in wasps). —Olberg, 1959. Das Verhalten der solitaeren Wespen
Mitteleuropas, 402 pp., 779 photos. (prey, nests). —Evans and Linsley, 1960. South. Calif.
Acad. Sci., Bul. 59: 30-37 (sleeping aggregations of aculeates). —Michener, 1961. Roy. Ent.
Soc. London, Symp. 1: 43-56 (aspects of social polymorphism). —Krombein, 1962. Ent. Soc.
Wash., Proc. 64: 11-19 (parasitism of several wasps and bees by acarid mites). —Linsley,
1962. Ent. Soc. Amer., Ann. 55: 148-164, 9 figs. (sleeping aggregations). —Evans, 1962.
Evolution 16: 468-483, 6 figs. (evolution of prey-carrying mechanisms in wasps). —Evans,
1963. Wasp farm, 178 pp., 25 pls., 16 text-figs. (popular account). —Hamilton, 1964. Jour.
Theoret. Biol. 7: 1-52 (genetic evolution of social behavior). —Evans, 1966. Ann. Rev. Ent.
11: 123-154, 2 figs. (behavior patterns of solitary wasps). —Krombein, 1967. Trap-nesting
wasps and bees: Life histories, nests and associates, 570 pp., 29 pls., 2 text-figs. Cited in
text as Trap-nesting wasps and bees. —Andrewes, 1969. The lives of wasps and bees, 204
pp., 16 pls., 15 text-figs. (popular account). —Evans and Eberhard, 1970. The wasps, 265
pp., 122 figs. (synthesis of data on life history, behavior, ecology). —F landers, 1970. Canad.
Ent. 102: 898-905 (cannibalistic infanticide in social Hymenoptera). —Wilson, 1971. Insect
societies, 548 pp., figs. (synthesis of insect sociology). —Iwata, 1972 (1971). Evolution of
instinct--comparative studies of Hymenoptera behavior, 503 pp., 50 figs. (in Japanese).
—Michener and Lin, 1972. Quart. Rev. Biol. 47: 131-159 (evolution of sociality).
—Spradbery, 1973. Wasps: an account of the biology and natural history of solitary and
social wasps, 408 pp., 28 pls., 131 text figs. —Hamilton, 1973. Ann. Rev. Syst. Ecol. 3:
193-232 (altruism in social insects). —Cazier and Linsley, 1974. Amer. Mus. Novitates 2546:
1-20, 6 figs. (foraging behavior of bees and wasps on Kallstroemia). —Schmidt et al.,
1974. Sozialpolymorphismus bei Insekten, 974 pp. —Trivers and Hare, 1976. Science 191:
249-263, 7 figs., 6 tabs. (haplodiploidy and evolution of social insects). —Iwata, 1976.

1202 Hymenoptera in America North of Mexico

Evolution of Instinct: Comparative Ethology of Hymenoptera, 535 pp., frontisp., 50 figs.
(English translation, Natl. Tech. Inform. Serv., PB 257052).

1203

Superfamily BETHYLOIDEA

By KARL V. KROMBEIN

Family BETHYLIDAE

North American members of this family are tiny or small wasps, rarely over 10 mm long.
They are considered to have been derived from a very primitive aculeate stock, and in some fea-
tures of behavior and development are more like the typical Parasitica rather than the Aculeata.
Usually the female stings the host larva a number of times until it is completely and per-
manently paralyzed. The host larva is left in situ or it may be dragged to a crevice. The bethylid
female usually deposits several eggs on each host specimen and the gregarious bethylid larvae
develop externally. The Bethylinae and a few Epyrinae prey upon lepidopterous larvae, prin-
cipally borers and seed-feeders, but also case bearers and leaf rollers. Most of the other
bethylids prey upon coleopterous larvae (occasionally pupae), especially species dwelling in the
soil, boring in wood or infesting seeds. There have been a few unconfirmed reports of hymenop-
terous larvae serving as hosts.

Despite Evans’ recent numerous revisions of many of the genera of North American
Bethylidae, we must expect that intensive, specialized collecting and biological observations in
the future will enlarge our native fauna by as much as 50 percent.

Revision: Evans, 1964. Mus. Compar. Zool., Bul. 132: 1-222, 144 figs. (generic reclassification
of New World spp.).

Taxonomy: Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 27-77. —Kieffer, 1914. Das Tierreich,
Lief. 41, pp. 1-595. —Richards, 1939. Roy. Ent. Soc. London, Trans. 89: 297-344.

Biology: Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 16-20.
Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 402-409, figs. 42-44, 46 (female,
male thorax).
SUBFAMILY BETHYLINAE

Genus PROSIEROLA Kieffer

Prosierola Kieffer, 1905. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 243.
Type-species: E pyris ? nasalis Westwood. Monotypic.
bicarinata (Brues). Ga., Fla., Tex.; Mexico. Host: Laspeyresia caryana (Fitch); Desmia
funeralis (Hbn.) in leaf rolls.
Parasierola bicarinata Brues, 1907. Wis. Nat. Hist. Soc., Bul. 5: 100. @.

Biology: Doutt, 1973. Ent. Soc. Amer., Ann. 66: 486-487, 1 fig. (brood tending).

1204 Hymenoptera in America North of Mexico

Genus PARASIEROLA Cameron

Parasierola Cameron, 1883. Ent. Soc. London, Trans., p. 197.
Type-species: Parasierola testaceicornis Cameron. Monotypic.
Perisierola Kieffer, 1914. Das Tierreich, Lief. 41, p. 533.
Type-species: Parasierola gallicola Kieffer. Desig. by Muesebeck and Walkley,
1951.

Several species have been reared from microlepidopterous pests such as the pink bollworm
and the Oriental fruit moth. Males are uncommon and thelytokous parthenogenesis may be com-
mon, four gererations of females having been reared from an unfertilized female.

alutacea Kieffer. Nev. (Ormsby Co.).
Parasierola cellularis var. alutacea Kieffer, 1906. Berlin. Ent. Ztschr. 50: 254. 3.
breviceps (Krombein). Calif. (Tracy). Host: Myelois venipars Dyar, larva.
Perisierola breviceps Krombein, 1954. Pan-Pacific Ent. 30: 259. 9.
cellularis (Say). Pa., N. J., Md., Va., Ohio, Mich., Ind., Iowa, Mo., Nebr., Kans.; Mexico (Nuevo
Leon). Host: Pyrausta ainsliei Heinr.; Ancylis comptana fragariae (W. and R.).
Bethylus cellularis Say, 1836. Boston Jour. Nat. Hist. 1: 279.

Biology: Guajardo and Ortiz, 1966. Soc. Nuevoleon Hist. Nat., Bol. 1: 41.
distinguenda Kieffer. Calif.; Nicaragua.
Parasierola distinguenda Kieffer, 1908. In Wytsman, Gen. Ins., fase. 76, p. 14. 2. N. name
for the taxon misidentified as cellularis (Say) by Kieffer, 1906, Berlin Ent. Ztschr. 50:
254.
emigrata (Rohwer). Tex.; Hawaii. Host: Pectinophora gossypiella (Saund.).
Perisierola emigrata Rohwer, 1917. Insecutor Inscitiae Menstruus 5: 1. 2, 6.

Biology: Busck, 1917. Insecutor Inscitiae Menstruus 5: 3-5. —Bridwell, 1919. Hawaii. Ent.
Soc., Proc. 4: 21. —Williams, 1927. U. S. Dept. Agr., Tech. Bul. 19: 11.
gracilicornis Kieffer. Oreg., Calif., Idaho, Nev., N. Mex., Tex.
Parasierola cellularis var. gracilicornis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 254. 2.
punctaticeps Kieffer. Calif., Tex. Host: Laspeyresia caryana (Fitch); Acrobasis sp., A. caryae
Grt.; Pectinophora gossypiella (Saund.).
Parasierola cellularis var. punctaticeps Kieffer, 1906. Berlin. Ent. Ztschr. 50: 254. 9.

Biology: Rude, 1937. Jour. Econ. Ent. 30: 840-841. —Nichols, Pierce and Pinkney, 1950. U. S.
Dept. Agr., Tech. Bul. 1011: 14.

Genus GONIOZUS Foerster

Goniozus Foerster, 1856. Hym. Stud., v. 2, p. 96.

Type-species: Bethylus claripennis Foerster. Desig. by Ashmead, 1893.
Progoniozus Kieffer; 1905. Soc. Sci. Bruxelles, Ann. 29: 105.

Type-species: Perisemus floridanus Ashmead. Orig. desig.

Most species whose hosts are known prey on microlepidopterous larvae. North American spe-
cies have been recorded from a number of economically important hosts. Males may be produced
from unfertilized eggs.

Revision: Fouts, 1928. Ent. Soc. Wash., Proc. 30: 127-132.

brevinervis Fouts. Ohio, N. J. Host: Coleophora sp.; Aroga trialbamaculella (Chamb.).
Goniozus brevinervis Fouts, 1928. Ent. Soc. Wash., Proc. 30: 127, 128. 9, d.

castaneicolor Evans. “United States.”
Bethylus castaneus Kieffer, 1907. Berlin. Ent. Ztschr. 51: 295. 2. Preoce. in Goniozus.
Goniozus castaneicolor Evans, 1964. Mus. Compar. Zool., Bul. 132: 201. N. name.

Taxonomy: Evans, 1962. Breviora, No. 150, p. 1 (generic transfer).

clarimontis Kieffer. Calif. (Claremont).
Gonziozus clarimontis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 253. °.

Superfamily BETHYLOIDEA 1205

columbianus Ashmead. N. J., D. C., Va., Utah. Host: Ancylis comptana fragariae (W. and R.),
Rhyacionia buoliana (Schiff.).
Goniozus columbianus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 76. 3, °.
Perisemus minimus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 71. 9.
electus Fouts. La. (Bogalusa). Host: Rhyacionia frustrana (Comst.), R. buoliana (Schiff.), R.
spp.
Goniozus electus Fouts, 1928. Ent. Soe. Wash., Proc. 30: 128, 132. 2, d.
flavipes Fouts. Kans. (Junction City).
Goniozus flavipes Fouts, 1928. Ent. Soe. Wash., Proc. 30: 128, 130. 2, 6.
floridanus (Ashmead). Fla. (Jacksonville).
Perisemus floridanus Ashmead, 1887. Ent. Amer. 3: 76. 6, 9.
foveolatus Ashmead. Ont. to Fla., west to Sask. and Tex., Ariz. Host: Acrobasis caryae Grote;
Polychrosis viteana (Clem.); Coleophora malivorella Riley; Ancylis comptana fragariae
(W. and R.); Anchylopera nubeculana Clem.; Grapholitha molesta (Busck); Acrobasis
caryivorella Rag.; Gracilaria negundella Chamb.; Nemapogon granella (L.);
Laspeyresia caryana (Fitch); Rhyacionia buoliana (Schiff.).
Goniozus foveolatus Ashmead, 1887. Ent. Amer. 3: 76. °.
Goniozus hortorum Brues, 1907. Wis. Nat. Hist. Soe., Bul. 5: 150. 9.

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99, No. 14: 44-46, pl. 15, figs. E, F
(male genitalia).
gallicola Fouts. Oreg., Calif. Host: Deoclona yuccasella Busck in Yucca seed pods; Melissopus
latiferreanus (Wlsm.)? in galls of Cynips maculipennis Gill.
Goniozus gallicola Fouts, 1942. Ent. Soc. Wash., Proc. 44: 168. 9.

Biology: Fouts, 1942. Ent. Soc. Wash., Proc. 44: 168 (host ?). —Gordh, 1976. U. S. Dept. Agr.,
Tech. Bul. 1524: 1-27, 5 figs. (behavior, life history, host).
hubbardi Howard. Fla. Host: Platynota rostrana (Wlkr.); Platoeceticus gloveri Pack.
Goniozus hubbardi Howard, 1885. In Hubbard, Ins. Affecting the Orange, p. 217. 9.
longiceps Kieffer. Tex. Host: Rhyacionia bushnelli (Busck), R. buoliana (Schiff.), R. spp.
Goniozus longiceps Kieffer, 1904. Arkiv for Zool. 1: 529. 9.
longinervis Fouts. Nebr., S. Dak., Calif. Host: Rhyacionia bushnelli (Busck), R. spp.,
Argyrotaenia citrana (Fern.).
Goniozus longinervis Fouts, 1928. Ent. Soc. Wash., Proce. 30: 128, 131. 2, d.
megacephalus Ashmead. Fla. (Key West).
Goniozus megacephalus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 74. 2.
Goniozus megalocephalus Schulz, 1906. Spolia Hym., p. 152. Emend.
mellipes (Ashmead). Fla. (Jacksonville).
Perisemus mellipes Ashmead, 1887. Ent. Amer. 3: 76. 9.
occipitalis Kieffer. Nev., Calif.
Goniozus occipitalis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 252. 9.
platynotae Ashmead. Conn., N. Y., to Fla. west to Kans. and La., also Utah and Ariz.; Puerto
Rico. Host: Platynota idaeusalis (Wlkr.); P. flavedana Clem.; P. stultana Wlsh.; Archips
argyrospila (Wlkr.); A. griseus (Robinson); A. rosaceana (Harr.); Archips sp.;
Coelostathma discopunctana Clem.?; Sparganothis sulphurana (F.); Canarsia sp.;
Argyrotaenia velutinana (Wlkr.); Pandemis limitata Rob.; Acrobasis caryae Grote;
Ancylis comptana fragariae (W. and R.); Agathodes designalis Guen. Parasite:
Perilampus fulvicornis Ashm. (secondary).
Goniozus platynotae Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 75. 3, 9.
Goniozus euliae Fouts, 1926. Ent. Soc. Wash., Proc. 28: 167. 2, d.

Biology: Fouts, 1928. Ent. Soc. Wash., Proc. 30: 129. —Woleott, 1948. Puerto Rico Univ. Jour.
Agr. 32: 848. —Bennett, 1961. Tenn. Acad. Sci., Jour. 36: 353. —Oatman and Jenkins, 1962.
Mo. Univ. Coll. Agr., Res. Bul. 789: 12. —Wilde and Semel, 1966. Jour. Econ. Ent. 59: 1040.
—Krombein, 1967. Trap-nesting Wasps and Bees, p. 69. —Prokopy, 1968. Jour. Econ. Ent.
61: 351. —Goertzen and Doutt, 1975. Ent. Soc. Amer., Ann. 68: 869-870 (female ovicidal
behavior).

1206 Hymenoptera in America North of Mexico

politus Ashmead. Md. (Virginia Beach).
Goniozus politus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 75. 9.

Genus BETHYLUS Latreille

Bethylus Latreille, 1802-03. Hist. Nat. Crust. Ins., v. 3, p. 315.
Type-species: Omalus fuscicornis Jurine. Desig. by Internat]. Comn. Zool. Nomencl.
Op. 153, 1944.
Perisemus Foerster, 1856. Hym. Stud., v. 2, pp. 95, 96.
Type-species: Perisemus triareolatus Foerster. Monotypic.
Episemus Thomson, 1862. Ofvers. k. Vetensk. Akad. Forh. 18: 452.
Type-species: Episemus variabilis Thomson. Desig. by Richards, 1939.
Digoniozus Kieffer, 1905. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 245.
Type-species: Perisemus oregonensis Ashmead. Monotypic.

Species of this genus prey upon larvae of Microlepidoptera, although there is a questionable

host record for a nitidulid beetle larva. Both sexes exhibit polymorphy in wing length, and no
fully winged North American specimens are known.

Revision: Evans, 1962. Breviora, No. 150, 12 pp., 3 figs., 1 map.
amoenus Fouts. N. W. T. south to Alta., east through Wis. and Ill. to N. Y., Maine, and N. S.
Host: Rhopobata naevana (Hbn.); ? Brachypterolus pulicarius (L.).
Bethylus amoenus Fouts, 1928. Ent. Soe. Wash., Proc. 30: 127. 2, d.
decipiens (Provancher). Alaska south to central Calif. eastward to N. Y., Maine and N. S.
Host: Cnephasia sp.
Gonatopus decipiens Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 179. @.
Perisemus oregonensis Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 70. @.
Arysepyris californicus Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 34. ¢.
Bethylus brachypterus Whittaker, 1928. Ent. Soc. London, Trans. 2: 385. °.
Bethylus flavicornis Whittaker, 1928. Ent. Soc. London, Trans. 2: 385. d, 2.

SUBFAMILY EPYRINAE
TRIBE EPYRINI

Genus RHABDEPYRIS Kieffer
Hosts of the American species are unknown. Two European species have been taken in ant
nests, but the association is probably fortuitous.
Revision: Evans, 1965. Mus. Compar. Zool., Bul. 133: 67-151, 7 pls.
Genus RHABDEPYRIS Subgenus RHABDEPYRIS Kieffer
Rhabdepyris Kieffer, 1904. Soc. Hist. Nat. Metz., Bul. (2)11: 32.
Type-species: Rhabdepyris myrmecophilus Kieffer. Desig. by Kieffer, 1906.

gracilis Evans. Calif.; Mexico (Durango).
Rhabdepyris (Rhabdepyris) gracilis Evans, 1965. Mus. Compar. Zool., Bul. 133: 76. 2, ¢.

mellipes Evans. Fla. (Orange Co.).
Rhabdepyris (Rhabdepyris) mellipes Evans, 1965. Mus. Compar. Zool., Bul. 133: 72, fig. 1.
OF

muesebecki Evans. Tex. (Brownsville); Mexico, Honduras, Costa Rica, Ecuador, Bolivia.
Rhabdepyris (Rhabdepyris) muesebecki Evans, 1965. Mus. Compar. Zool., Bul. 133: 74. 9°,
3.

nigripilosus (Ashmead). Ariz. (Huachuca Mts.); Mexico (Nayarit).
Mesitius nigripilosus Ashmead, 1895. Calif. Acad. Sci., Proc. (2)5: 539. °.
Rhabdepyris (Rhabdepyris) huachucae Evans, 1965. Mus. Compar. Zool., Bul. 138: 73. 2.

Taxonomy: Evans, 1965. Psyche 72: 268.

Superfamily BETHYLOIDEA 1207

Genus RHABDEPYRIS Subgenus TRICHOTEPYRIS Kieffer

Rhabdepyris subg. Trichotepyris Kieffer, 1906. In Andre, Spec. Hym. Eur. Alg., v. 9, p.
376,
Type-species: Rhabdepyris pallidipennis Kieffer. Desig. by Muesebeck and
Walkley, 1951.

SPECIES GROUP MEGACEPHALUS

angusticeps Evans. Ariz. (Tucson).
Rhabdepyris (Trichotepyris) angusticeps Evans, 1965. Mus. Compar. Zool., Bul. 133: 102,
figs. 8, 48. 9.
apache Evans. Ariz. (Pima Co.); Mexico (Sonora, Sinaloa, Nayarit, Jalisco).
Rhabdepyris (Trichotepyris) apache Evans, 1965. Mus. Compar. Zool., Bul. 133: 91, figs. 5,
ZO PrGr.
carolinianus Evans. S. C., Fla.
Rhabdepyris (Trichotepyris) carolinianus Evans, 1965. Mus. Compar. Zool., Bul. 133: 100,
figs. 7, 28, 46. 2.
megacephalus (Ashmead). Tex., N. Mex., Ariz., Calif.
Epyris megacephalus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 61. 9.
Epyris megalocephala Schulz, 1906. Spolia Hym., p. 153. Emend.
texanus Evans. Tex., Ariz.; Mexico (Sonora, Sinaloa, Morelos).
Rhabdepyris (Trichotepyris) texanus Evans, 1965. Mus. Compar. Zool., Bul. 133: 93, figs. 6,
LBW 29 OenCr-
werneri Evans. Ariz. (Gila Co., Pima Co.); Mexico (Chihuahua).
Rhabdepyris (Trichotepyris) werneri Evans, 1965. Mus. Compar. Zool., Bul. 133: 89, figs. 4,
17, 25, 44. 9, d.

SPECIES GROUP PULCHRIPENNIS

amabilis Fouts. Mass.,.N. Y., Md., D. C., Va., Fla., Ark., Mo.
Rhabdepyris amabilis Fouts, 1927. Ent. Soc. Wash., Proc. 29: 165. 9.

Genus RHABDEPYRIS Subgenus CHLOREPYRIS Kieffer

Chlorepyris Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 108.
Type-species: E pyris semiviridis Kieffer. Desig. by Kieffer, 1914.

SPECIES GROUP VIRIDISSIMUS

fulgens (Brues). Tex. (Brownsville); Honduras.
Epyris fulgens Brues, 1907. Wis. Nat. Hist. Soc., Bul. 5: 99. 9°.

Genus ANISEPYRIS Kieffer

Anisepyris Kieffer, 1905. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 248.
Type-species: Epyris amazonicus Westwood. Monotypic.
Rhabdepyris subg. Lophepyris Evans, 1959. Ent. Soc. Wash., Proce. 61: 201.
Type-species: Rhabdepyris (Lophepyris) bridwelli Evans. Orig. desig.
Revision: Evans, 1959. Ent. Soc. Wash., Proc. 61: 97-120, 28 figs. (U. S. species). —Evans,

1959. Ent. Soe. Wash., Proc. 61: 201-204 (Lophepyris). —Evans, 1966. Studia Ent. 9: 1-120,
151 figs. (New World species).

SPECIES GRouUP AENEUS
The Aeneus Species Group is equivalent to subgenus Lophepyris Evans.

aeneus Kieffer. Southern Tex. to Panama.

Anisepyris aeneus Kieffer, 1906. Soc. Sci. Bruxelles, Ann. 30: 138. 2.

Anisepyris sublevis Kieffer, 1906. Soc. Sci. Bruxelles, Ann. 30: 140. ¢.

Rhabdepyris (Lophepyris) bridwelli Evans, 1959. Ent. Soc. Wash., Proc. 61: 202. 2, ¢.
bradleyi (Evans). Fla., Tex.

Rhabdepyris (Lophepyris) bradleyi Evans, 1959. Ent. Soe. Wash., Proce. 61: 204. 2.

1208 Hymenoptera in America North of Mexico

SPECIES GROUP AURICHALCEUS

aurichalceus (Westwood). Fla. (Miami); Cuba, Puerto Rico, St. Thomas, St. Kitts.
Epyris aurichalceus Westwood, 1874. Thesaurus Ent. Oxon., p. 160, pl. 31, fig. 3. 9.
Anisepyris viridis Kieffer, 1907. Soc. Sci. Bruxelles, Ann. 32: 12. ¢. Preoce.
Anisepyris viridellus Kieffer, 1914. Das Tierreich, Lief. 41, p. 438. N. name.
Anisepyris cubensis Fouts, 1928. Ent. Soe. Wash., Proc. 30: 125. °.

SPECIES GROUP PROTEUS

arapaho Evans. Tex. (Culberson Co.).
Anisepyris arapaho Evans, 1966. Studia Ent. 9: 62, figs. 44, 65. 2.
dietrichorum Evans. Southern Ariz.; Mexico (Sinaloa, Nayarit, Jalisco, Veracruz).
Anisepyris dietrichorum Evans, 1959. Ent Soc. Wash., Proc. 61: 118, fig. 21. 2.
papago Evans. Ariz. (Pima Co.).
Anisepyris papago Evans, 1966. Studia Ent. 9: 63, fig. 45. @.
rugosicollis Brues. Tex. (Brownsville).
Anisepyris rugosicollis Brues, 1908. Wis. Nat. Hist. Soc., Bul. (2) 6: 48. ?.

SPECIES GROUP OCCIDENTALIS

arizonicus Evans. Ariz. (Pima Co., Cochise Co.).
Anisepyris arizonicus Evans, 1959. Ent. Soc. Wash., Proc. 61: 112, fig. 15. °.
gibbosifrons Evans. N. J., Fla.
Anisepyris gibbosifrons Evans, 1959. Ent. Soc. Wash., Proc. 61: 116, figs. 23, 25, 26. 2, d.
laticeps Evans. Southeastern Ariz.
Anisepyris laticeps Evans, 1959. Ent. Soc. Wash., Proc. 61: 113, figs. 18, 19, 24. 2, 6.
Anisepyris latifrons (!) Evans, 1959. Ent. Soc. Wash., Proc. 61: 115, fig. 24 legend. Lapsus.
occidentalis (Ashmead). Wash., Oreg., Idaho, Utah, Nev., Calif., Ariz.; Mexico (Aguascalientes,
Guadalajara, Guerrero, Chiapas).
Epyris occidentalis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 58, 59. 6, @.
Anisepyris punctaticeps Kieffer, 1906. Soc. Sci. Bruxelles, Ann. 30: 139. 6.
subviolaceus Kieffer. Mass., Mich., Alta. and B. C. south to Fla., Tex. and Ariz.; Mexico
(Sinaloa, Nayarit, Morelos, Veracruz).
Anisepyris subviolaceus Kieffer, 1910. Soc. Ent. France, Ann. 79: 39. 2.

SPECIES GROUP COLUMBIANUS

columbianus (Ashmead). Idaho and Wash. south through Ariz. and Calif. to Costa Rica.
Epyris columbianus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 58, 60. 6, 9.
Anisepyris pulchellus Fouts, 1928. Ent. Soc. Wash., Proc. 30: 126. 2.

grandis (Ashmead). Fla.
Goniozus grandis Ashmead, 1887. Ent. Amer. 3: 76. 6, 2.
Epyris aeneiceps Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 59. 9..

williamsi Evans. Utah, Idaho and Oreg., south through Ariz. and Calif. to southern Mexico.
Anisepyris williamsi Evans, 1959. Ent. Soc. Wash., Proc. 61: 106, fig. 9. 2, 6.

SPECIES GROUP AMAZONICUS

analis (Cresson). N. C. to Fla., Tenn., Tex.; Mexico (Hidalgo).
Epyris analis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 193. 9.

Genus EPYRIS Westwood

Epyris Westwood, 1832. London, Edinb. and Dublin Phil. Mag. and Jour. (ser. 3) 1 (2): 129.
Type-species: E pyris niger Westwood. Monotypic.

Muellerella Saussure, 1892. In Grandidier, Hist. Nat. Madagascar 20: pl. 25, fig. 20.
Type-species: Muellerella amabilis Saussure. Monotypic.

Empyris (!) Melander and Brues, 1903. Biol. Bul. 5: 23.

Parepyris Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 108.
Type-species: Epyris interruptus Kieffer. Desig. by Kieffer, 1914.

Psilepyris Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 108.
Type-species: E pyris indivisus Kieffer. Desig. by Kieffer, 1914.

Superfamily BETHYLOIDEA 1209

Several species are known to parasitize soil-dwelling beetle larvae, especially Tenebrionidae.
A few species hibernate gregariously beneath or in bark of trees. Several species of which E.
californicus (Ashm.) is an example, have a very painful sting and can cause severe allergic reac-
tions in humans.

Revision: Evans, 1969. Amer. Ent. Soe., Trans. 95: 181-352, 168 figs., 24 maps (New World

species).
SPECIES GROUP CONNEXUS
connexus Evans. Southern Ariz. and Calif.
Epyris connexus Evans, 1967. Ent. News 78: 97. 2, 6.
spissus Evans. Ont., Mass., south to Fla.
Epyris spissus Evans, 1969. Amer. Ent. Soc., Trans. 95: 193, figs. 1, 3, 7, map 1. 9, 6.
SPECIES GRouP TRICOSTATUS
cariniceps Evans. Central Calif., southern Ariz., western Tex. to Costa Rica.
Epyris cariniceps Evans, 1969. Amer. Ent. Soc., Trans. 95: 216, figs. 17, 41, 49, map 5. @,
3.

eriogoni Kieffer. Calif., Ariz., N. Mex., Tex. south to Veracruz.
Epyris eriogoni Kieffer, 1906. Berlin. Ent. Ztschr. 50: 245. 6.
Epyris erigoni (!) Brues, 1907. Wis. Nat. Hist. Soc., Bul. 5: 97.

Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 220, figs. 22, 25, 40, 50, map 6. 2, 6.
oriplanus Kieffer. Va. to Fla., Tex. to Calif., south to Chiapas.
Epyris oriplanus Kieffer, 1911. Soc. Sci. Bruxelles, Ann. 35: 228. °.
Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 213, figs. 10, 11, 19, 33, 34, map 4. 2, d.
tricostatus Evans. Pa. to Fla., Kans., Tex.
Epyris tricostatus Evans, 1969. Amer. Ent. Soc., Trans. 95: 208, figs. 16, 39, 47, map 3. 2,
é.
vogti Evans. Md. (Nanjemoy).
Epyris vogti Evans, 1969. Amer. Ent. Soc., Trans. 95: 211, fig. 24, map 5. °.

SPECIES GROUP RUFIPES

brachypterus (Ashmead). Md., D. C., Va., “Carolina”.
Mesitius brachypterus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 66. °.
Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 273, map 11. °.

californicus (Ashmead). Pa. to Fla., IIl., Kans., Colo., Tex., Oreg., Calif. and Ariz. south to
Honduras.
Mesitius californicus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 65. 9.
Mesitius minutus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 65. 3.

Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 260, figs. 63, 64, 72, 98-101, map 10. °,
Ge
Biology: Essig and Michelbacher, 1932. Science 76: 407-408.

clarimontis Kieffer. Fla., N. Dak. to Wash. south to Calif. and Tex., thence to central Mexico.
Epyris clarimontis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 243. °.
Epyris indivisus Kieffer, 1905. Berlin. Ent. Ztschr. 50: 243. d.

Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 280, figs. 73, 74, 75, 86, 108-111, map 14.
Oni
cochise Evans. Idaho, Utah, Nev., Ariz., N. Mex., Tex.
Epyris cochise Evans, 1969. Amer. Ent. Soc., Trans. 95: 285, figs. 84, 85, 114, map 15. d, °.

deficiens Krombein. N. Y. to Ga., W. Va., ?Kans.
Epyris deficiens Krombein, 1956. Ent. Soc., Wash., Proce. 58: 156, fig. 3. °.
Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 288, figs. 68, 69, 78, 115, map 15. @, ¢.

guatemalensis Cameron. Southern Calif. through western and central Mexico to Guatemala.
Epyris guatemalensis Cameron, 1888. Biol. Cent.-Amer., Hym. 1: 453, pl. 19, fig. 20. °.

1210 Hymenoptera in America North of Mexico

Epyris guatamalensis (!) Evans, 1969. Amer. Ent. Soe., Trans. 95: 264, fig. 104, map 17. ?,
3.
monticola (Ashmead). Colo.; Mexico (Durango).

Epyris monticola Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 8. d.

Taxonomy: Evans, 1969. Amer. Ent. Soe., Trans. 95: 256, figs. 58, 59, 96, map 16. 6.

myrmecophilus (Brues). Vt., Wis. and Colo. south to Fla., Tex., Ariz., southern Calif.
Mesitius myrmecophilus Brues, 1903. Amer. Ent. Soc., Trans. 29: 124. ?.

Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 269, figs. 65, 105, 107, map 12. ?, 6.
rufipes (Say). Que., B. C., U.S. south at higher elevations to Guatemala.

Bethylus rufipes Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 329. @ (?).

Bethylus formicoides Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 179. 2.

Epyris rufipes Cameron, 1888. Manchester Lit. and Phil. Soc., Mem. and Proe. 1: 173. °.
Preoce.

Epyris montezuma Cameron, 1888. Manchester Lit. and Phil. Soc., Mem. and Proc. Errata
1: vii. N. name.

Mesitius vancouverensis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 64. 2.

Mesitius nevadensis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 64. °.

Mesitius bifoveolatus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 66. 3, °.

Epyris longicollis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 244. 2. Preoce.

Epyris nudicornis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 245. 6.

Epyris gracilicollis Kieffer, 1908. In Wytsman, Gen. Ins., fase. 76, p. 28. N. name.

Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 250, figs. 53, 54, 91-94, 97, map 9. 2, d.

sculleni Evans. Alta., Wash., Idaho, Utah, Oreg., Calif., Ariz., N. Mex.; Mexico (Hidalgo).
Epyris sculleni Evans, 1969. Amer. Ent. soc., Trans. 95: 266, figs. 66, 67, 82, 102, 103, map
Mok, 65 Ge
sepulchralis Evans. Mass., Md., Fla., Tenn., Okla., Tex., southern Calif.; Mexico (Guanajuato,
Jalisco, Veracruz).
Epyris sepulchralis Evans, 1969. Amer. Ent. Soc., Trans. 95: 275, figs. 76, 79, 80, 112, map
1Sacn ee
texanus (Ashmead). N. Y., Va., N. C., Ill, Tex.
Mesitius texranus Ashmead, 18938. U.S. Natl. Mus., Bul. 45: 67. °.
Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 290, map 16. °.
vierecki Krombein. Mass., N. Y., Md., Va., W. Va., N. C., S. C., Ga.
Epyris vierecki Krombein, 1962. Biol. Soc. Wash., Proc. 75: 1. d, 2.
Taxonomy: Evans, 1969. Amer. Ent. Soc., Trans. 95: 277, figs. 77, 81, 106, map 14. 2, d.
SPECIES GROUP DEPRESSIGASTER
alachua Evans. Fla. (Alachua Co.).
Epyris alachua Evans, 1969. Amer. Ent. Soc., Trans. 95: 337, fig. 1538, map 22. 2.

corticinus Evans. Pa., Md., Va.
Epyris corticinus Evans, 1969. Amer. Ent. Soc., Trans. 95: 338, figs. 148, 149, map 22. 2.

festivus Evans. Fla. (Lake Co., Dixie Co.).
Epyris festivus Evans, 1969. Amer. Ent. Soc., Trans. 95: 339, map 22. 2.

SPECIES GROUP IDIONOTUM

idionotum Evans. Fla.
Epyris idionotum Evans, 1969. Amer. Ent. Soc., Trans. 95: 340, figs. 146, 156, map 23. d.

NOMEN NUDUM IN EPYRIS WESTWOOD

Epyris monticola Ashmead in Cockerell, 1889. Colo. Biol. Assn., 10th Rept., [p. 2] (probably
published originally in Custer County Courant newspaper).

Genus BAKERIELLA Kieffer

Bakeriella Kieffer, 1910. Soe. Ent. France, Ann. 78: 288.
Type-species: Bakeriella flavicornis Kieffer. Monotypic.

Superfamily BETHYLOIDEA 1211

floridana Evans. Fla. (Paradise Key).
Bakeriella floridana Evans, 1964. Mus. Compar. Zool., Bul. 182: 134, fig. 84. 6, ?.

Genus HOLEPYRIS Kieffer

Holepyris Kieffer, 1904. Mus. Civ. Stor. Nat. Genova, Ann. 41: 390.
Type-species: Holepyris africanus Kieffer. Desig. by Kieffer, 1906.
Holepyris subg. Rysepyris Kieffer, 1906. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 341.
Type-species: Holepyris (Rysepyris) numidicus Kieffer. Desig. by Kieffer, 1914.
Misepyris Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 108.
Type-species: Holepyris remotus Kieffer. Desig. by Kieffer, 1914.
Parepyris Brethes, 1913. Buenos Aires Mus. Nac. de Hist. Nat., An. 24: 87.
Type-species: Parepyris sylvanidis Brethes. Monotypic.

Authenticated host records indicate that members of this genus parasitize coleopterous lar-
vae, although in the laboratory they can be induced to attack Microlepidoptera also. One North
American species attacks larvae infesting grain, and another was reared from a beetle larva
beneath bark.

coronatus (Ashmead). Md. (Bladensburg). Host: Catogenus rufus (F.).
Apenesia coronata Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 47. 6.
floridanus (Ashmead). Fla.
Isobrachium floridanum Ashmead, 1887. Ent. Amer. 3: 76. d.
haemorrhoidalis (Kieffer). Tex.
Rhabdepyris haemorrhoidalis Kieffer, 1904. Arkiv. for Zool. 1: 528. 2.
marylandicus Fouts. Md., N. C.
Holepyris marylandicus Fouts, 1928. Ent. Soc. Wash., Proc. 30: 125. 2.
punctifrons Fouts. Fla. (Hawthorne).
Holepyris punctifrons Fouts, 1927. Ent. Soc. Wash., Proc. 29: 166. °.
subapterus (Melander and Brues). Mass.
Empyris (!) subapterus Melander and Brues, 1903. Biol. Bul. 5: 23. 2.
sylvanidis (Brethes). Widespread in U. S.; cosmopolitan. Host: Oryzaephilus surinamensis
(L.); Sitophilus oryza (L.); Tribolium castaneum (Hbst.); T. confusum Duv.;
Laemophloeus ferrugineus (Steph.).
Parepyris sylvanidis Brethes, 1913. Buenos Aires Mus. Nac. de Hist. Nat., An. 24: 87. d,
2

Rhabdepyris zeae Turner and Waterson, 1921. Rpt. Grain Pests (War) Com. [Gt. Brit.], No.
hip Zone

Genus LAELIUS Ashmead

Laelius Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 50.
Type-species: Laelius trogodermatis Ashmead. Orig. desig.
Paralaelius Kieffer, 1905. Soe. Sci. Bruxelles, Ann. 29: 129.
Type-species: Bethylus pedatus Say. Desig. by Kieffer, 1914.

All authenticated host records for North American Laelius are larvae of Dermestidae, usually
in and around buildings.

Revision: Muesebeck, 1939. Biol. Soc. Wash., Proc. 52: 171-176.

centratus (Say). Fla., Ind., Colo.
Bethylus centratus Say, 1836. Boston Jour. Nat. Hist. 1: 281.
occidentalis Whittaker. B. C. (Chilliwack).
Laelius occidentalis Whittaker, 1928. Ent. Soc. London, Trans. p. 387. °.
pedatus (Say). N. J., N. Y., D. C., Va., S. C., Fla., Ind., Colo. south to Mexico (Morelos). Host:
Anthrenus verbasci (L.).
Bethylus pedatus Say, 1836. Boston Jour. Nat. Hist. 1: 280.
Ateleopterus nubilipennis Ashmead, 1887. Ent. Amer. 3: 97. 2.
Laelius tricarinatus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 51. d, °.
Laelius rufipes Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 51. 2. Preoce.

1212 Hymenoptera in America North of Mexico

Laelius nigripilosus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 52. d, °.
Bethylus constrictus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 53. 2.
Laelius ashmeadi Kieffer, 1908. In Wytsman, Gen. Ins., fase. 76, p. 38. N. name.
Laelius fumipennis Brues, 1910. Wis. Nat. Hist. Soc., Bul. (Ser. 2) 8: 45. 2.
trogodermatis Ashmead. Que., N. Y., D. C., Va., Fla. Host: Trogoderma parabile Beal, T.
simplex Jayne, T. versicolor Creutz.
Laelius trogodermatis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 51. d, 2.

Biology: Howard, 1901. The Insect Book, pp. 34-36, fig. 18. —Krombein, 1955. Brooklyn Ent.
Soe., Bul. 50: 13-14.
utilis Cockerell. Ont., Mass., Md., D. C., Va., N. C. Host: Trogoderma versicolor Creutz.;
Bruchus brachialis Fahr.?
Laelius utilis Cockerell, 1920. Canad. Ent. 52: 34. °.
voracis Muesebeck. D. C., Va.; India. Host: Anthrenus vorax (Waterh.).
Laelius voracis Muesebeck, 1939. Biol. Soe. Wash., Proc. 52: 172. 2, d.

Biology: Back, 1940. Ent. Soc. Wash., Proc. 42: 110-113, 1 pl. —Kurian, 1954. Agra Univ.,
Jour. Res. (Sci.) 3: 422, figs. 9-13.

TRIBE CEPHALONOMIINI

Genus PLASTANOXUS Kieffer

Plastanoxus Kieffer, 1905. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 244.
Type-species: Anoxus chittendenti Ashmead. Desig. by Gahan, 1931.

These tiny wasps are parasitic on coleopterous larvae.
Revision: Gahan, 1931. Wash. Acad. Sci., Jour. 21: 213-218, figs. 1-3.

Taxonomy: Evans, 1964. Mus. Compar. Zool., Bul. 132: 151-152 (key to species).
chittendenii (Ashmead). Transcontinental in southern Canada and U. S.; adventive in England.
Host: Cis fuscipes Mellie?
Anoxus Chittendenii Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 68. 3.
incompletus Evans. Ariz. (Mesa).
Plastanoxus incompletus Evans, 1964. Mus. Compar. Zool., Bul. 132: 152. 2.
laevis (Ashmead). D. C.; ?Brazil.
Anoxus laevis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 68. °.
westwoodi (Kieffer). D. C., Ga.; probably cosmopolitan. Host: Laemophloeus pusillus (Schonh.).
Cephalonomia westwoodi Kieffer, 1914. Das Tierreich, Lief. 41, p. 248. d, 9.
Plastanoxus kiefferi Gahan, 1931. Wash. Acad. Sci., Jour. 21: 217. 2, d (? Lapsus for
westwoodi.)

‘Genus CEPHALONOMIA Westwood

Cephalonomia Westwood, 1853. Mag. Nat. Hist. 6: 420.
Type-species: Cephalonomia formicifornis Westwood. Monotypic.

Holopedina Foerster, 1850. Naturh. Ver. Rheinlande Verh. 7: 502.
Type-species: Holopedina polypori Foerster. Monotypic.

Cephalonomia subg. Cephaloderma Hoffer, 1936. Festchr. Embrik Strand, v. 1, p. 459.
Type-species: Cephalonomia (Cephaloderma) strandi Hoffer. Monotypic.

So far as known these tiny wasps parasitize the larvae or pupae of small Coleoptera occurring
in cryptic situations. One group attacks pests of stored grains, another parasitizes Ciidae occur-
ring in fungi, and the third attacks bark beetles. Four types of species may be distinguished
based on development of the wings in both sexes: One in which both sexes are macropterous;
one in which the males are macropterous, the females macropterous or brachypterous; a third in
which the males are macropterous or apterous and the females apterous; and the last in which
the males are macropterous or apterous, the females macropterous, micropterous or apterous.

Biology: Evans, 1964. Mus. Compar. Zool., Bul. 1382: 158-159.

Superfamily BETHYLOIDEA 1213

Morphology: Grandi, 1929. Bol. Lab. Ent. Bologna 2: 301-314. —van Emden, 1931. Ztschr. f.
Morph. u. Oekol. der Tiere 23: 425-574. —Grandi, 1932. Bol. Lab. Ent. Bologna 5: 13-21.
cynipsiphila (Ashmead). Fla.
Sclerochroa cynipsiphila Ashmead, 1887. Ent. Amer. 3: 75. °.
gallicola (Ashmead). Widely distributed in North America; probably cosmopolitan. Host:
Stegobium paniceum (L.); Lasioderma serricorne (F.); Ptinus sp.; Araecerus
fasciculatus de Geer.
Sclerochroa gallicola Ashmead, 1887. Ent. Amer. 3: 75. 2.
Holopedina nubilipennis Ashmead, 1887. Ent. Amer. 3: 97. 2.
2Cephalonomia xambeui Giard, 1898. Soc. Ent. France, Bul., p. 50. 2, 3.
Cephalonomia quadridentata Duchaussoy, 1917. Soc. Hist. Nat. Afrique du Nord, Bul. 9:
ili, ©.
Cephalonamia caesarorum van Emden, 1931. Ztschr. Morph. u. Oekol. Tiere 23: 431.
Cephalonomia (Cephaloderma) strandi Hoffer, 1936. Festschr. Embrik Strand, v. 1, p. 460.
OF

Biology: van Emden, 1931. Ztschr. Morph. u. Oekol. Tiere 23: 425-574. — Kearns, 1934. Jour.
Econ. Ent. 27: 801-806. —Kearns, 1934. Ent. Soc. Amer., Ann. 27: 533-541. —Cabal Concha,
1956. Rev. Facult. Nac. Agron. [Colombia] 18: 64.

Morphology: van Emden, 1931. Ztschr. Morph. u. Oekol. Tiere 23: 425-574. —Grandi, 1932.
Lab. Ent. R. Ist. Sup. Agr. Bologna, Bol. 5: 113-115. —Reid, 1941. Roy. Ent. Soc. London,
Trans. 91: 403-404.

hyalinipennis Ashmead. Mass., Conn., Fla., Calif. Host: Conophthorus coniperda (Schw.);
Hypothenemus spp.; Pissodes terminalis Hopping; Pityophthorus sp.; Scolytus
rugulosus (Ratz.). Also recorded from galls of Amphibolips cinerea (Ashm.) and
Disholcaspis omnivora (Ashm.), where, Dee gunably, it parasitizes bark beetles invading
the woody tissues of the galls.
Cephalonomia hyalinipennis Ashmead, 1898. U.S. Natl. Mus., Bul. 45: 49. ¢, @.

Biology: Stark and Wood, 1964. Canad. Ent. 96: 1217. —Godwin and Odell, 1965. Ent. Soe.
Amer., Ann. 58: 218.
perpusilla Evans. Oreg., Calif., Ariz.; Mexico (Baja California, Nayarit). Host: Ciid beetle
larvae in fungi.
Cephalonomia perpusilla Evans, 1963. Psyche 70: 152. d, 9.
tarsalis (Ashmead). Widely distributed in U. S.; probably cosmopolitan. Host: Oryzaephilus
surinamensis (L.); Sitophilus oryzae (L.).
Ateleopterus tarsalis Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 45. 9, ¢.
Cephalonomia carinata Kieffer, 1907. Berlin. Ent. Ztschr. 51: 295. 2.
Cephalonomia kiefferi Fouts, 1920. Ent. Soc. Wash., Proc. 22: 77. 2, d. -

Biology: Richards and Herford, 1930. Ann. Appl. Biol. 17: 382. —Powell, 1938. Ent. Soc.
Amer., Ann. 31: 44-49.

utahensis Brues. Utah, Calif. Host: Conophthorus lambertianae Hopk.; C. monophyllae Hopk.;
C. ponderosae Hopk.; C. radiatae Hopk.
Cephalonomia utahensis Brues, 1909. Wis. Nat. Hist. Soc., Bul. 6: 154. °.

Biology: Ruckes, 1956. Pan-Pacifie Ent. 32: 184-185, 1 fig. —Schaefer, 1962. Ent. Soc. Amer.,
Ann. 55: 574-575, fig. 6. —Ruckes, 1963. Pan-Pacifie Ent. 39: 45.

waterstoni Gahan. Widely distributed in North America; probably cosmopolitan. Host:
Laemophloeus ferrugineus (Steph.).
Cephalonomia waterstoni Gahan, 1931. Wash. Acad. Sci., Jour. 21: 219. 9.

Biology: Sheppard, 1936. Colo. Expt. Sta., Tech. Bul. 17: 16-17. —Rilett, 1949,. Canad. Jour.
Res., Sect. D, Zool. Sci. 27: 1-27. —Finlayson, 1950. Bul. Ent. Res. 41: 79.

TRIBE SCLERODERMINI

Genus NESEPYRIS Bridwell
Nesepyris Bridwell, 1920. Hawaiian Ent. Soc., Proc. 4: 309.

1214 Hymenoptera in America North of Mexico

Type-species: Nesepyris ewa Bridwel!. Monotypic.

The small rare species belonging to this genus are probably parasitic on borers in twigs or
bark.
floridanus Evans. Fla.; Mexico.
Nesepyris floridanus Evans, 1964. Mus. Compar. Zool., Bul. 132: 165, 111, 112, °.
virginianus Evans. Md., Va. Bred from redbud (Cercis canadensis) and found under hickory

bark.
Nesepyris virginianus Evans, 1964. Mus. Compar. Zool., Bul. 132: 164, fig. 110. 2.

Genus GLENOSEMA Kieffer

Glenosema Kieffer, 1905. Soc. Sci. Bruxelles, Ann. 29: 100. No species.
Type-species: Glenosema nigra Kieffer. First included species.
Arysepyris Kieffer, 1905. Soc. Sci. Bruxelles, Ann. 29: 102.
Type-species: Arysepyris merceti Kieffer. Orig. desig.
crandalli Evans. Ariz. (Santa Catalina Mts.).
Glenosema crandalli Evans, 1964. Mus. Compar. Zool., Bul. 132: 170, fig. 116. °.

Genus SCLERODERMUS Latreille

Sclerodermus Latreille, 1809. Gen. Crust. Ins., v. 4, p. 118.

Type-species: Sclerodermus domesticus Latreille. Monotypic.
Scleroderma(!) Oken, 1817. Isis 8: 1178. Emend.
Schlerochroa Foerster, 1850. Naturh. Ver. Rheinlande, Verh. 7: 501. Unnecessarily

proposed as a new name for Sclerodermus Latr.

Schleroderma (!) Costa, 1862. Annuario Mus. Zool. Napoli 2: 131. Misspelling.
Neoscleroderma Kieffer, 1905. Soc. Sci. Bruxelles, Ann. 29: 106.

Type-species: Ateleopterus virginiensis Ashmead. Desig. by Kieffer, 1914.

The North American species are parasitic on wood-boring beetle larvae, frequently those in-
festing houses. The adult wasps have frequently been reported as stinging humans in infested
houses.

carolinensis (Ashmead). Conn. and N. Y. to Ga. Host: Dicerca lepida Lac.; Urographis fasciata
(de Geer).
Dissomphalus carolinensis Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 42, 43. 9.
Ateleopterus virginiensis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 45. 3, 9.
macrogaster (Ashmead). D. C., Va., N. C., Fla., Miss., Tex.; Cuba, Hispaniola, Jamaica, Brazil.
Host: Megacyllene antennatus (White), Scolytus rugulosus (Ratz.), Xyletinus peltatus

(Harr.).
Sclerochroa macrogaster Ashmead, 1887. Ent Amer. 3: 75. 2.

SUBFAMILY PRISTOCERINAE
Genus PRISTOCERA Klug
Genus PRISTOCERA Subgenus PRISTOCERA Klug

Pristocera Klug, 1808. Mag. Gesell. Naturf. Freunde Berlin 2: 49.
Type-species: Bethylus depressus Fabricius. Monotypic.

Mangesia Kieffer, 1911. Soc. Sci. Bruxelles, Ann. 35: 209.
Type-species: Mangesia fuscipennis Kieffer. Orig. desig.

Trichelobrachium Kieffer, 1914. Das Tierreich, Lief. 41, p. 425.
Type-species: Pristocera obliterata Kieffer. Monotypic.

The typical subgenus does not occur in North America.
Genus PRISTOCERA Subgenus ACREPYRIS Kieffer

Acrepyris Kieffer, 1905. Jn Andre, Spec. Hym. Eur. Alg., v. 9, p. 249.
Type-species: E pyris reticulatus Kieffer. Orig. desig.

Superfamily BETHYLOIDEA 1215

Pristocera subg. Neopristocera Yasumatsu, 1955. Fac. Agr. Kyushu Univ., Jour. 10: 248.
Type-species: Pristocera japonica Yasumatsu. Orig. desig.

These wasps are parasitic on the larvae of Elateridae (wireworms) in the soil. Males carry the
females in flight during mating.

Revision: Evans, 1963. Mus. Compar. Zool., Bul. 129: 241-290, 60 figs.
armifera (Say). N. H. and Vt. south to Fla., west to S. Dak., Colo. and Tex. Host: Aeolus sp.;
Limonius agonus (Say); Melanotus sp.
Bethylus armiferus Say, 1828. Contrib. Maclur. Lye. Phila. 1: 80. d.
Scleroderma thoracica Westwood, 1839. Ent. Soe. London, Trans. 2: 167. °.
Scleroderma contracta Westwood, 1839. Ent. Soc. London, Trans. 2: 169, pl. 12, fig. 11. 2.
Epyris laeviventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 193. d.
Epyris reticulatus Kieffer, 1904. Arkiv for. Zool. 1: 527. ¢.

Biology: Hyslop, 1916. Ent. Soc. Wash., Proc. 18: 170, pl. 11, figs. 1-3.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14: 44-46, pl. 15, figs. A-D
(male genitalia).
atra Klug. N. C. to Fla., west to N. Mex.
Pristocera atra Klug, 1810. Beitr. Naturk., v. 2, p. 206. ¢.

Taxonomy: Evans, 1963. Mus. Compar. Zool., Bul. 129: 273-276, figs. 5, 12, 17, 48, 53, 54. d, @.

bridwelli Evans. Ark. (Dodd City).

Pristocera (Acrepyris) bridwelli Evans, 1963. Mus. Compar. Zool., Bul. 129: 267. 6, °.
californica Evans. Calif., Utah, Wyo.

Pristocera (Acrepyris) californica Evans, 1963. Mus. Compar. Zool., Bul. 129: 263. d.
chihuahua Evans. Ariz.; Mexico (Chihuahua, Zacatecas).

Pristocera (Acrepyris) chihuahua Evans, 1963. Mus. Compar. Zool., Bul. 129: 284. 6.
cockerelli Evans. Tex. to Calif.; Mexico (Sonora, Oaxaca).

Pristocera (Acrepyris) cockerelli Evans, 1963. Mus. Compar. Zool., Bul. 129: 264. 3, °.
fraterna Evans. N. C. to Fla., Kans.

Pristocera (Acrepyris) fraterna Evans, 1963. Mus. Compar. Zool., Bul. 129: 261. d, 2 (?).
hyalina Brues. Ark., La., Tex., N. Mex., south to central Mexico.

Pristocera hyalina Brues, 1906. Wis. Nat. Hist. Soc., Bul. 4: 148. ¢.

Genus APENESIA Westwood

Apenesia Westwood, 1874. Thesaurus Ent. Oxon., p. 170.

Type-species: A penesia amazonica Westwood. Desig. by Westwood, 1881.
Aeluroides Tullgren, 1904. Ark. Zool. 1: 428.

Type-species: Aeluroides sjostedti Tullgren. Monotypic.
Propristocera Kieffer, 1905. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 247.

Type-species: Propristocera interrupta Kieffer. Desig. by Kieffer, 1914.
Cleistepyris Kieffer, 1910. Soc. Ent. France, Ann. 79: 48.

Type-species: Cleistepyris punctatus Kieffer. Desig. by Kieffer, 1914.
Dipristocera Kieffer, 1914. Das Tierreich, Lief. 41, p. 471.

Type-species: Dipristocera microchela Kieffer. Monotypic.
Neopristocera Benoit, 1957. Explor. Pare Nat. Albert, Mission DeWitte, fasc. 88, p. 44.

Preoce.
Type-species: Neopristocera triloba Benoit. Orig. desig.

Hosts are unknown for the North American species, but two Old World species have been
reared from weevil larvae (Curculionidae) in roots or stems.

Revision: Evans, 1963. Mus. Compar. Zool., Bul. 130: 249-359, 138 figs.
SPECIES GROUP EXILIS

cochise Evans. Ariz. (Cochise Co.).
Apensia cochise Evans, 1963. Mus. Compar. Zool., Bul. 130: 296, figs. 27, 32. ¢.

1216 Hymenoptera in America North of Mexico

exilis Evans. Ariz., Calif.
Apenesia exilis Evans, 19638. Mus. Compar. Zool., Bul. 130: 293, figs. 28, 30, 33, 34. 3.

martini Evans. Fla. (Manatee Co.).
Apenesia martini Evans, 1963. Mus. Compar. Zool., Bul. 130: 294, fig. 29. ¢.

pima Evans. Ariz. (Cochise Co.).
Apenesia pima Evans, 1963. Mus. Compar. Zool., Bul. 130: 292, figs. 26, 31, 35, 36. d.

SPECIES GROUP DISSOMPHALOIDES

dissomphaloides Evans. Ariz.; Mexico (Sinaloa).
Apenesia dissomphaloides Evans, 1963. Mus. Compar. Zool., Bul. 130: 297, figs. 37, 42. 3.

SPECIES GROUP LAEVIGATA

pallidula Evans. Ariz. (Cochise Co.).
Apenesia pallidula Evans, 1963. Mus. Compar. Zool., Bul. 130: 300, figs. 38, 43, 46, 50. 3.

SPECIES GROUP MEXICANA

chiricahua Evans. Tex., Ariz.; Mexico (Sinaloa, Durango, Mexico).
Apenesia chiricahua Evans, 1963. Mus. Compar. Zool., Bul. 130: 306, figs. 57, 60, 69, 80, 84.
3.
malinche Evans. Ariz. (Tucson Mts.); Mexico (Sinaloa, Puebla).
Apenesia malinche Evans, 1963. Mus. Compar. Zool., Bul. 130: 309, figs. 59, 62. ¢.
mohave Evans. Calif.; Mexico (Baja California).
Apenesia mohave Evans, 1963. Mus. Compar. Zool., Bul. 130: 308, figs. 58, 61. d.

SPECIES GROUP BRASILIENSIS

parapolita Evans. Ark., La., Ala., and Ga., north to IIl., W. Va. and N. Y.
Propristocera polita Evans, 1958. Ent. Soc. Wash., Proc. 59: 294, figs. 2, 6. d. Preoce.
Apenesia parapolita Evans, 1963. Mus. Compar. Zool., Bul. 130: 322 (male), 347 (female),
figs. 81, 96, 127, 1386. N. name.

Genus PARASCLERODERMA Kieffer

Parascleroderma Kieffer, 1904. Mus. Civ. Stor. Nat. Genova, Ann. 41: 376.
Type-species: Parascleroderma fulviceps Kieffer. Orig. desig.

No host information is available for the single North American species. Two Old World spe-
cies have been recorded as attacking the larvae of bostrichid and clerid beetles. The normal
hosts apparently are larvae occurring under bark or in borings in wood.

insolita (Evans). Tex. (Brownsville).
Apenesia insolita Evans, 1963. Mus. Compar. Zool., Bul. 130: 350, figs. 130, 138. 9.

Genus DISSOMPHALUS Ashmead

Dissomphalus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 41.
Type-species: Dissomphalus xanthopus Ashmead. Orig. desig.
Ecitopria Wasmann, 1899. Zoologica, h. 26, p. 55.
Type-species: Ecitopria crassicornis Wasmann. Monotypic.
Dissemphalus (!) Ashmead, 1902. Canad. Ent. 34: 271.
? Psilobethylus Kieffer, 1906. In Andre, Spec Hym. Eur. Alg., v. 9, p. 461.
Type-species: Psilobethylus luteus Kieffer. Monotypic.
Thaumatepyris Kieffer, 1910. Soc. Ent. France, Ann. 79: 47.
Type-species: Thaumatepyris punctatus Kieffer. Monotypic.
Glenobethylus Kieffer, 1910. Soc. Ent. France, Ann. 79: 50.
Type-species: Glenobethylus montanus Kieffer. Monotypic.
Parecitopria Ogloblin, 1930. Soc. Ent. Arg., Rev. 3: 15.
Type-species: Parecitopria azarai Ogloblin. Monotypic.

Associated data with several species suggest that these minute wasps are probably parasitic
on larvae of both myrmecophilous and wood-boring beetles. The wingless females are apparently
carried by the males during mating flights.

Superfamily BETHYLOIDEA 1217

Revision: Evans, 1954. Ent. Soe. Wash., Proc. 56: 288-309, 21 figs. (North American species).

Taxonomy: Evans, 1962. Ent. Soe. Wash., Proc. 64: 65-78, 4 figs. (revised key, North
American and West Indian species).

SPECIES GROUP KANSANUS

arizonicus Evans. Ariz. (Portal).
Dissomphalus arizonicus Evans, 1962. Ent. Soc. Wash., Proc. 64: 70, fig. 3. 6.

kansanus Evans. Pa., Fla., Ill., Kans.
Dissomphalus kansanus Evans, 1954. Ent. Soe. wash., Proc. 56: 302, figs. 4, 11, 17. d.

SPECIES GROUP XANTHOPUS

barberi Evans. Maine, Md., N. C.
Dissomphalus barberi Evans, 1954. Ent. Soe. Wash., Proc. 56: 298, figs. 3, 10, 16, 19. 3, 2.

californicus Ashmead. Ariz., Calif.
Dissomphalus californicus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 42. 6.

Taxonomy: Evans, 1967. Ent. News 78: 17. ¢.
xanthopus Ashmead. U. and L. Austr. Zones east of Rockies, south in Mexico to Chiapas.
Dissomphalus xanthopus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 42. 2, 3.
Psilobethylus lucidus Brown and Cheng, 1952. Psyche 58: 146, figs. 2, 3. 2.

SPECIES GROUP APERTUS

altivolans Evans. IIl., Ala., La., Tex., Ariz., Calif.; Mexico (Sonora).
Dissomphalus altivolans Evans, 1954. Ent. Soc. Wash., Proc. 56: 307, figs. 7, 21. 6.

Taxonomy: Evans, 1962. Ent. Soc. Wash., Proc. 64: 76. 9.
apertus Kieffer. N. Y., Ohio and Kans., south into Mexico (Cordoba).
Dissomphalus apertus Kieffer, 1914. Soe. Ent. France, Bul. 1: 50. 6.
foveolatus (Brown and Cheng). Md., Va., Ga., Ill., Miss., Ark.
Psilobethylus foveolatus Brown and Cheng, 1952. Psyche 58: 143, fig. 1. 2.

nigrescens Evans. Tex., ?Ark., La., Ala.
Dissomphalus nigrescens Evans, 1954. Ent. Soc. Wash., Proe. 56: 308, figs. 8, 14. 3.

Taxonomy: Evans, 1962. Ent. Soc. Wash., Proc. 64: 75-76 (putative female).
Genus PSEUDISOBRACHIUM Kieffer

Genus PSEUDISOBRACHIUM Subgenus PS9EUDISOBRACHIUM Kieffer

Pseudisobrachium Kieffer, 1904. Mus. Civ. Stor. Nat. Genova, Ann. (8a) 1: 368.
Type-species: Pseudisobrachium laticeps Kieffer. Desig. by Kieffer, 1906.
Monepyris Kieffer, 1905. Soc. Sci. Bruxelles, Ann. 29: 101, 124.
Type-species: Epyris halidayi Westwood. Monotypic.
Xestobethylus Cameron, 1909. Amer. Ent. Soc., Trans. 35: 449.
Type-species: Xestobethylus pallidipes Cameron. Monotypic.
Plutobethylus Kieffer, 1910. Soc. Ent. France, Ann. 79: 51.
Type-species: Plutobethylus distans Kieffer. Orig. desig.
Lyssepyris Kieffer, 1913. Portici Lab. Zool. Gen. e Agr., Bol. 7: 108.
Type-species: Holepyris flavicornis Kieffer. Monotypic.
Xantepyris Kieffer, 1913. Portici Lab. Zool. Gen. e Agr., Bol. 7: 108.
Type-species: Epyris flaviventris Kieffer. Monotypic.
Lissepyris (!) Kieffer, 1914. Das Tierreich, Lief. 41, p. 236. Typ. error.
Xanthepyris Kieffer, 1914. Das Tierreich, Lief. 41, p. 417. Emend.
Parisobrachium Kieffer, 1914. Das Tierreich, Lief. 41, p. 424.
Type-species: Rhabdepyris (?) albipes Kieffer. Monotypic.

Only the typical subgenus occurs in North America.

Females of several species have been found in ant nests, but unsuccessful laboratory testing
on ant larvae suggests that the wasps parasitize myrmecophilous beetle larvae occurring in the
ant nests.

1218 Hymenoptera in America North of Mexico

Revision: Evans, 1961. Mus. Compar. Zool, Bul 126: 211-318, 5 pls.
Species Group CRassuM

crassum Evans. Tex. (Brownsville, San Antonio).

Pseudisobrachium crassum Evans, 1961. Mus. Compar. Zool, Bul. 126: 239, fig. 13. ¢-.
texanum Evans. Tex. (Harlingen, Brownsville, Wharton, San Antonio).

Pseudisobrachium teranum Evans, 1961. Mus. Compar. Zool, Bul. 126: 240, figs. 48, 53. ¢.

Species Group OCCIDENTALE
eastaneum Evans. Calif. (San Diego).
Pseudisobrachium castaneum Evans, 1961. Mus. Compar. Zool, Bul. 126: 248, fig. 56. ¢.
matthewsi Evans. Tex. N. Mex. Ariz.
Pseudisobrachium matthewsi Evans, 1961. Mus. Compar. Zool, Bul. 126: 249, fig. 36. ¢,
(?).
occidentale Evans. Calif, Ariz. Associated with dolichoderine ants.
Pseudisobrachium occidentale Evans, 1961. Mus. Compar. Zool, Bul. 126: 244, figs. 14, 55.
6, 2 (@.
SPEciES Group OBSCURUM

cearolinianum Evans. S. C., Fla.

Pseudisobrachium carolinianum Evans, 1961. Mus. Compar. Zool, Bul. 126: 254. ¢.
gibbosum Evans. N. Mex. (Rodeo).

Pseudisobrachium gibbosum Evans, 1961. Mus. Compar. Zool, Bul. 126: 255. 3.
obscurum Evans. Tex., Ariz, Nev., Utah, Calif; Mexico (Chihuahua, Baja California, Nayarit).

Pseudisobrachium obscurum Evans, 1961. Mus. Compar. Zool, Bul. 126: 258, figs. 16, 36.

6, 2 ().

otiosum Evans. Ariz.; Mexico (Sonora, Nayarit).

Pseudisobrachium otiosum Evans, 1961. Mus. Compar. Zool, Bul. 126: 256. 3.
pallidum Evans. Ariz. Calif.; Mexico (Sonora).

Pseudisobrachium pallidum Evans, 1961. Mus. Compar. Zool, Bul. 126: 258, fig. 17. 3.
werneri Evans. Ariz.; Mexico (Sonora, Sinaloa).

Pseudisobrachium werneri Evans, 1967. Ent. News 78: 20. 3.

SPEcIES Group PROLONGATUM

arenarium Evans. N. J., Pa. (?), N. C., Ga, Fla. (?), Ala, Miss. Mo. (?), IIL Associated with
ponerine ants.
Pseudisobrachium arenarium Evans, 1961. Mus. Compar. Zool, Bul. 126: 263, figs. 20, 39,
57. 3, 2 (2).
prolongatum (Provancher). N. B. and N. S. west to B. C, and Wash., south to N. C, Ky. IL
and Iowa. Associated with formicine and myrmicine ants.
Bethylus prolongatus Provancher, 1881. Nat. Canad. 12: 265.“2”= ¢.
Isobrachtum magnum Ashmead, 1893. U. S. Natl Mus., Bul. 45: 36. ¢.
Isobrachium myrmecophilum Ashmead, 1893. U. S. Natl Mus. Bul. 45: 37. 2, (¢ misdet.).
Isobrachium mandibulare Ashmead, 1893. U. S. Natl. Mus., Bul = 38. 2, (¢ misdet.).
Isobrachium montanum Ashmead, 1893. U. S. NatL Mus., Bul. 45: 39. 2, (¢ misdet.).
Pseudisobrachium rugosulum Fouts, 1928. Ent. Soc. Wash. Pe 30: 124. 3.
Pseudisobrachium agilis Whittaker, 1928. Roy. Ent. Soe. London, Trans. 76: 386, pl. 13,
fig. 4. ¢

SPEcIES Group CARBONARIUM

ashmeadi Evans. Mass., Ont., Mich. and Kans., south to Fla, Ala. and La. Host: Associated
with formicine and dolichoderine ants.
Pseudisobrachium ashmeadi Evans, 1961. Mus. Compar. Zool, Bul. 126: 275, figs. 23, 33,

58, 63. d, 2 (2).
carbonarium (Ashmead). Md. south to Ga, Ala, W. Va. Ky. (?), IIL, Kans. N. Dak. (?).
Epyris carbonarius Ashmead, 1893. U. S. Natl Mus. Bul. 45: 59, pl. 4, fig. 4. ¢.

Taxonomy: Evans, 1961. Mus. Compar. Zool, Bul. 126: 271, figs. 23, 37. ¢, 2 (2).

Superfamily BETHYLOIDEA 1219

krombeini Evans. Tex., N. Mex.; Mexico (Sonora).
Pseudisobrachium krombeini Evans, 1961. Mus. Compar. Zool., Bul. 126: 287, figs. 27, 50.
é.
minimum Evans. N. Mex., Ariz.; Mexico (Sinaloa, Michoacan, Mexico).
Pseudisobrachium minimum Evans, 1961. Mus. Compar. Zool., Bul. 126: 280, fig. 24. d.
minutissimum Evans. N. Mex., Ariz.; Mexico (Morelos, Baja California), Guatemala.
Pseudisobrachium minutissimum Evans, 1961. Mus. Compar. Zool., Bul. 126, fig. 26. 3.
navajo Evans. Ariz. (Coconino and Yavapai Counties).
Pseudisobrachium navajo Evans, 1961. Mus. Compar. Zool., Bul. 126: 283, fig. 25. 3.
persimile Evans. Calif., B. C. (?).
Pseudisobrachium persimile Evans, 1961. Mus. Compar Zool., Bul. 126: 284, fig. 2. 3d, 2
(?).

SPECIES GROUP RUFIVENTRE

apache Evans. Ariz. (Santa Cruz, Gila and Pinal Counties).

Pseudisobrachium apache Evans, 1961. Mus. Compar. Zool., Bul. 126: 299, fig. 46. 3.
comanche Evans. Ariz., Tex.; Mexico (Sinaloa).

Pseudisobrachium comanche Evans, 1961. Mus. Compar. Zool., Bul. 126: 298, fig. 59. ¢.
emarginatum Evans. Tex. (Kerrville).

Pseudisobrachium emarginatum Evans, 1961. Mus. Compar. Zool., Bul. 126: 303, fig. 49. 3.
flavinervis Fouts. Tex., Ariz., Nev., Calif.; Mexico (Sonora, Nayarit).

Pseudisobrachium flavinervis Fouts, 1928. Ent. Soc. Wash., Proc. 30: 123. ¢.

Taxonomy: Evans, 1961. Mus. Compar. Zool., Bul. 126: 305, figs. 6, 31, 38, 62. d, @ (?).
flaviventre (Kieffer). Del. to Fla., west to Tex., Kans., Ill. Host: Associated with myrmicine

ants.
Epyris flaviventris Kieffer, 1904. Arkiv for Zool. 1: 526. ¢.

Taxonomy: Evans, 1961. Mus. Compar. Zool., Bul. 126: 288, figs. 41, 60. d, Q(?).

foutsi Evans. Tex., N. Mex. (?), Ariz. (?), Calif. (?); Mexico (Sonora, Coahuila).

Pseudisobrachium foutsi Evans, 1961. Mus. Compar. Zool., Bul. 126: 300, figs. 29, 61. ¢.
macrops Evans. Tex. (Cameron and Hidalgo Counties).

Pseudisobrachium macrops Evans, 1961. Mus. Compar. Zool., Bul. 126: 309. d.
pusillum Evans. La. (Shreveport), Ark. (?).

Pseudisobrachium pusillum Evans, 1961. Mus. Compar. Zool., Bul. 126: 297. 3, 2 (?).
rectangulatum Evans. Tex., Kans., Nebr.; Mexico (Nuevo Leon).

Pseudisobrachium rectangulatum Evans, 1961. Mus. Compar. Zool., Bul. 126: 304. ¢.
rufiventre (Ashmead). Mass. and N. Y. south to Fla. and La. Host: Associated with formicine

ants.

Isobrachium rufiventre Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 38. 9, d.

Pseudisobrachium flavicoxis Fouts, 1928. Ent. Soc. Wash., Proc. 30: 122. 2.

Pseudisobrachium puncticeps Fouts, 1928. Ent. Soc. Wash., Proc. 30: 123. d.

UNPLACED TAXA OF PSEUDISOBRACHIUM SUBGENUS PSEUDISOBRACHIUM KIEFFER

anomalum Evans. N. J., Va., Fla., Ill.
Pseudisobrachium anomalum Evans, 1961. Mus. Compar. Zool., Bul. 126: 311, figs. 8, 54,
64, 67. 3.
paucipunctata Fouts. Utah, Calif.
Pseudisobrachium paucipunctata Fouts, 1928. Ent. Soe. Wash., Proce. 30: 122. 2.
UNPLACED TAXA OF BETHYLIDAE

Bethylus musculus Say, 1836. Boston Jour. Nat. Hist. 1: 280. Ind.
Family SCLEROGIBBIDAE

So far as is known, members of this family are parasites of Embioptera. The wasp larvae
develop as external parasites of embiid nymphs in much the same manner as larval Dryinidae
and Rhopalosomatidae.

1220 Hymenoptera in America North of Mexico

Revision: Richards, 1939. Roy. Ent. Soc. London, Proc., Ser. B, Taxonomy, 8: 211-223.

Morphology: Reid, 1941. Roy. Ent. Soe. London, Trans. 91: 412-414, figs. 53-54 (female, male
thorax).

Genus PROBETHYLUS Ashmead

Probethylus Ashmead, 1902. Canad. Ent. 34: 270.
Type-species: Probethylus schwarzi Ashmead. Orig. desig.

Taxonomy: Shetlar, 1973. Ent. News 84: 205-206 (generic redescription, key to spp.).

Biology: Callan, 1939. Roy. Ent. Soc. London, Proc., Ser. B, Taxonomy, 8: 223-224
(Probethylus callani Rich. from Trinidad).

schwarzi Ashmead. Ariz.
Probethylus schwarzi Ashmead, 1902. Canad. Ent. 34: 270. 6.

Taxonomy: Shetlar, 1973. Ent. News 84: 206-208, figs. 1-4 (redescription male, female).
Biology: Shetlar, 1973. Ent. News 84: 208, fig. 5 (host, life history).

Family CHRYSIDIDAE

The cuckoo or ruby-tailed wasps constitute one of the most attractive families of Hymenop-
tera because of their brilliantly metallic coloration. Almost all of our species are either purple,
blue or green, but many Palaearctic species are marked also with golden or ruby in specific pat-
terns.

The behavior and life history also make this a very fascinating group. All species are parasitic,
and, as one common name implies, many of them exhibit behavior in the host nests similar to
that of cuckoos in the nests of their bird hosts. Members of the small subfamilies Cleptinae and
Amiseginae parasitize, respectively, the resting larvae of sawflies in their cocoons and eggs of
walking sticks. Almost all species belonging to the Elampinae, Chrysididinae and Parnopinae
have as their hosts solitary wasps or bees which nest in the ground or in cavities in wood or
which build mud cells; one exotic species of Chrysis L. is known to parasitize the resting larva of
the oriental moth in its cocoon. The parasite egg is deposited in the host cell while it is being
provisioned by the mother wasp or bee. In most species the newly hatched chrysidid larva
devours the host egg or young larva and then feeds on the provisions stored for the host. How-
ever, in species of Chrysura Dahlb. the parasite larva attaches to the host larva, sucks only a
small amount of body fluids, and does not devour the host larva until the latter attains full
growth and spins its cocoon. Eggs of Parnopes Latr. and Chrysis pellucidula Aar. are deposited
in the host nest while the host wasp is provisioning; these parasite larvae also devour the rest-
ing host larva in its cocoon. The female of Chrysis fuscipennis Br. chews a hole in the host mud
cell, oviposits therein, and the parasite larva develops on the resting larva of the host.

The arrangement of genera in the Chrysididinae and division into Species Groups in Chrysis
Linnaeus are by R. M. Bohart who also contributed new synonymy and distribution in both the
Chrysididinae and Elampinae.

SUBFAMILY CLEPTINAE

So far as is known, members of this subfamily are parasitic on resting larvae of sawflies in
their cocoons.

Revision: Aaron, 1885. Amer. Ent. Soe., Trans. 12: 209-248 (No. Amer. spp.). —Mocsary, 1889.
Monog. Chrysid., 643 pp. (spp. of world). —Ashmead, 1902. Canad. Ent. 34: 221-231 (gen.).
—Bischoff, 1913. Gen. Ins., Fase. 151, 86 pp., 5 pls. —Linsenmaier, 1959. Schweiz. Ent.
Ges., Mitt. 32: 1-232, 711 figs. (European spp.).

Biology: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 141-168, 2 pls. (spp. of coastal N. C.).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 437-475, figs. 87, 88, 128-134 (life
history of spp. attacking wasps and bees nesting in borings in wood.).

Superfamily BETHYLOIDEA 1221

Genus CLEPTES Latreille

Genus CLEPTES Subgenus CLEPTES Latreille

Cleptes Latreille, 1802. Hist. Nat. Crust. Ins. 3: 316.
Type-species: Sphex semiaurata Linnaeus. Monotypic.

semiauratus (Linnaeus). New Jersey (Metuchen); Europe. Adventive from Europe. Host:
Pristiphora sp., Nematus spp., Pachynematus spp. in Europe.
Sphex semiaurata Linnaeus, 1761. Fauna Suec., Ed. 2, p. 413. d.
Biology: Gauss, 1964. Ztschr. f. Angew. Ent. 54: 225-282, figs. 1-4 (life history, European
hosts).

Genus CLEPTES Subgenus LEIOCLEPTES Moczar

Cleptes subg. Leiocleptes Moczar, 1962. Acta. Zool., Acad. Sci. Hungaricae 8: 118.
Type-species: Cleptes nitidulus Fabr. Orig. desig.

alienus Patton. Mont., Wyo.
Cleptes aliena Patton, 1879. Canad. Ent. 11: 66. 9.

blaisdelli Bridwell. Calif.
Cleptes blaisdelli Bridwell, 1919. Hawaii. Ent. Soc., Proe. 4: 37. ¢.

insperatus Aaron. Mont.
Cleptes insperata Aaron, 1885. Amer. Ent. Soc., Trans. 12: 212. 2.

Genus CLEPTES Subgenus MELANOCLEPTES Moczar

Cleptes subg. Melanocleptes Moczar, 1962. Acta Zool., Acad. Sci. Hungaricae 8: 121.
Type-species: Cleptes morawitzi Rad. Orig. desig.
provancheri Aaron. Canada, “Wash. Ter.”, Idaho, Colo. Host: Neodiprion sp. near scutellatus
Rohw.
Cleptes americana Provancher, 1881. Nat. Canad. 12: 304. 2. Preoce.
Cleptes Provancheri Aaron, 1885. Amer. Ent. Soe., Trans. 12: 212. N. name.
Biology: Smith, 1962. Pan-Pacifie Ent. 38: 189.
purpuratus Cresson. Wash., Oreg., Calif., Colo., Nev. Host: Neodiprion sp. in fulviceps
complex, N. sp.
Cleptes purpurata Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. x. 2.
Cleptes Americana Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. x. d, 2.
Biology: Dahlsten, 1961. Canad. Ent. 93: 192-193. —Dahlsten, 1967. Ecology 48: 287.

speciosus Aaron. Mont.
Cleptes speciosa Aaron, 1885. Amer. Ent. Soc., Trans. 12: 212. 2.

SUBFAMILY AMISEGINAE

Members of this subfamily are parasitic in the eggs of walking sticks.

Taxonomy: Krombein, 1957. Amer. Ent. Soc., Trans. 82: 147-215, 4 pls. (generic
reclassification of world fauna).

Genus MESITIOPTERUS Ashmead

Mesitiopterus Ashmead, 1902. Canad. Ent. 34: 231.
Type-species: Mesitiopterus Kahlii Ashmead. Orig. desig.

floridensis Krombein. Fla. (Gainesville).
Mesitiopterus floridensis Krombein, 1960. Amer. Ent. Soc., Trans. 86: 28, fig. 6. d, °.

kahlii Ashmead. N. Y. south to N. C., Wis., Minn., Kans., Ariz. Host: Diapheromera femorata
Say, eggs.
Mesitiopterus Kahlii Ashmead, 1902. Canad. Ent. 34: 231. d.

Biology: Milliron, 1950. Ent. Soc. Wash., Proe. 52: 47.

1222 Hymenoptera in America North of Mexico

Genus MICROSEGA Krombein

Microsega Krombein, 1960. Amer. Ent. Soc., Trans. 86: 31.
Type-species: Microsega bella Krombein. Orig. desig.

bella Krombein. Tex., Okla.
Microsega bella Krombein, 1960. Amer. Ent. Soc., Trans. 86: 32, figs. 1, 2, 5. 2.

Genus ADELPHE Mocsary

Adelphe Mocsary, 1890. Termes. Fuzetek 13: 46.
Type-species: Adelphe mexicana Mocsary. Monotypic.
Pseudepyris Ducke, 1902. Ztschr. Hym. Dipt. 2: 204.
Type-species: Pseudepyris paradoxa Ducke. Monotypic.
Adelpha Schulz, 1906. Spolia Hym., p. 153. Proposed unnecessarily for Adelphe Mocsary.
Preoce.
Parachrysis Kieffer, 1910. Soc. Ent. France, Ann. 78: 287. Preocc.
Type-species: Parachrysis metallica Kieffer. Monotypic.
anisomorphae Krombein. Ga., Fla., La. Host: Anisomorpha ferruginea (Beauv.), eggs.
Adelphe anisomorphae Krombein, 1960. Amer. Ent. Soc., Trans. 86: 35, fig. 3. d, 2.

SUBFAMILY ELAMPINAE

These predominantly small wasps are parasites of solitary ground- and wood-nesting wasps
and bees.

Genus OMALUS Panzer

Most of these species parasitize twig-nesting wasps although one has been reared from a
ground-nesting species. Authenticated host records all pertain to aphid-storing pemphredonine
wasps. European records of other wasps and bees as hosts of Omalus are probably erroneous
and based on mixed nests where supersedure by or of the pemphredonine had occurred.

Revision: Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53: 235-250, 30 figs.
Genus OMALUS Subgenus OMALUS Panzer

Omalus Panzer, 1801. Faunae Ins. German. H. 85, no. 13.
Type-species: Chrysis aenea Fabricius. Monotypic.
Homalus Saunders, 1873. Ent. Soe. London, Trans. p. 411. Emend.
Ellampus subg. Dictenulus Semenov, 1932. Soc. Ent. Ross., Hor. 42: 6.
Type-species: Ellampus (Dictenulus) specularis Semenov. Orig. desig.
aeneus (Fabricius). Canada and U. S. south to Ga., N. Mex. and Ariz., chiefly in Canadian and
Transition Zones, occasionally Upper Austral; Europe. Host: Passaloecus cuspidatus
Sm.; P. sp.; Pemphredon sp. Predator: Philanthus pulcher D. T.
Chrysis aeneus Fabricius, 1787. Mant. Ins. 1: 284.
Omalus laeviventris Cresson, 1865. Ent. Soc. Phila., Proc. 4: 303. 2.
Elampus cyanescens Provancher, 1881. Nat. Canad. 12: 303. 2.
Omalus diversus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 213. °.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 97 (host record). —Krombein, 1967.
Trap-nesting wasps and bees, p. 442 (life history). —Evans, 1973. Great Basin Nat. 33: 155
(host).

butleri Bohart and Campos. Ariz., N. Mex., Utah, Colo., B. C., Ind., N. Y.
Omalus (Omalus) butleri Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53: 240, figs. 6,
ie Or
glomeratus (du Buysson). Calif. to Wash., east to Idaho, Colo., N. Mex. Host: Stigmus i.

inordinatus Fox.
Ellampus glomeratus du Buysson, 1901. K. K. Naturhist. Hofmus., Ann. 16: 98. ¢.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 95 (host record).

Superfamily BETHYLOIDEA 1223

iridescens (Norton). Throughout Canada and-U. S., in Canadian, Transition and Austral Zones.
Host: Stigmus i. inordinatus Fox, S. americanus Pack., Psenulus trisuleus (Fox),
Passaloecus sp.
Elampus iridescens Norton, 1879. Amer. Ent. Soe., Trans. 7: 234. d.
Elampus marginatus Provancher. 1881. Nat. Canad. 12: 304. 2. Preoce.

Biology: Davidson, 1895. Psyche 7: 271 (host record). —Krombein, 1958. U. S. Dept. Agr.,
Monog. 2, Sup. 1: 94. —Krombein, 1967. Trap-nesting wasps and bees, pp. 442-443 (life
history).

Genus OMALUS Subgenus PSEUDOMALUS Ashmead

Pseudomalus Ashmead, 1902. Canad. Ent. 34: 229.
Type-species: Omalus semicircularis Aaron. Monotypic.
auratus (Linnaeus). N. Y. south to S. C., Ohio, Ind.; widely distributed in the Palearctic
Region. Undoubtedly adventive in U. S. Host: Pemphredon |. lethifer (Shuck.),
Passaloecus turionum Dahlb.; Perithous divinator (Rossi).
Sphex aurata Linnaeus, 1758. Syst. Nat., ed. 10, p. 572.
Hedychrum sinuosum Say, 1828. Contrib. Maclur. Lye. Phila. 1: 82. 2.

Taxonomy: Krombein, 1959. Brooklyn Ent. Soc., Bul. 54: 95-96.

Biology: Thomas, 1962. Ent. News 73: 217-218.
janus (Haldeman). Canada and U. S. south to Calif., Ariz., N. C., chiefly in Canadian and
Transition Zones. Host: Pemphredon concolor Say, P. errans Roh.
Hedychrum janus Haldeman, 1844. Acad. Nat. Sci. Phila., Proc. 1: 53. @.
Elampus coruscans Norton, 1879. Amer. Ent. Soc., Trans. 7: 234. 2.
Omalus corruscans (!) Aaron, 1885. Amer. Ent. Soc., Trans. 12: 214.

Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 94 (host record).
purpuratus (Provancher). Alaska, Canada and U. S. south to Calif., Ariz. and Ga. Host:

Stigmus americanus Pack., Pemphredon 1. lethifer (Shuck.), P. harbecki Roh. ?,
Passaloecus cuspidatus Sm. 7

Elampus purpuratus Provancher, 1881. Nat. Canad. 12: 302. 2.

Elampus purpurascens (!) Provancher, 1881. Nat. Canad. 12: 303.

Omalus (Pseudomalus) macswaini Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53:
244, fig. 22. g, 2. N. syn. (W. R. M. Mason).

Biology: Krombein, 1960. Ent. News 71: 32-33 (host record). —Parker and Bohart. 1966.
Pan-Pacifie Ent. 42: 94 (host record). —Evans, 1973. Great Basin Nat. 33: 155 (host).
semicircularis Aaron. Colo., Nev.
Omalus semicircularis Aaron, 1885. Amer. Ent. Soc., Trans. 12: 215. ¢.
trilobatus Bohart and Campos. B. C. and Idaho south to Calif., Ariz. and Colo., in Canadian,
Transition and Upper Sonoran Zones. Host: Pemphredon giffardi (Roh.).
Omalus (Pseudomalus) trilobatus Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53:
245, figs. 4, 15, 25, 28. 6, @.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (host record).

Genus OMALUS Subgenus DIPLORRHOS Aaron

Diplorrhos Aaron, 1885. Amer. Ent. Soc., Trans. 12: 216.
Type-species: Diplorrhos plicatus Aaron. Monotypic.
cressoni (Aaron). Oreg., Calif., Ariz., Colo., Utah, Mont., Nebr. Host: Diodontus occidentalis
Fox, Pemphredon giffardi (Roh.), Stigmus i. inordinatus Fox, Passaloecus cuspidatus
Sm.
Elampus cressoni Aaron, 1885. Amer. Ent. Soc., Trans. 12: 215. d, 2.

Biology: Powell, 1964. Wasmann Jour. Biol. 21: 172-173 (host record). —Parker and Bohart,
1966. Pan-Pacific Ent. 42: 94-95 (host record). —Evans, 1973. Great Basin Nat. 33: 155
(host).

1224 Hymenoptera in America North of Mexico

downeyi Bohart and Campos. Oreg., Calif., Nev., chiefly in Canadian Zone.
Omalus (Diplorrhos) downey Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53: 246,
fiess Dy Saree
granti Bohart and Campos. Calif., Ariz., N. Mex., Nev., Colo.
Omalus (Diplorrhos) granti Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53: 247, fig.
1955 ee
intermedius (Aaron). Mont., Md. Host: Diodontus virginianus (Roh.).
Notozus intermedius Aaron, 1885. Amer. Ent. Soc., Trans. 12: 218. 3.

Biology: Krombein, 1963. Ent. Soc. Wash., Proc. 65: 264 (host record).
krombeini Bohart and Campos. Ariz., southern Calif.
Omalus (Diplorrhos) krombeini Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53: 247,
figs. 3, 13. 5, 9.
pice eS Canada, western and northern U. S. south to Calif., N. Mex., Minn.,
ich., N. Y.
Diplorrhos plicatus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 216. 3, @.
seminudus (Aaron). Alaska, B. C. south to Calif., Idaho, Ont.
Notozus seminudus Aaron, 1885. Amer. Ent. Soe., Trans. 12: 218. 6, 9°.
speculum (Say). Wash., Oreg., Calif., Colo.
Hedychrum speculum Say, 1836. Boston Jour. Nat. Hist. 1: 225. d.
telfordi Bohart and Campos. Tex. to southern Calif., Nev.
Omalus (Diplorrhos) telfordi Bohart and Campos, 1960. Ent. Soc. Amer., Ann. 53: 249, figs.
VASAT:
variatus (Aaron). Western N. Amer. in Canadian, Transition and Upper Sonoran Zones. Host:
Stigmus i. inordinatus Fox.
Elampus variatus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 215. d, °.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 95 (host record).
Genus NOTOZUS Foerster

Elampus Spinola, 1806. Insectorum Liguriae, v. 1, p. 10.
Type-species: Chrysis panzeri Fabricius. Desig. by Latreille, 1810.
Notozus Foerster, 1853. Naturh. Ver. Rheinlande, Verh. 10: 351.
Type-species: Notozus frivaldszkii Foerster. Desig. by Bodenstein, 1939 (~Chrysis
panzeri Fabricius).
Ellampus (!) Agassiz, 1846. Nomencl. Zool., Index Univ., pp. 135, 136.

The few available host records indicate that species of this genus parasitize ground-nesting
sphecoid wasps.
Taxonomy: Huber, 1975. Bul. Zool. Nomencl. 32: 181-187 (petition to suppress Elampus
Spinola, 1806, and place Notozus Foerster, 1853, on Official List of Generic Names).
aaroni (Bodenstein), n. comb. Man., Sask., Alta., Mont., Colo., Calif.
Notozus productus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 219. 2. Preoce.
Elampus aaroni Bodenstein, 1951. U. S. Dept. Agr., Monog. 2: 719. N. name.
connexus Viereck. Kans., Calif.
Notozus connexus Viereck, 1906. Amer. Ent. Soc., Trans. 32: 192.
hyalinus Aaron. N. C., Fla., Mich., Ill., Minn., Iowa, Mo., S. Dak., Kans., Tex., Okla., Colo., N.
Mex., Mont., Nev., Utah, Ariz. :
Notozus hyalinus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 218. °.
marginatus Patton. Entire U. S. and Canada.
Notozus marginatus Patton, 1879. Canad. Ent. 11: 66.
nitidus Aaron. West and Southwest, east to Iowa., Man.; Mexico (Sonora).
Notozus nitidus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 218. d.
versicolor (Norton), n. comb. Calif., Mont., Ariz., N. Mex., “Dakota,” Kans., Man.; Mexico (Baja

California).
Elampus versicolor Norton, 1879. Amer. Ent. Soc., Trans. 7: 235. 6.

Superfamily BETHYLOIDEA 1225

viridicyaneus (Norton), n. comb. Transcont. in Canad., Transit. and Austral Zones. Host:
Hoplisoides costalis (Cress.). Predator: Philanthus pulcher D. T.
Elampus viridicyaneus Norton, 1879. Amer. Ent. Soc., Trans. 7: 235. 9.
Elampus spinosus Provancher, 1881. Nat. Canad. 12: 302. °.

Genus HOLOPYGA Dahlbom

Holopyga Dahlbom, 1845. Dispos. Method. Spec. Hym., pt. 2, p. 4.
Type-species: Holopyga amoenula Dahlbom. Desig. by Ashmead, 1902 (=Chrysis
gloriosa Fabricius).
Halopyga (!) Tournier, 1878. Schweiz. Ent. Gesell., Mitt. 5: 305.

Members of this genus are parasites of ground-nesting sphecid wasps.

hora Aaron. Calif., Idaho, Nev., N. Mex., Colo., Wyo., Ariz., Mont., Kans., Tex.; Mexico (Baja
California, Oaxaca).
Holopyga horus Aaron, 1885. Amer. Ent. Soc., Trans. 12: 220. 3, 9.
Hedychrum continuum Aaron, 1885. Amer. Ent. Soc., Trans. 12: 224. ¢, 2. N. syn. (R. M.
Bohart).
lazulina Dahlbom. IIl., Tex. south to Argentina.
Holopyga lazulina Dahlbom, 1854. Hym. Europaea, v. 2, p. 49.
ventralis (Say). Widely distributed throughout U. S., Canada and north. Mexico. Host: Bicyrtes
fodiens (Handl.), B. quadrifasciata (Say). Predator: Philanthus pulcher D. T.
Hedychrum ventrale Say, 1824. In Keating, Narr. Long’s 2nd. Exped., v. 2, App., p. 330.
Hedychrum asperum Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 52. 2. N. syn. (R. M.
Bohart).
Holopyga Dohrni Dahlbom, 1854. Hym. Europaea, v. 2, p. 48.
Holopyga compacta Cresson, 1865. Ent. Soc. Phila., Proe. 4: 304.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 160, 169 (hosts).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14, pp. 42-48, pl. 13 (male
genitalia).

Genus MUESEBECKIDIUM Krombein

Muesebeckidium Krombein, 1969. Ent. Soc. Wash., Proce. 71: 352.
Type-species: Hedychrum obsoletum Say. Orig. desig.
obsoletum (Say). Widely distributed in North America in U. and L. Austr. Zones east of 100th
meridian.

Hedychrum obsoleta Say, 1836. Boston Jour. Nat. Hist. 1: 284.? @.

Hedychrum zimmermanni Dahlbom, 1845. Dispos. Method. Spec. Hym., pt. 2, p. 2. d.
occidentale Krombein. N. Mex., Ariz.; Mexico (Sonora, Durango).

Muesebeckidium occidentale Krombein, 1969. Ent. Soc. Wash., Proc. 71: 357, fig. 13. d, 2.

Genus PSEUDOLOPYGA Krombein

Pseudolopyga Krombein, 1969. Ent. Soc. Wash., Proc. 71: 357.
Type-species: Holopyga taylori Bodenstein. Orig. desig.
taylori (Bodenstein). Mo., N. Mex., Ariz., Nev., Idaho, Calif. Host: Solierella affinis blaisdelli
(Bridw.); S. peckhami (Ashm.).
Holopyga taylori Bodenstein, 1939. Ent. News 50: 19. 3, @.
Hedychridium carrilloi Bohart and Brumley, 1967. Pan-Pacifie Ent. 43: 232, figs. 1-4. 6,
Ok

Taxonomy: Carrillo and Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 677-679, figs. 8-11 (egg,
larvae, pupa, cocoon).

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 443 (host). —Carrillo and
Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 677-679, figs. 8-12 (life historv)

1226 Hymenoptera in America North of Mexico
Genus HEDYCHRIDIUM Abeille de Perrin

Hedychridium Abeille de Perrin, 1878. Diagn. Chrysid. Nouv., p. 3.
Type-species: Hedychrum minutum Lepeletier. Desig. by Ashmead, 1902
(=Chrysis ardens Coquebert).

Most species of Hedychridium parasitize ground-nesting wasps and bees, but one has been
recorded as parasitizing a twig-nesting sphecid wasp.

amabile Cockerell. Calif., Ariz., Nev., Utah, N. Mex.; Mexico (Baja California, Sonora).
Hedychridium amabile Cockerell, 1903. Canad. Ent. 35: 262.
caeruleum (Norton). N. J., Ind., Ill., lowa, N. Dak., Mont.
Hedychrum caeruleum Norton, 1879. Amer. Ent. Soc., Trans. 7: 239. 2.
cockerelli du Buysson. Colo., S. Dak.
Hedychridium cockerelli du Buysson, 1906. Rev. Ent. Caen 25: 111. 6.
dimidiatum (Say). Throughout U.S. and Canada; Mexico.
Hedychrum dimidiatum Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, App., p. 330,
n. 2.
Chrysis Mexicana Cameron, 1888. Biol. Cent.-Amer., Hym., v. 1, p. 460. N. syn. (R. M.
Bohart).
Holopyga (Hedychridium) mexicana Moesary, 1911. Mus. Nat. Hungarici, Ann. 9: 448. 6.
N. syn. (R. M. Bohart).
fletcheri Bodenstein. Western U. S. and Canada to Ind. and Sask., Ala., Fla. Host: Tachysphex
similis Roh. Predator: Philanthus pulcher D. T.
Hedychrum viride Cresson, 1865. Ent. Soc. Phil., Proc. 4: 306. Preoce.
Hedychridium fletcheri Bodenstein, 1951. U. S. Dept. Agr. Monog., 2: 720. N. name.

Biology: Kurezewski, 1967. Kans. Ent. Soc., Jour. 40: 278-284, 1 fig. (life history, behavior).
semirufum (Cockerell). N. Mex., Ariz.
Holopyga semirufa Cockerell, 1896. Psyche (sup.) 7: 17. 9.
solierellae Bohart and Brumley. Western U. S., B. C.; Mexico (Baja California). Host:
Solierella affinis blaisdelli (Bridw.); S. peckhami (Ashm.).
Hedychridium solierellae Bohart and Brumley, 1967. Pan-Pacific Ent. 43: 234. 6, 9.

Taxonomy: Carrillo and Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 679-680, figs. 13-15 (egg,
larvae, pupa, cocoon).

Biology: Carrillo and Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 679-680, figs. 13-15 (life
history).

Genus HEDYCHRUM Latreille

Hedychrum Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 317.
Type-species: Hedychrum lucidulum Latreille. Monotypie (=Sphex nobile Scopoli).
Hedycrum (!) Say, 1825. In Keating, Narr. Long’s 2nd Exped., v. 2, App., p. 65.

Wasps of this genus are parasitic on ground-nesting wasps and bees.

confusum du Buysson. U.S. and Canada west to N. Mex., Wyo., Sask. Host: Cerceris halone
Bks., C. architis Mick.
Hedychrum confusum du Buysson, 1891. Rev. Ent. Caen 10: 30. 6, 2.
Hedychrum nearcticum Moesary, 1914. Mus. Nat. Hungarici, Ann. 12: 11. 9. N. syn. (R. M.
Bohart).
cupricolle Cresson. Alta., Colo., Kans., Nebr., Minn.
Hedychrum cupricolle Cresson, 1865. Ent. Soc. Phila., Proc. 4: 305. 9.
louisianiae Norton. N. C., Fla. west to N. Mex., Mo., Kans., Colo.; Mexico (Mexico).
Hedychrum Louisianiae Norton, 1879. Amer. Ent. Soc., Trans. 7: 238. ¢.
Hedychrum Louisianae(!) Aaron, 1885. Amer. Ent. Soc., Trans. 12: 223. Emend.
nigropilosum Mocsary. Western U.S. and Canada to Wyo., Colo., Alta.
Hedychrum nigropilosum Moesary, 1889. Monog. Chrysid., p. 162. 3.

parvum Aaron. Transcontinental in U. S., Man., Alta., B. C., N. W. T.
Hedychrum violaceum var. parvum Aaron, 1885. Amer. Ent. Soc., Trans. 12: 2238.

Superfamily BETHYLOIDEA 1227

Hedychridium polygoni Rohwer, 1909. Psyche 16: 87. °. N. syn. (R. M. Bohart).
violaceum Brulle. Throughout U. S., Canada and northern Mexico.
Hedychrum violaceum Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 51. d.
Hedychrum affinissimum Bischoff, 1910. Berlin Zool. Mus., Mitt. 4: 444. °. N. syn. (R. M.
Bohart).
wiltii Cresson. Ark., Tex., Colo., Ariz.
Hedychrum Wiltii Cresson, 1865. Ent. Soe. Phila., Proc. 4: 305. d.

SUBFAMILY CHRYSIDIDINAE

Members of this subfamily have as hosts a wide variety of wasps and bees, including species
that nest in the ground, in cavities in wood, or that construct nests of mud.

Genus CHRYSIS Linnaeus

Genus CHRYSIS Subgenus CHRYSIS Linnaeus

Chrysis Linnaeus, 1761. Fauna Suecica, Ed. 2, p. 414.
Type-species: Sphex ignita Linnaeus. Desig. by Latreille, 1810.
Chrysis subg. Tetrachrysis Lichtenstein, 1876. Petites Nouv. Ent., v. 2, p. 27.
Type-species: Chrysis aeruginosa Dahlbom. Desig. by Ashmead, 1902 (=Chrysis
succincta Linnaeus).
Chrysis subg. Hexachrysis Lichtenstein, 1876. Petites Nouv. Ent., v. 2, p. 27.
Type-species: Chrysis micans Rossi. Desig. by Bodenstein, 1939 (=Chrysis
sexdentata Christ).
Chrysis subg. Actinochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,
Abhandl. u. Ber. 23: 74.
Type-species: Chrysis succincta Linnaeus. Orig. desig.
Chrysis subg. Cymatochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,
Abhandl. u. Ber. 23: 74.
Type-species: Chrysis viridula Linnaeus. Orig. desig.

Only the typical subgenus occurs in North America, but it has been divided into a number of
species groups.

Taxonomy: Bohart, 1962. Acta Hym. 1: 361-375, 25 figs. (hexadentate species).
SPECIES GRoUP DORSALIS

alfkenella du Buysson. Western U.S. to Nebr., Ark.
Chrysis (Tetrachrysis) Alfkenella du Buysson, 1904. Rev. Ent. Caen 23: 266. d.

dorsalis Aaron. Transcontinental in U. S. and Canada. Host: Photopsis sp.
Chrysis dorsalis Aaron, 1885. Amer. Ent. Soc., Trans. 12: 234. 6, °.

Biology: Ferguson, 1962. Univ. Calif. Pubs. Ent. 27: 29 (host record).
meta Aaron. Colo., Wyo., Mont., Alta.
Chrysis meta Aaron, 1885. Amer. Ent. Soc., Trans. 12: 234. d, °.
montana Aaron. Ont. and Maine south to Fla., Tex., Mich., Ill., Iowa, Mo., Kans., Nebr., Colo.,
Mont., Idaho, Utah,Calif.
Chrysis montana Aaron, 1885. Amer. Ent. Soc., Trans. 12: 234. ?.
Chrysis hirsuta Aaron, 1885. Amer. Ent. Soc., Trans. 12: 235. d, 2. Preoce. N. syn. (R. M.
Bohart).
Chrysis (Tetrachrysis) aaroni Moesary, 1889. Monog. Chrysid., p. 386.
Chrysis (Tetrachrysis) equidens Viereck, 1906. Amer. Ent. Soc., Trans. 32: 193. N. syn. (R.
M. Bohart).
Chrysis (Hexachrysis) nana Moesary, 1913. Mus. Nat. Hungarici, Ann. 11: 33. d. N. syn.
(R. M. Bohart).

Taxonomy: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 145-146, fig. 6 (equidens
redescription). —Bohart, 1962. Acta Hym. 1: 375 (synonymy of hirsuta and nana).

provancheri Schulz. Maine, Ont., Minn., S. Dak., N. Dak., Alta., N. W. T.
Chrysis aurichalcea Provancher, 1881. Nat. Canad. 12: 300. 2. Preoce.

1228 Hymenoptera in America North of Mexico

Chrysis Provancheri Schulz, 1906. Spolia Hym., p. 154. N. name.
scitula Cresson. Colo., Utah, Calif.
Chrysis scitula Cresson, 1865. Ent. Soe. Phila., Proc. 4: 308. °.
Chrysis Californica Gribodo, 1879. Mus. Civ. Stor. Nat. Genova, Ann. 14: 336. 2. N. syn.
(R. M. Bohart).
tenuicornis Taylor. Oreg., Calif., Ariz., N. Mex.; Mexico (Baja California, Jalisco, Durango).
Chrysis (Tetrachrysis) tenuicornis Taylor, 1924. Calif. Acad. Sci., Proc. (4)18: 327. 9.

Taxonomy: Bohart, 1962. Acta Hym. 1: 378, figs. 3, 15.
SPECIES GROUP CONICA

conica Brulle. D. C. to Fla., Ky., Tenn., Tex., Ariz.; Mexico. Host: Eumenes fraternus Say in
mud nests.
Chrysis conica Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 32. 6, @.
Chrysis peracuta Aaron, 1885. Amer. Ent. Soc., Trans. 12: 237. 2.
Chrysis peregrina du Buysson, 1887. Rev. d’Ent. 6: 188. 9.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14, pp. 42, 48, pl. 13, fig. O
(male genitalia).

SPECIES GROUP DERIVATA

amala Rohwer. Mass., Iowa, Minn., Kans., Colo., N. Mex., Utah, Nev., Mont., Wash.

Chrysis (Tetrachrysis) amala Rohwer, 1909. Psyche 16: 91. @.
antennalis Mocsary. Tex., N. Mex., Ariz.; northern Mexico.

Chrysis (Tetrachrysis) antennalis Mocsary, 1912. Mus. Nat. Hungarici, Ann. 10: 564. d.
barri Bohart. Idaho, Nev., Calif. Host: Parancistrocerus toltecus (Sauss.) nesting in twig

cavities.
Chrysis barri Bohart, 1966. Biol. Soc. Wash., Proc. 79: 1382. d, @.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (host record).
derivata du Buysson. Transcontinental in U. S. and southern Canada. Host: Ancistrocerus
catskill halophilus Vier.; Euodynerus foraminatus apopkensis (Robt.), E. f. scutellaris
(Sauss.); Symmorphus cristatus nevadensis (Cam.). Undoubtedly parasitizes a number
of other eumenid wasps nesting in cavities in twigs or borings in wood.
Chrysis (Tetrachrysis) derivata du Buysson, 1891. Rev. Ent. Caen 10: 38. ¢.
Chrysis (Tetrachrysis) decepta Rohwer, 1909. Psyche 16: 90. 2. N. syn. (R. M. Bohart).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 92-94 (host record). —Krombein, 1967.
Trap-nesting wasps and bees, p. 462 (life history).

irwini Bohart. Calif., Oreg. Host: Eumenidae in twigs.
Chrysis irwini Bohart, 1966. Biol. Soc. Wash., Proc. 79: 133. d, 2.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 98 (host record).
pattoni Aaron. Transcontinental in U. S., Sask., B. C. Host: Ancistrocerus spilogaster Cam.;
Leptochilus washo Prkr.; Microdynerus bakerianus (Cam.). Hosts nest in cavities in

twigs.
Chrysis Pattoni Aaron, 1885. Amer. Ent. Soc., Trans. 12: 235. °.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (host record). —Parker and
Bohart, 1968. Pan-Pacific Ent. 44: 2-3 (host records). —Parker, 1970. Pan-Pacifie Ent. 46:
242 (host record).
snowi Viereck. U.S. and Canada west of 100th meridian.
Chrysis (Tetrachrysis) snowi Viereck, 1906. Amer. Ent. Soc., Trans. 32: 195.
stenodyneri Krombein. Transcontinental in U. S., south to Panama. Host: Stenodynerus h.
histrionalis (Robt.), S. krombeini Boh., S. lineatifrons Boh.; Parancistrocerus histrio
(Lep.); Microdynerus bakerianus (Cam.).
Chrysis (Chrysis) stenodyneri Krombein, 1958. Amer. Ent. Soc., Trans. 84: 151, figs. 1, 2, 8.
ONG

Superfamily BETHYLOIDEA 1229

Biology: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 155-160 (life history.). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 471-472 (life history). —Parker, 1970. Pan-Pacific Ent.

46: 242 (host record).
SPECIES GROUP SMARAGDULA

Taxonomy: Bohart, 1962. Acta Hym. 1: 362-363, figs. 16, 21-25 (group characters).
arizonica Bohart. Tex., N. Mex., Ariz.; northern Mexico. Host: Euodynerus guerrero (Sauss.);

Parancistrocerus toltecus (Sauss.). Hosts nest in borings in wood.
Chrysis arizonica Bohart, 1962. Acta Hym. 1: 366, figs. 2, 10, 18, 19. 6, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 455-456 (life history).
bequaerti Bohart. D. C., Ga., Ark., Mo., Iowa, IIL, Ind., S. Dak. to Tex., N. Mex.
Chrysis bequaerti Bohart, 1962. Acta Hym. 1: 366, figs. 7, 9, 20, 21, 25. d, °.

clara Cresson. S. Dak., Colo., Wash.
Chrysis clara Cresson, 1865. Ent. Soe. Phila., Proc. 4: 313. 2.

Taxonomy: Bohart, 1962. Acta Hym. 1: 367.
inaequidens Dahlbom. Over most of U. S., N. Y. to Wash. and south into Mexico (Chihuahua,

Nuevo Leon, Baja California). Host: Euodynerus foraminatus apopkensis (Robt.), E.
foraminatus scutellaris (Sauss.), E. megaera (Lep.); Monobia quadridens (L.);
Pachodynerus erynnis (Lep.); Stenodynerus pulvinatus surrufus Krom.;
Parancistrocerus salcularis rufulus (Boh.). Undoubtedly parasitizes a number of other
eumenid wasps nesting in borings in wood or abandoned mud-dauber nests.

Chrysis inaequidens Dahlbom, 1854. Hym. Europaea, v. 2, p. 334. d.

Chrysis Texana Gribodo, 1879. Mus. Civ. Stor. Nat. Genova, Ann. 14: 829. d, ?.

Taxonomy: Bohart, 1962. Acta Hym. 1: 368, fig. 14.

Biology: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 161-163 (life history). —Moore and
Parker, 1962. Pan-Pacific Ent. 38: 14 (host record). —Krombein, 1967. Trap-nesting wasps
and bees, pp. 463-465 (life history).

intricata Brulle. Tex. (Brownsville); Mexico south to Brazil.
Chrysis intricata Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 25. °.
Chrysis smidti Dahlbom, 1845. Hym. Europaea v. 2, p. 317. °.
Chrysis cognata Gribodo, 1879. Mus. Civ. Stor. Nat. Genova, Ann. 14: 328. °.
Chrysis proxima Cameron, 1888. Biol. Cent.-Amer., Hym. v. 1, p. 465. °.
Chrysis (Hexachrysis) schulthessi Mocsary, 1889. Monog. Chrysid., p. 572. d.
Chrysis (Hexachrysis) aenescens Mocsary, 1889. Monog. Chrysid., p. 577. °.
Chrysis aperta du Buysson, 1898 (1897). Ann. Soc. Ent., France 66: 559. d.

Taxonomy: Bohart, 1962. Acta Hym. 1: 368, fig. 16.

oraria Bohart. Ariz., N. Mex., Tex.
Chrysis oraria Bohart, 1962. Acta Hym. 1: 370. d, °.
praestans du Buysson. Ky., Colo. to Tex., west to Wash. and Calif., B. C., Alta., Man.; West
Indies (Grenada).
Chrysis praestans du Buysson, 1898. Rev. Ent. Caen 17: 144. ?.
Taxonomy: Bohart, 1962. Acta Hym. 1: 370, fig. 8.
serrata Taylor. U. S. west of 100th meridian, B. C.
Chrysis (Hexachrysis) serrata Taylor, 1924. Calif. Acad. Sci., Proc. (4)13: 329. d, °.
Taxonomy: Bohart, 1962. Acta. Hym. 1: 371, fig. 12.
smaragdula Fabricius. N. J. to Fla., Ky., Mo., La., N. Mex., Ariz., Utah. Host: Monobia
quadridens Fabr. Hosts nest in borings in wood.

Chrysis smaragdula Fabricius, 1775. Systema Ent., p. 357.
Chrysis pensylvanica Lepeletier, 1846. Hist. Nat. Ins. Hym., v. 4, p. 24. °.

Taxonomy: Bohart, 1962. Acta Hym. 1: 371, fig. 13.

Biology: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 163 (life history). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 469-471 (life history).

1230 Hymenoptera in America North of Mexico

wasbaueri Bohart. Generally distributed over U. S., Mass. to Wash., south to Mexico
(Chihuahua, Nuevo Leon, Durango).
Chrysis wasbaueri Bohart, 1962. Acta Hym. 1: 372, figs. 11, 22, 23, 24. 6, @.

SPECIES GROUP LAUTA

Taxonomy: Bohart, 1964. Biol. Soc. Wash., Proc. 77: 223-228, figs. 3, 6, 17, 20, 21, 24, 25 (group
characters and descriptions of all species except tripartita Aar.).
coloradica Bohart. Oreg., Calif., Nev., Idaho, Wyo., Colo., Utah, Ariz. Host: Anthidiwm
collectum Huard, in ground nests.
Chrysis pulcherrima Cresson, 1865. Ent. Soc. Phila., Proc. 4: 311. ¢. Preoce.
Chrysis coloradica Bohart, 1964. Biol. Soc. Wash., Proc. 77: 224. N. name.

Biology: Bohart, 1964. Biol. Soc. Wash., Proc. 77: 225 (host record).
florissanticola Rohwer. Alaska, Yukon, Alta., Wash., Calif., Utah, Wyo., Colo. Host:
Anthidium banningense Ckll.
Chrysis (Tetrachrysis) florissanticola Rohwer, 1909. Psyche 16: 88. ¢.
Chrysis florissantensis (!) Bohart, 1962. Acta Hym. 1: 363.

Biology: Grigarick and Stange, 1968. Calif. Insect Survey, Bul. 9: 15 (host record).
lauta Cresson. N. C., Ga., Miss., Tex., Kans., Nebr., Colo., N. Mex., Ariz.; Mexico (Morelos,
Jalisco, San Luis Potosi, Oaxaca.) Host: Anthidium porterae Ckll.
Chrysis lauta Cresson, 1865. Ent. Soc. Phila., Proc. 4: 310 9.
Chrysis prasinus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 310. 2. Preoce.
Chrysis chlorophana Mocsary, 1887. Termes. Fuzetek 11: 16. N. name.
Chrysis (Tetrachrysis) clypeata Mocsary, 1889. Monog. Chrysid., p. 393. ¢.
Chrysis (Tetrachrysis) falsifica du Buysson, 1891. Rev. Ent. Caen 10: 38. N. name.

Taxonomy: Bohart, 1964. Biol. Soc. Wash., Proc. 77: 224 (synonymy).

Biology: Custer, 1928. Ent. News 39: 123-134.
tripartita Aaron. Ga., Mo., Kans., Colo., Tex. west to Calif.; northern Mexico. Host: Anthidium
collectum Huard; A. maculosum Cr. Hosts nest in ground or in twig cavities.
Chrysis tripartita Aaron, 1885. Amer. Ent. Soc., Trans. 12: 238. °.
Chrysis (Tetrachrysis) indeterminabilis Bischoff, 1910. Berlin Zool. Mus., Mitt. 4: 476. 9°.
N. syn. (R. M. Bohart).

Biology: Grigarick and Stange, 1968. Calif. Insect Survey, Bul. 9:19 (host record). —Horning,

1971. Ent. Soe. Wash., Proc. 73: 43 (host record).
tularensis Bohart. Calif. (Tulare Co.).

Chrysis tularensis Bohart, 1962. Acta Hym. 1: 373, figs. 4, 6. d, °.
vagabunda Bohart. Calif., Oreg., Wash., Idaho, Nev., Utah, Ariz. Host: Anthidiuwm collectum

Huard.

Chrysis vagabunda Bohart, 1964. Biol. Soc. Wash., Proc. 77: 225, figs. 24, 25. 3, 9.
xerophila Bohart. Calif.

Chrysis xerophila Bohart, 1964. Biol. Soc. Wash., Proc. 77: 226, figs. 3, 20, 21. d, @.

SPECIES GROUP NISSERI

nisseri Dahlbom. N. J. to Fla., Mo., Ark., Tex.; Mexico, Central and South America.

Chrysis nisseri Dahlbom, 1845. Dispos. Method. Spec. Hym., pt. 2, p. 14. °.

Chrysis (Tetrachrysis) propinqua Moesary, 1889. Monog. Chrysid., p. 348. ¢, 2. N. syn. (R.
M. Bohart).

Chrysis (Tetrachrysis) laminifera Bischoff, 1910. Berlin Zool. Mus., Mitt. 4: 460. 2. N. syn.
(R. M. Bohart).

Chrysis (Tetrachrysis) chiriquensis Bischoff, 1910. Berlin Zool. Mus., Mitt. 4: 461. 9. N.
syn. (R. M. Bohart).

Chrysis (Tetrachrysis) palifera Bischoff, 1910. Berlin Zool. Mus., Mitt. 4: 461. ¢. N. syn.
(R. M. Bohart).

Taxonomy: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 149, fig. 3.

Superfamily BETHYLOIDEA 1231

SPECIES GROUP FUSCIPENNIS

fuscipennis Brulle. D. C., Md., Va., Ky., Calif.; Hawaii; Palaearctic, Oriental, Australian.

Probably adventive from Hawaii. Host: Sceliphron caementarium (Dru.) in U. S.;
Eumenes conica (F.) in India, Sceliphron spp. in Japan. Hosts are resting mature larvae
of mud-daubers.

Chrysis fuscipennis Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 38. 9.

Chrysis erratica Abeille de Perrin, 1887. Jn Buysson, Rev. d’Ent. 6: 189. d, 9.

Chrysis pulchellus Cameron, 1888. Manchester Lit. and Phil. Soc., Mem. and Proc. 26: 126.
Preoce.

Taxonomy: Krombein, 1956. Ent. Soc. Wash., Proc. 58: 275.
Biology: Stage, 1960. Pan-Pacifie Ent. 36: 191-195, 1 pl. (life history).
SPECIES GROUP VENUSTA
Taxonomy: Bohart, 1964. Biol. Soc. Wash., Proc. 77: 233-235, figs. 8, 9, 10, 12, 13 (group
characters and descriptions of all species except venusta Cr.).
astralia Bohart. Oreg., Calif., Nev., Utah, Ariz., N. Mex.; Mexico (Baja California, Sonora,

Zacatecas, Morelos, Jalisco).
Chrysis astralia Bohart, 1964. Biol. Soc. Wash., Proc. 77: 233, figs. 8, 13. d, 2.

venusta Cresson. Kans., Colo., Tex., N. Mex., Calif.
Chrysis venusta Cresson, 1865, Ent. Soc. Phila., Proc. 4: 311. 6.
Chrysis (Tetrachrysis) frey-gessneri Gribodo, 1879. Mus. Civ. Stor. Nat. Genova, Ann. 14:
333. 2. N. syn. (R. M. Bohart).

Taxonomy: Bohart, 1964. Biol. Soc. Wash., Proc. 77: 232, fig. 10 (male genitalia).

venustella Bohart. Wash., Oreg., Calif., Nev., Idaho, Wyo.
Chrysis venustella Bohart, 1964. Biol. Soc. Wash., Proc. 77: 235, figs. 9, 12. d, @.

SPECIES GROUP PROPRIA
Taxonomy: Bohart, 1964. Biol. Soc. Wash., Proc. 77: 228-233, figs. 1, 2, 5, 7, 11, 17 (group
characters and descriptions of new species only).
aridula Bohart. Ariz., southern Calif.
Chrysis aridula Bohart, 1962. Acta Hym. 1: 364, figs. 1,5, 17. d, 9.
crotema Bohart. N. Mex., Ariz.; Mexico (Chihuahua, Zacatecas).
Chrysis crotema Bohart, 1964. Biol. Soc. Wash., Proc. 77: 228, figs. 5, 18, 19. d, °.

prolata Bohart. Kans., Iowa, IIl., Wis.
Chrysis prolata Bohart, 1964. Biol. Soc. Wash., Proc. 77: 229. 2.

propria Aaron. Transcontinental in U. S., Alta., B. C.; Mexico (Chihuahua, Durango).
Chrysis propria Aaron, 1885. Amer. Ent. Soc., Trans. 12: 238. d, 2.
Chrysis (Tetrachrysis) kahli Viereck, 1906. Amer. Ent. Soc., Trans. 32: 194. ¢.
Chrysis (Tetrachrysis) pattonella Viereck, 1906. Amer. Ent. Soc., Trans. 32: 194. d.

Taxonomy: Krombein, 1957. Ent. News 68: 191.
rivalis Bohart. Calif., Oreg., Wash., Idaho, Mont., Wyo., Utah, Colo., Nev.
Chrysis rivalis Bohart, 1964. Biol. Soc. Wash., Proc. 77: 229, figs. 4, 22, 28. d, °.
submontana Rohwer. N. Mex., Colo., Utah, Wash., Oreg., Calif.
Chrysis (Tetrachrysis) submontana Rohwer, 1909. Psyche 16: 91.

tensa Bohart. Calif., Utah, Ariz., N. Mex., Tex., Kans., Nebr.; Mexico (Sonora, Chihuahua).
Chrysis tensa Bohart, 1964. Biol. Soe. Wash., Proc. 77: 230, figs. 2, 14, 15. d, 9.

vibex Bohart. Calif., Ariz., N. Mex., Tex., Okla., Kans.; Mexico (Chihuahua, Nuevo Leon,
Jalisco).
Chrysis vibex Bohart 1964. Biol. Soc. Wash., Proc. 77: 231, figs. 1, 7, 11, 16. d, °.
SPECIES GROUP COERULANS
Taxonomy: Moore, 1966. Ent. Soc. Amer., Ann. 59: 1125-1131, 9 figs. (group characters,
synonymy and description of both sexes of included species).

canadensis du Buysson. Transcontinental in U. S. and Canada; Mexico (Guerrero).
Chrysis (Tetrachrysis) canadensis du Buysson, 1891. Rev. Ent. Caen 10: 37. 2.

1232 Hymenoptera in America North of Mexico

Chrysis (Tetrachrysis) altivolans Moesary, 1914. Mus. Nat. Hungarici, Ann. 12: 53. 6. N.
syn. (R. M. Bohart).
Chrysis (Tetrachrysis) coerulans var. nanula Rohwer, 1909. Psyche 16: 88. 2. N. syn (R.
M. Bohart).
cembricola Krombein. Transcontinental in U. S. and Canada. Host: Symmorphus canadensis
(Sauss.). Hosts nest in borings in wood.
Chrysis (Chrysis) cembricola Krombein, 1958. Ent. Soe. Wash., Proc. 60: 53, fig. 1. 2, d.

Biology: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 57-58 (life history). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 456-457 (life history).
coerulans Fabricius. Widely distributed in southern Canada and U.S. Host: Ancistrocerus a.
antilope (Panz.); A. c. catskill (Sauss.); A. c. albophaleratus (Sauss.); A. spilogaster
Cam.; A. a. adiabatus (Sauss.); A. a. cytainus (Cam.); A. tuberculocephalus sutterianus
(Sauss.); Huodynerus foraminatus apopkensis (Robt.); E. f. foraminatus (Sauss.); E. f.
scutellaris (Sauss.); E. leucomelas (Sauss.); E. megaera (Lep.); Parancistrocerus
acarophorus (Boh.); P. saleularis rufulus (Boh.); Symmorphus albomarginatus (Sauss.);
S. c. cristatus (Sauss.); Hoplitis anthocopoides (Schenk). Parasite: Melittobia chalybii
Ashm. Hosts nest in cavities in twigs, borings in wood, and in old mud-dauber nests.
Chrysis coerulans Fabricius, 1805. Systema Piezatorum, p. 172. 9.
Chrysis Servillei Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 37. N. syn. (R. M. Bohart).
Chrysis bella Cresson, 1865. Ent. Soc. Phila., Proc. 4: 312. 2.
Chrysis Nortoni Aaron, 1885. Amer. Ent. Soc., Trans. 12: 237. d, 2. N. syn. (R. M.
Bohart).
Chrysis (Tetrachrysis) conserta du Buysson, 1891. Rev. Ent. Caen 10: 37. °.
Chrysis (Tetrachrysis) praticola Mocsary, 1914. Mus. Nat. Hungarici, Ann. 12: 50. d, ?.N.
syn. (R. M. Bohart).
Chrysis (Tetrachrysis) sejuncta Mocsary, 1914. Mus. Nat.. Hungarici, Ann. 12: 51. 2. N.
syn. (R. M. Bohart).

Biology: Walsh, 1869. Amer. Ent. 1: 135 (host record, erroneous). —Ashmead, 1894. Psyche 7:
79 (host record, erroneous). —Parker, 1962. Pan-Pacific Ent. 38: 140 (host record).
—Medler, 1964. Ent. News 75: 26-27 (host record). —Medler, 1964. Ent. Soc. Wash., Proc.
66: 209-215 (life history). —Medler, 1964. Ent. Soc. Amer., Ann. 57: 59 (host record).
—Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 93 (host records). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 457-462 (life history). —Eickwort, 1973. Search, Agr. 3:
24 (host). —Evans, 1973. Great Basin Nat. 33: 152 (host).

inflata Aaron. Tex., Colo. and N. Mex. west to Calif. and Oreg., Idaho, B. C.; Mexico. Host:

Ancistrocerus durangoensis Cam.; A. lineativentris Cam.; A. tuberculocephalus
sutterianus (Sauss.); A. t. tuberculocephalus (Sauss.); Euodynerus guerrero (Sauss.).
Hosts nest in cavities in twigs and in borings in wood.

Chrysis inflata Aaron, 1885. Amer. Ent. Soc., Trans. 12: 237. d, °.

Chrysis Dugesi du Buysson, 1898 (1897). Soc. Ent. France, Ann. 66: 582, pl. 18, fig. 7. 2. N.
syn. (R. M. Bohart).

Chrysis (Tetrachrysis) nokomis Rohwer, 1909. Psyche 16: 89. d, 2. N. syn. (R. M. Bohart).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (host record). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 465-466 (life history).
nitidula Fabricius. Transcontinental in U. S. and Canada. Host: Ancistrocerus a. antilope

(Panz.); A. c. catskill (Sauss.); A. c. albophaleratus (Sauss.); A. a. adiabatus (Sauss.);
Euodynerus f. foraminatus (Sauss.); E. schwarzi (Krom.); Symmorphus c. cristatus
(Sauss.). Hosts nest in cavities in stems, borings in wood, and abandoned mud cells of
mining bees.

Chrysis nitidula Fabricius, 1775. Systema Ent., p. 359.

Chrysis cyanea Villers, 1789. Caroli Linnaei Ent., v. 3, p. 257.

Chrysis (Tetrachrysis) cessata du Buysson, 1891. Rev. Ent. Caen 10: 36.

Chrysis (Tetrachrysis) chalcopyga Moesary, 1914. Mus. Nat. Hungarici, Ann. 12: 48. ?.

Biology: Harrington, 1896. Canad. Ent. 28: 79 (host record). —Hobbs, Nummi and Virostek,
1961. Canad. Ent. 93: 147 (host record). —Medler, 1964. Ent. Soc. Amer., Ann. 57: 59 (host

Superfamily BETHYLOIDEA 1233

record). —Medler, 1964. Ent. Soc. Wash., Proc. 66: 209-215 (life history). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 466-468 (life history).
parkeri Moore. Que., Alta., B. C., Mont., Idaho, Utah, Nev., Oreg., Calif. Host: Ewodynerus
foraminatus scutellaris (Sauss.); Leptochilus rufinodus (Cr.); Microdynerus bakerianus
(Cam.); Parancistrocerus acarigaster (Boh.). Hosts nest in cavities in twigs and in
abandoned mud-dauber nests.
Chrysis parkeri Moore, 1966. Ent. Soc. Amer., Ann. 59: 1130, fig. 8. d, 2.
Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 93 (host records). —Parker, 1970.
Pan-Pacifie Ent. 46: 242 (host record).

pellucidula Aaron. Transcontinental in U.S. and southern Canada. Host: Trypargilum
collinum rubrocinctum (Pack.), T. t. tridentatum (Pack.). Hosts nest in cavities in twigs

and in borings in wood.
Chrysis pellucidula Aaron, 1885. Amer. Ent. Soc., Trans. 12: 235. 9.
Biology: Hicks, 1934. Univ. Colo. Studies 21: 267 (host record). —Parker and Bohart, 1966.
Pan-Pacific Ent. 42: 95 (host record). —Krombein, 1967. Trap-nesting wasps and bees, p.
473 (life history). —Medler, 1967. Amer. Midland Nat. 78: 351 (host record).
remissa Mocsary. Pa., Va., Tex., N. Mex., Ariz.; Mexico (Zacatecas, Veracruz).
Chrysis (Tetrachrysis) remissa Mocsary, 1914. Mus. Nat. Hungarici, Ann. 12: 52. °.

Genus CHRYSIS Subgenus PRAESTOCHRYSIS Linsenmaier

Chrysis subg. Praestochrysis Linsenmaier, 1959. Schweiz. Ent. Ges., Mitt. 32: 164.
Type-species: Chrysis shanghaiensis Smith. Orig. desig.
This subgenus is not a member of the Nearctic fauna.
shanghaiensis Smith. Liberated in Mass. in 1917-18 but not recovered since 1919; Japan,
China, Thailand, India, East Indies. Introduced from China. Host: Diapausing larva,
Cnidocampa flavescens (Wlkr.), the oriental moth.

Chrysis Shanghaiensis Smith, 1874. Ent. Soc. London, Trans., p. 460. 2.
Chrysis (Pentachrysis) himalayensis Moesary, 1888. In Radoszkowski, Hor. Soc. Ent.

Ross. 23: 31, fig. 63. 3.
Chrysis (Pentachrysis) crassiscuta Mocsary, 1889. Monog. Chrysid., p. 524. °.
Biology: Parker, 1936. Jour. Agr. Res. 52: 449-458 (life history, behavior).

Morphology: Parker, 1936. Jour. Agr. Res. 52: 452-455, 5 figs. (egg, larva).
Genus CERATOCHRYSIS Cooper

Chrysis subg. Ceratochrysis Cooper, 1952. Amer. Ent. Soc., Trans. 78: 138.
Type-species: Chrysis (Ceratochrysis) enhuycki Cooper. Orig. desig.

Biology and host relationships are known for only a few of the included species. These meager
data indicate that species of this genus parasitize eumenid and sphecoid wasps and megachilid
bees nesting in cavities in twigs.

Taxonomy: Cooper, 1952. Amer. Ent. Soc., Trans. 78: 137-148. —Bohart, 1966. Kans. Ent.

Soe., Jour. 39: 112-113 (redefined and raised to generic rank).

alveata Bohart. Calif., Ariz.
Ceratochrysis alveata Bohart, 1966. Kans. Ent. Soc., Jour. 39: 118, figs. 7, 11, 14. d, 2.

antyga Bohart. Calif., Nev., Ariz., N. Mex. Host: Trypargilum t. tridentatum (Pack.);
Pisonopsis birkmanni Roh.
Ceratochrysis antyga Bohart, 1966. Kans. Ent. Soc., Jour. 39: 115, figs. 6, 13. d, 2.
Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 94-95 (host records).

bitumida Bohart. N. Mex. (Hidalgo Co.).
Ceratochrysis bitumida Bohart, 1966. Kans. Ent. Soc., Jour. 39: 116, fig. 1. 2.

cavicantha Bohart. Calif., Ariz.
Ceratochrysis cavicantha Bohart, 1966. Kans. Ent. Soc., Jour. 39: 116, fig. 8. d, °.

1234 Hymenoptera in America North of Mexico

crossata Bohart. Oreg., Calif., Nev., Utah, Ariz., N. Mex., Tex.; Mexico (Hidalgo, San Luis

Potosi, Puebla, Mexico).
Ceratochrysis crossata Bohart, 1966. Kans. Ent. Soc., Jour. 39: 117, figs. 2, 9, 12. d, °.

cyanosoma Mocsary. Wash., Oreg., Calif., Nev., Utah, Wyo.
Chrysis (Holochrysis) cyanosoma Moesary, 1914. Mus. Nat. Hungarici, Ann. 12: 17. 9.

declinis Bohart. N. B., N. S., Ont., Maine, N. H., Conn., N. Y., Mich., Wis., Minn., Sask., Colo.,

Alta.
Ceratochrysis declinis Bohart, 1966. Kans. Ent. Soc., Jour. 39: 119, figs. 10, 15. d, @.

enhuycki (Cooper). N. Y. to Fla., Mo., Tex., Utah, Ariz. Host: Leptochilus ornatus Sauss.
Reared from sumac twigs.
Chrysis (Ceratochrysis) enhuycki Cooper, 1952. Amer. Ent. Soc., Trans. 78: 140, figs. 3, 5,
6, 8, 9. 2, 6.
Biology: Krombein, 1959. Ent. News 70: 17-23 (life history).
faceta (Aaron). Colo., Tex., N. Mex., Calif.
Chrysis faceta Aaron, 1885. Amer. Ent. Soc., Trans. 12: 229. ¢.
Taxonomy: Cooper, 1952. Amer. Ent. Soc., Trans. 78: 144-147, figs. 1, 4, 10 (redescription
male). —Bohart, 1966. Kans. Ent. Soc., Jour. 39: 119.
kansensis (Viereck). Generally distributed in southern Canada and U. S. west to Alta., Wyo.,
Colo. and Ariz.; Mexico (Jalisco).
Chrysis (Gonochrysis) kansensis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 193.
Taxonomy: Bohart, 1966. Kans. Ent. Soc., Jour. 39: 120, fig. 19.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14, pp. 42-48, pl. 18, figs.
J-N (male genitalia).
longiceps Bohart. Nev., Ariz.; Mexico (Guanajuato).
Ceratochrysis longiceps Bohart, 1966. Kans. Ent. Soc., Jour. 39: 120, fig. 4. d, 2.
nearctica (Mocsary). Tex., Ariz., Calif.; Mexico (Sonora).
Spintharis nearctica Mocsary, 1911. Mus. Nat. Hungarici, Ann. 9: 462. 9.
Taxonomy: Bohart, 1966. Kans. Ent. Soc., Jour. 39: 115, 121.
perpulchra (Cresson). U. S. and Canada west of 100th meridian, south to Costa Rica. Host:
Ammophila aberti Hald.
Chrysis perpulchra Cresson, 1865. Ent. Soc. Phila., Proc. 4: 308. 9.
Taxonomy: Bohart, 1966. Kans. Ent. Soc., Jour. 39: 122.

Biology: Hicks, 1932. Canad. Ent. 64: 150-151 (host record).
quadrituberculata (Cameron). Idaho, Tex., Ariz., south to El Salvador, Honduras.
Chrysis quadrituberculata Cameron, 1888. Biol. Cent.-Amer., Hym., v. 1, p. 461, pl. 20, figs.
Went
Taxonomy: Cooper, 1952. Amer. Ent. Soc., Trans. 78: 142-144, figs. 2, 7, 11. d, 2. —Bohart,
1966. Kans. Ent. Soc., Jour. 39: 119.
thysana Bohart. Oreg., Calif., Nev. Host: Leptochilus periallis Prkr.
Ceratochrysis thysana Bohart, 1966. Kans. Ent. Soc., Jour. 39: 120, figs. 5, 16. ¢, 9.
Biology: Parker and Bohart, 1968. Pan-Pacific Ent. 44: 2 (host record).
trachypleura Bohart. B. C. and Mont. south to Calif., Ariz. and Colo. Host: Ammophila aberti
Hald.
Ceratochrysis trachypleura Bohart, 1966. Kans. Ent. Soc., Jour. 39: 121, figs. 3, 17. d, 9.
Biology: Hicks, 1932. Canad. Ent. 64: 150 (host; parasite misdet. as perpulchra Cr.).
tuberella Bohart. Oreg., Calif., Nev., Utah, Colo. Host: Leptochilus rufinodus (Cr.).
Ceratochrysis tuberella Bohart, 1966. Kans. Ent. Soc., Jour. 39: 122, fig. 18. 6, 9.
Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (host record).

unita (Mocsary). Tex., Colo., Calif.; Mexico.
Chrysis (Olochrysis) unita Mocsary, 1889. Monog. Chrysid., p. 211. 2.

Superfamily BETHYLOIDEA 1235

Genus EUCHROEUS Latreille
Taxonomy: Linsenmaier, 1968. Schweiz.Ent. Ges., Mitt. 41: 38-47.
Genus EUCHROEUS Subgenus EUCHROEUS Latreille

Euchroeus Latreille, 1809. Gen. Crust. Ins., v. 4, p. 578.
Type-species: Chrysis purpuratus Fabricius. Monotypic.

The typical subgenus does not occur in North America.

Genus EUCHROEUS Subgenus PSEUDOSPINOLIA Linsenmaier

Euchroeus subg. Pseudospinolia Linsenmaier, 1951. Schweiz. Ent. Ges., Mitt. 24: 27.
Type-species: Chrysis uniformis Dahlbom. Orig. desig.
neglectus (Shuckard). Alta., Man., Sask., Minn., N. Dak., Nebr., Mont., Colo., N. Mex.; Europe,

Asia. Host: Odynerus spinipes (L.), O. reniformis (Gmel.) in Europe; putative host in
North America is O. dilectus Sauss.

Chrysis neglecta Shuckard, 1837. Ent. Mag. 4: 169.

Olochrysis semicuprea Viereck, 1903. Amer. Ent. Soc., Trans. 29: 70. 2. N. syn. (R. M.
Bohart).

Genus SPINTHAROSOMA Zimmermann

Spintharosoma Zimmermann, 1959. Deut. Ent. Ztschr. 6: 32.
Type-species: Spintharosoma chrysonota Dahlbom. Orig. desig.

Euchroeus subg. Hyalichroeus Linsenmaier, 1959. Schweiz. Ent. Ges., Mitt. 32: 70.
Type-species: Spintharosoma chrysonota Dahlbom. Orig. desig.

Dahlbom misidentified Spintharis Klug. The names listed above were proposed for Spintharis
sensu Dahlbom.

mesillae (Cockerell). Southern Tex. to southern Calif., Utah; Mexico (Sonora). Host:
Ammophila breviceps Sm.; A. femurrubra Fox. The host species nest in soil.
Chrysis (Dichrysis) mesillae Cockerell, 1894. Ent. News 5: 125. °.
Chrysis (Dichrysis) bigeloviae Cockerell, 1897. Ann. and Mag. Nat. Hist. 19: 401. d.
Spintharis annulipes Mocsary, 1911. Mus. Nat. Hungarici, Ann. 9: 462. d.

trochilus (du Buysson). U.S. west of 100th meridian; Mexico (Sonora).
Chrysis (Spintharis) trochilus du Buysson, 1891. Rev. Ent. Caen 10: 32. 9.

Genus CHRYSURA Dahlbom

Chrysura Dahlbom, 1845. Dispos. Method. Spec. Hym., pt. 2, p. 6.
Type-species: Chrysis austriaca Fabricius. Desig. by Bodenstein, 1939.
Chrysogona Foerster, 1853. Naturh. Ver. Rheinlande, Verh. 10: 327.
Type-species: Chrysogona gracillima Foerster. Monotypic.
Chrysis subg. Olochrysis Lichtenstein, 1876. Petites Nouv. Ent., v. 2, p. 27.
Type-species: Chrysis aerata Dahlbom. Desig. by Ashmead, 1902 (=Chrysis
trimaculata Foerster).
Chrysis subg. Monochrysis Lichtenstein, 1876. Petites Nouv. Ent., v. 2, p. 27.
Type-species: Chrysis hybrida Lepeletier. Desig. by Ashmead, 1902.
Holochrysis Rye, 1878. Zool. Record for 1876, v. 13, Ins., p. 134. Emend.
Chrysoura Dalla Torre, 1892. Cat. Hym., v. 6, p. 40. Emend.
Chrysis subg. Arctochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,
Abhandl. u. Ber. 23: 72.
Type-species: Chrysis austriaca Fabricius. Orig. desig.
Chrysis subg. Taeniochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,
Abhandl. u. Ber. 23: 72.
Type-species: Chrysis dichroa Dahlbom. Orig. desig.
Chrysis subg. Selenochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,
Abhandl. u. Ber. 23: 72.
Type-species: Chrysis candens Germar. Orig. desig.

1236 Hymenoptera in America North of Mexico

Chrysis subg. Ischnochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,

Abhandl. u. Ber. 23: 73.
Type-species: Chrysis gracillima Foerster. Orig. desig.

The known life histories indicate that the female Chryswra deposits her egg in a host cell at
the time that it is being provisioned. The parasite egg hatches after that of the host and the
newly hatched Chrysura larva attaches to the host body, imbibes a small amount of body fluids,
but does not molt until the host larva spins its cocoon and enters the resting stage. The Chrysu-
ra larva then molts, devours the host larva, and spins its cocoon within that of the host. Develop-
ment to the adult stage and emergence from the nest is synchronized with the host. In vernal
univoltine species the host and parasite transform to adults late in the summer and overwinter
in the cocoons.

In addition to the hosts listed below for some of the species, Thorp (1968. Kans. Ent. Soc.,
Jour. 41: 324-331) published life history notes on a species near pacifica (Say) which parasitized
Proteriades bunocephala (Mich.) in California.

boharti Horning. B. C., Wash., Idaho, Calif., Wyo.
Chrysura boharti Horning, 1971. Pan-Pacific Ent. 47: 29, figs. 2, 5, 8, 9, 11, 18. d, @.
cobaltina (Aaron). Mass., Mich., Ind., Ill., Mont., Wyo., Utah, Calif., Wash., B. C.
Chrysis cobaltina Aaron, 1885. Amer. Ent. Soc., Trans. 12: 228. 9.
Chrysis lateri-dentata Aaron, 1885. Amer. Ent. Soc., Trans. 12: 228. 9.
crescentis Horning. U. Sonor. and Transit. Zones in central and southern California.
Chrysura crescentis Horning, 1971. Pan-Pacifie Ent. 47: 27, figs. 1, 3, 4, 6, 7, 10, 12. 6, 9.
densa (Cresson). Wash., Idaho, Oreg., Calif., Nev., Utah, Ariz., Wyo., Colo., Tex. Host:
Pseudomasaris edwardsii (Cr.); P. vespoides (Cr.); P. zonalis (Cr.). Hosts nest in mud
cells attached to rocks.
Chrysis densa Cresson, 1865. Ent. Soc. Phila., Proc. 4: 307. @.

Biology: Hicks, 1927. Canad. Ent. 59: 78 (host record). —Hicks, 1929. Canad. Ent. 61: 122

(host record).

inusitata Aaron. B. C., Oreg., Calif., Utah, Colo., N. Y., Va.
Chrysis inusitata Aaron, 1885. Amer. Ent. Soc., Trans. 12: 227. 3.
Chrysis optima Aaron, 1885. Amer. Ent. Soc., Trans. 12: 227. 2.
Chrysis (Holochrysis) interfata du Buysson, 1908. Rev. Ent. Caen 27: 208.

kyrae Krombein. Md. to Ga., Ind., Mo., Colo., Utah, Idaho, B. C., Wash., Oreg., Calif. Host:

Osmia l. lignaria Say in borings in wood.

Chrysura kyrae Krombein, 1963. Ent. News 74: 150. 6, °.

Biology: Krombein, 1963. Ent. News 74: 152 (life history). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 444-445, figs. 87, 88, 128-130 (life history).
martia (Patton). Lower Canada, Mich.
Chrysis martia Patton, 1879. Canad. Ent. 11: 67-68.
pacifica (Say). Transcontinental in U. S. and Canada. Host: Osmia kincaidvi Ckll., O. pumila
Cr., O. tanneri Sandh., O. georgica Cr., O. nigrifrons Cr., O. coloradensis Cr. Predator:
Philanthus pulcher D. T.
Chrysis pacifica Say, 1828. Contrib. Maclur. Lye. Phila. 1: 82.
Chrysis hilaris Dahlbom, 1854. Hym. Europaea, v. 2, p. 103.
Chrysis halictula Gribodo, 1874. Mus. Civ. Stor. Genova, Ann. 6: 359.
Chrysis resecta Gribodo, 1879. Mus. Civ. Stor. Nat. Genova., Ann. 14: 336. N. syn. (R. M.
Bohart).

Taxonomy: Krombein, 1963. Ent. News 74: 149-150. —Horning, 1971. Pan-Pacifie Ent. 47: 31.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 97 (host record). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 446-447, fig. 131 (life history). —Parker, 1975.
Pan-Pacifiec Ent. 51: 182 (host). —Hawkins, 1975. Kans. Ent. Soc., Jour. 48: 498 (host).

smaragdicolor (Walker) B. C. to Calif., Idaho, Wyo., Utah, N. Mex., Colo., N. Dak. Host:

Osmia longula Cr.
Chrysis smaragdicolor Walker, 1866. Jn Lord, Naturalist in Vancouver Isl. and B. C., v. 2,

p. 343.

Superfamily BETHYLOIDEA 1237
Taxonomy: Krombein, 1963. Ent. News 74: 149.

Biology: Leech, 1948. Ent. Soc. Brit. Columbia, Proc. 44: 27 (host record).
sonorensis (Cameron). Idaho, Oreg., Calif., Utah, Ariz., N. Mex.; Mexico (Baja California).
Host: Ashmeadiella aridula Ckll.; A. bigeloviae (Ckll.); A. bucconis denticulata (Cr.); A.
occipitalis Mich.; Hoplitis producta gracilis (Mich.); Anthocopa c. copelandica (Ckll.); A.

c. albomarginata (Ckll.); Osmia kincaidii Ckll. Hosts nest in cavities in twigs or in
borings in wood.

Chrysis Sonorensis Cameron, 1888. Biol. Cent.-Amer., Hym., v. 1, p. 461.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 96-97 (host records). —Parker, 1967.
Pan-Pacifie Ent. 43: 215 (host record). —Krombein, 1967. Trap-nesting wasps and bees, pp.
447-449 (life history). —Parker and Bohart, 1968. Pan-Pacifie Ent. 44: 4 (host record).
—Parker, 1975. Pan-Pacifie Ent. 51: 119 (host).

tota (Aaron). Md., Iowa, Colo., Idaho, Utah, Calif., B. C.

Chrysis integra Cresson, 1865. Ent. Soc. Phila., Proc. 4: 306-307. ¢. Preoce.
Chrysis tota Aaron, 1885. Amer. Ent. Soc., Trans. 12: 228. N. name.
Chrysis elongata Mocsary, 1887. Termes., Fuzetek 11: 15. N. name.

Genus NEOCHRYSIS Linsenmaier
Taxonomy: Bohart, 1966 (1963). Brooklyn Ent. Soc., Bul. 58: 139-144 (generic reclassification).

Genus NEOCHRYSIS Subgenus NEOCHRYSIS Linsenmaier

Pleurocera subg. Neochrysis Linsenmaier, 1959. Schweiz. Ent. Ges., Mitt. 32: 74.
Type-species: Chrysis punctatissima Spinola. Orig. desig.
montezuma (Cameron). Southern Ariz. south to Yucatan.
Chrysis montezuma Cameron, 1888. Biol. Cent.-Amer., Hym., v. 1, p. 463. °.

Genus NEOCHRYSIS Subgenus EXOCHRYSIS Bohart

Neochrysis subg. Exochrysis Bohart, 1966 (1963). Brooklyn Ent. Soc., Bul. 58: 141.
Type-species: Chrysis panamensis Cameron. Orig. desig.
panamensis (Cameron). Pa. to Fla., Ind., Okla., south to Panama. Ecology: Hosts nest in
borings in wood. Host: Podium rufipes (F.); P. luctuosum Sm.?
Chrysis Panamensis Cameron, 1888. Biol. Cent.-Amer., Hym., v. 1, p. 464, pl. 20, fig. 8. 2.
Chrysis (Tetrachrysis) alabamensis Moesary, 1914. Mus. Nat. Hungarici, Ann. 12: 49. 2.

Taxonomy: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 149, fig. 4. —Bohart and Menke,

1963. Univ. Calif. Pubs. Ent. 30: 108 (synonymy). —Bohart, 1966 (1963). Brooklyn Ent.
Soc., Bul. 58: 141. 2, 6d.

Biology: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 147-149 (life history). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 473-475 (life history).

Genus NEOCHRYSIS Subgenus IPSIURA Linsenmaier

Ipsiura Linsenmaier, 1959. Schweiz. Ent. Ges., Mitt. 32: 74.
Type-species: Chrysis marginalis Brulle. Orig. desig.
genbergi (Dahlbom). Fla.; Brazil, Argentina.
Chrysis genbergi Dahlbom, 1854. Hym. Europaea, v. 2, p. 319. 2.
neolateralis Bohart. D. C., Va., Ga., Ill., Kans., Ark. south to E] Salvador.
Neochrysis (Ipsiura) neolateralis Bohart, 1966 (1963) Brooklyn Ent. Soc., Bul. 58: 148. 3,
OF

pilifrons (Cameron). Southern Tex. to southern Mexico.
Chrysis pilifrons Cameron, 1888. Biol. Cent.-Amer., Hym., v. 1, p. 465. d.
Chrysis (Hexachrysis) stenops Mocsary, 1889. Monog. Chrysid., p. 571. °.

Genus TRICHRYSIS Lichtenstein
Chrysis subg. Trichrysis Lichtenstein, 1876. Petites Nouv. Ent., v. 2, p. 27.

1238 | Hymenoptera in America North of Mexico

Type-species: Sphex cyanea Linnaeus. Monotypice.
Chrysis subg. Alocochrysis Haupt, 1956. [Dresden] Mus. f. Tierkunde u. Voelkerk.,

Abhandl. u. Ber. 23: 73.
Type-species: Chrysis cyaneus Linnaeus. Monotypic.

The hosts of Trichrysis, so far as known, nest in cavities in twigs, borings in wood or aban-
doned nests of mud-daubers.

areolata (Mocsary). Md., Tenn., Tex.; Mexico (Guerrero).
Chrysogona areolata Mocsary, 1911. Mus. Nat. Hungarici, Ann. 9: 463. 9.
carinata (Say). Mass. to Fla., W. Va., Wis., Mo., Kans., Colo., Tex. Host: Trypargilum c.
collinum (Sm.); T. collinum rubrocinctum (Pack.); T. clavatum (Say); T. striatum
(Prov.).
Chrysis carinata Say, 1828. Contrib. Maclur. Lye. Phila. 1: 82.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 450-454, figs. 132, 133 (?), 134 (life
history).
deversor Bohart. B. C. to Calif., Idaho, Mont., Nev., Utah, Colo. Host: Trypargilum t.

tridentatum (Pack.).
Trichrysis deversor Bohart, 1966. Biol. Soc. Wash., Proc. 79: 131. d, 2.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 95 (host record).
doriae (Gribodo). Southern Canada and U. S. south to Panama. Host: Trypargilum collinum

rubrocinctum (Pack.); Trypoxylon backi Sandh.; T. bidentatum Fox; T. fastigium Fox;
T. frigidum Sm.; T. scullent Sandh.

Chrysis Doriae Gribodo, 1874. Mus. Civ. Stor. Nat. Genova, Ann. 6: 359.

Chrysis verticalis Patton, 1879. Canad. Ent. 11: 67. 6.

Chrysis striatellus Norton, 1879. Amer. Ent. Soc., Trans. 7: 241. 2.

Chrysis discreta Aaron, 1885. Amer. Ent. Soc., Trans. 12: 230. 9.

Biology: Thomas, 1962. Amer. Midland Nat. 67: 365-366 (life history). —Thomas, 1963. Mich.
Acad. Sci., Arts, Letters, Papers 48: 127-130 (life history). —Parker and Bohart, 1966.
Pan-Pacifie Ent. 42: 95 (host records). —Krombein, 1967. Trap-nesting wasps and bees, pp.
449-450 (life history). —Medler, 1967. Amer. Midland Nat. 78: 350 (host record).

mucronata (Brulle). ? Canada, Tex., Ariz., Calif.; south to Venezuela. Host: Trypargilum t.
tridentatum (Pack.); Euodynerus p. pratensis (Sauss.).
Chrysis mucronata Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 45. °.

Taxonomy: Bohart, 1966. Biol. Soc. Wash., Proc. 79: 132.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 454-455 (life history).
tridens (Lepeletier). U. S. east of Rocky Mts., south to Argentina. Host: Trypargilum politum

(Say).

Pyria tridens Lepeletier, 1825. In Olivier, Encycl. Meth., Dict. Ins., v. 10, p. 495. 2.

Chrysis truncata Guerin, 1842. Rev. Zool., p. 146. d, 2.

Chrysis Mexicana Guerin, 1842. Rev. Zool., p. 147.

Chrysis tridentata Dahlbom, 1845. Dispos. Method. Spec. Hym., pt. 2, p. 18.

Chrysis unicolor Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 42. 2. Preoce.

Chrysis olivieri Brulle, 1846. Hist. Nat. Ins. Hym., v. 4, p. 48. 2. Type locality erroneous.
N. syn. (R. M. Bohart).

Chrysis virens Cresson, 1865. Ent. Soc. Phila. Proc. 4: 309. 2. Preoce.

Chrysis Brullei Abeille de Perrin, 1879. Soe. Linn. Lyon, Ann. 26: 42. N. name for unicolor
Br.

Taxonomy: Bohart, 1966. Biol. Soc. Wash., Proc. 79: 132.
Biology: Johnson, 1974. Ent. Soc. Wash., Proc. 76: 448-449 (host).

Genus STILBUM Spinola

Stilbum Spinola, 1806. Insectorum Liguriae, v. 1, p. 9.
Type-species: Stilbwm calens Spinola. Desig. by Latreille, 1810 (=Chrysis cyanuwra
Foerster).

Superfamily BETHYLOIDEA 1239

This genus is not an established member of the Nearctic fauna. A specimen from Ontario in
the Provancher collection is either mislabeled or is a stray which did not become established.

cyanurum var. amethystinum (Fabricius). One specimen from Ont., but not established;
Mediterranean area, Asia, East Indies.
Chrysis amethystina Fabricius, 1775. Systema Ent., p. 359.
Stilbum splendidulum Westwood, 1842. In Donovan, Epitome Nat. Hist. Ins. India, p. 88.

SUBFAMILY PARNOPINAE

Genus PARNOPES Latreille

Parnopes Latreille, 1796. Precis. Caract. Gen. Ins., p. 126. No species —Latreille, 1802.
Hist. Nat. Crust. Ins., v. 3, p. 317.
Type-species: Chrysis grandior Pallas. First included species (~Chrysis carnea
Fabricius).

Species of Parnopes are parasitic on ground-nesting sphecoid wasps belonging to the
Bembicini. The egg of Parnopes is deposited during provisioning of the nest by the host wasp,
but the Parnopes larva does not develop until the host larva has spun its cocoon.

Revision: Viereck, 1904. Amer. Ent. Soc., Trans. 30: 245-250. —Telford, 1964. Univ. Calif.
Pubs. Ent. 36: 1-42, 5 pls., 1 text fig.
borregoensis Telford. Southern Calif.; Mexico (Baja California).
Parnopes borregoensis Telford, 1964. Univ. Calif. Pubs. Ent. 36: 9, pl. 1, fig. 2. 3, @.
chrysoprasinus Smith. N. Y. south to Fla., west to Wis. and Tex. Host: Bembix nubilipennis
Cr.; B. americana spinolae Lep.?, Steniolia obliqua (Cr.)?.
Parnopes chrysoprasinus Smith, 1874. Ent. Soc. London, Trans. p. 454, n. 8. d.
Parnopes aglaspidula Melander and Brues, 1902. Biol. Bul. 3: 38. °.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 307-308, 314-315
(life history).
concinnus Viereck. Calif. to western Tex., Nev.; Mexico (Baja California, Jalisco). Host:
Bembix multipicta Sm.?; Glenostictia scitula (Fox)?; Steniolia duplicata Prov.?
Parnopes concinna Viereck, 1904. Amer. Ent. Soc., Trans. 30: 248. °.
Parnopes arizonensis Viereck, 1904. Amer. Ent. Soc., Trans. 30: 249. 6, 9.
Parnopes digueti du Buysson, 1904. Rev. d’Ent. Caen 23: 274. ¢.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 205, 344.
edwardsii (Cresson). Alaska to California, east to the Rocky Mts.; Mexico (Baja California).

Host: Bembix amoena Handl.?; B. americana comata Prkr.; B. americana spinolae
Lep.?; B. pruinosa Fox.?; Steniolia obliqua (Cr.).

Euchroeus Edwardsii Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. iv. 2.

Parnopes boutheryi Brethes, 1902. Mus. Nac. Buenos Aires, An. 8 (8): 287. 2, d.

Parnopes hageni Viereck, 1904. Amer. Ent. Soc., Trans. 30: 246. d, @.

Parnopes henshawi Viereck, 1904. Amer. Ent. Soc., Trans. 30: 247. 2.

Biology: Bohart and MacSwain, 1940. Pan-Pacific Ent. 16: 92-93. —Evans and Gillaspy, 1964.
Amer. Midland Nat. 72: 277, fig. 14. —Evans, 1966. Compar. ethology and evolution of
sand wasps, pp. 219, 288, 315, 349.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14, pp. 42-43, pl. 13, figs. P,
R (male genitalia).

excurvatus Viereck. B. C., Calif., Nev., Ariz.
Parnopes excurvata Viereck, 1904. Amer. Ent. Soc., Trans. 30: 250. “2” = ¢.

festivus Cockerell. Colo., Ariz. to western Tex.; Mexico (Chihuahua).
Parnopes festivus Cockerell, 1894. Ent. News 5: 328. 2.

fulvicornis atlanticus Krombein. Md., Va., N. C., ? Fla. Host: Microbembex monodonta (Say)?
Parnopes westcottii atlanticus Krombein, 1958. Amer. Ent. Soc., Trans. 84: 164. 2, d.

Biology: Krombein, 1958. Amer. Ent. Soc., Trans. 84: 166-167.

1240 Hymenoptera in America North of Mexico

fulvicornis fulvicornis Cameron. Idaho and Iowa south to Calif. and Tex., and further south to
Guatemala. Host: Microbembex nigrifrons (Prov.); M. monodonta (Say)?
Parnopes fulvicornis Cameron, 1888. Biol. Cent.-Amer. Hym., v. 1, p. 466, pl. 20, figs. 1, la.
2

Parnopes westcottii Melander and Brues, 1902. Biol. Bul. 3: 39. d, 9.
Parnopes diadema Viereck, 1904. Amer. Ent. Soc., Trans. 30: 248. 6, °.
Parnopes taeniata Viereck, 1904. Amer. Ent. Soc., Trans. 30: 249. 9.

Biology: Bohart and MacSwain, 1940. Pan-Pacifie Ent. 16: 92-93 (host record for nigrifrons
misdet. as aurata). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp.
387-388.

Family DRYINIDAE

The North American fauna is very poorly known and probably a large number of taxa remain
to be collected and described. Except in the Aphelopinae the sexes are strongly dimorphic and
difficult to associate except by rearing. Males appear to be extremely rare or lacking in many
species. Classification of the family is based entirely on characters of the females.

These small wasps parasitize nymphs of Homoptera, principally species belonging to the Ful-
goridae, Cercopidae, Membracidae and Cicadellidae. The wasp larvae develop endoparasitically
in the host abdomen. In the later stages of development the parasite protrudes from the host ab-
domen as a cyst, formed from the moulted skins of the parasite in the Anteoninae, Dryininae
and Gonatopodinae, but from the host tissues in the Aphelopinae. Pupation takes place in a
cocoon spun on the host’s food plant or in the soil.

Taxonomy: Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 80-102. —Kieffer, 1914. Das Tierreich,
Lief. 41, pp. 7-222. —Richards, 1939. Roy. Ent. Soe. London, Trans. 89: 188-293 (British
species). —Richards, 1953. Roy. Ent. Soc. London, Trans. 104: 51-70 (keys to some genera
of Anteoninae and Dryininae). —Ponamarenko, 1970. Rev. d’Ent. de PUSSR 49: 423-427
(family reclassification; English translation, 1971. Ent. Rev. 49: 254-256).

Biology: Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1(1): 6-20.
—Haupt, 1916. Ztschr. f. Wiss. Insektenbiol. 12: 217-223. —Fenton, 1918. Ohio Jour. Sci. 18:
182-257.

Morphology: Swezey, 1903. Ohio Naturalist 3: 448-451. —Haupt, 1932. Zool. Anz. 99: 1-18.
—Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 409-412, figs. 47-50 (female, male thorax).

SUBFAMILY ANTEONINAE
Genus ANTEON Jurine
Genus ANTEON Subgenus ANTEON Jurine

Anteon Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 302.
Type-species: Anteon jurineanus Latreille. First included species.
Antaeon Haliday, 1833. Ent. Mag. 1: 275. Emend.
Prosanteon Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1: 66.
Type-species: Prosanteon chelogynoides Perkins. Orig. desig.
Liodryinus Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 325.
Type-species: Anteon doddi Kieffer. Orig. desig.
Xenanteon Kieffer, 1913. Soc. Ent. France, Bul., p. 300.
Type-species: Xenanteon reticulatuws Kieffer. Desig. by Muesebeck and Walkley,
1951.
Allanteon Kieffer, 1914. Das Tierreich, Lief. 41, p. 198.
Type-species: Anteon punctatus Kieffer. Monotypic.
arizonensis Perkins. Ariz. (Nogales). Host: Cicadellidae nymph.
Anteon arizonensis Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Buls 4375 ior Oe
canadensis (Ashmead). Que., Ont., Md.
Chelogynus canadensis Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 98. 9.

Superfamily BETHYLOIDEA 1241

hirtifrons Whittaker. B. C. (Hollyburn).
Anteon hirtifrons Whittaker, 1930. Ent. Soc. Wash., Proc. 32: 68. d.
minutus Ashmead. D. C.
Anteon minutus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 97. °.
pallidicornis Ashmead. Utah (Utah Lake).
Anteon pallidicornis Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 97. °.
politus Ashmead. Que., Ont.
Anteon politus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 96. 2.
popenoei (Ashmead). Kans. Assigned questionably to Anteon.
Dryinus popenoei Ashmead, 1888. Kans. State Col. Agr., Bul. 3: App., p. I. d.
puncticeps Ashmead. D. C., Va.
Anteon puncticeps Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 97. d.
rugosiceps Kieffer. Calif. (Santa Clara).
Anteon rugosiceps Kieffer, 1906. Berlin. Ent. Ztschr. 50: 238. d.
tibialis Say. D. C., Ind. Assigned questionably to Anteon.
Anteon tibialis Say, 1836. Boston Jour. Nat. Hist. 1: 284. d.
unifasciatus Ashmead. Fla. (Biscayne Bay).
Anteon unifasciatus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 95. 2.
whittakeri Muesebeck and Walkley. B. C.
Anteon flaviscapus Whittaker, 1930. Ent. Soc. Wash., Proce. 32: 67. ¢. Preoce.
Anteon whittakeri Muesebeck and Walkley, 1951. U. S. Dept. Agr., Monog. 2: 1040. N.
name.

Genus ANTEON Subgenus CHELOGYNUS Haliday

Chelogynus Haliday, 1838. Ent. Mag. 5: 518.
Type-species: Dryinus infectus Haliday. Desig. by Muesebeck and Walkley, 1951.
Neochelogynus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1:
60.
Type-species: Neochelogynus typicus Perkins. Orig. desig.
Lasianteon Kieffer, 1913. Soc. Ent. France, Bul. p. 200.
Type-species: Anteon rubrifrons Kieffer. Desig. by Muesebeck and Walkley, 1951.
Callianteon Kieffer, 1913. Soc. Ent. France, Bul., p. 300.
Type-species: Anteon bifasciatus Kieffer. Monotypic.
funestus Perkins. Ariz. Host: Cicadellidae nymph.
Chelogynus funestus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 48. 2, 3.
lusus Perkins. Ariz. (Tucson). Host: Cicadellidae nymph.
Chelogynus lusus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul.
4: 50. 2.
melanacrias Perkins. Ariz. (Nogales).
Chelogynus melanacrias Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 49. 2.
minimus Fenton. N. Y. (Barneveld).
Chelogynus minimus Fenton, 1927. U. S. Natl. Mus., Proce. 72 (8): 13. °.
osborni Fenton. Ohio (Sandusky). Host: Chlorotettix uwnicolor (Fitch).
Chelogynus osborni Fenton, 1918. Ohio Jour. Sci. 18: 256, 272. 2.
rugulosus Fenton. N. B. (Saint John).
Chelogynus rugulosus Fenton, 1927. U. S. Natl. Mus., Proce. 72 (8): 14. °.
virginiensis Fenton. Va. (Rosslyn).
Chelogynus virginiensis Fenton, 1927. U.S. Natl. Mus., Proc. 72 (8): 14. °.
vivariensis Bradley. N. Y., R. I.
Chelogynus vivariensis Bradley, 1926. Biol. Soc. Wash., Proc. 39: 7. °.
xanthothorax Bradley. N. Y.
Chelogynus xanthothorax Bradley, 1926. Biol. Soc. Wash., Proc. 39: 8. °.

1242 Hymenoptera in America North of Mexico

Genus DEINODRYINUS Perkins

Deinodryinus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4:

45.
Type-species: Deinodryinus paradoxus Perkins. Desig. by Muesebeck and Walkley,

1951.
atriceps (Brues). N. Y., Pa., Va., W. Va., Iowa, Kans.
Bocchus atriceps Brues, 1904. Canad. Ent. 36: 118. 9.
Deinodryinus variabilis var. carinatus Fenton, 1927. U. S. Natl. Mus., Proce. 72 (8): 11. @.
atriventris (Cresson). Tex.
Dryinus atriventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 193. 2.
ferrugineus (Brues). Tex.
Chelogynus ferrugineus Brues, 1905. Wis. Nat. Hist. Soc., Bul. 3: 183. @.
grandis (Brues). Mass., N. Y., W. Va.
Chelogynus grandis Brues, 1905. Wis. Nat. Hist. Soc., Bul. 3: 184. 9.
henshawi (Ashmead). Mass. N. Y., Wis., Nebr.
Chelogynus Henshawi Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 93. 9.
paradoxus Perkins. Ariz. (Nogales). Host: Cicadellidae sp. on oak.
Deinodryinus paradoxus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 46. 2, do.
pilosus Fenton. Ariz. (Chiricahua Mts.).
Deinodryinus pilosus Fenton, 1927. U.S. Natl. Mus., Proc. 72 (8): 12. °.
quercicola Perkins. Ariz. (Nogales).
Deinodryinus quercicola Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 47. @.
schaefferi (Brues). Ariz., Tex.
Chelogynus schaefferi Brues, 1907. Wis. Nat. Hist. Soc., Bul. 5: 101. @.
variabilis Fenton. Va. (Falls Church).
Deinodryinus variabilis Fenton, 1927. U. S. Natl. Mus., Proc. 72 (8): 10. 2.

Genus PRENANTEON Kieffer

Prenanteon Kieffer, 1913. Soc. Ent. France, Bul. p. 301.
Type-species: Anteon crassiscapus Kieffer. Desig. by Kieffer, 1914.
americanus Kieffer. Wis.
Prenanteon americanum Kieffer, 1914. Soc. Ent. France, Bul. p. 91.

bakeri (Kieffer). Nev. (Ormsby Co.).
Anteon bakeri Kieffer, 1906. Berlin. Ent. Ztschr. 50: 239. °.

micropunctatus Fenton. N. B. (Nerepis).
Prenanteon micropunctatus Fenton, 1927. U. S. Natl. Mus., Proc. 72 (8): 15. @.

Genus TRISANTEON Kieffer

Trisanteon Kieffer, 1913. Soc. Ent. France, Bul. p. 300.
Type-species: Anteon hirticornis Kieffer. Monotypic.

rugosus (Ashmead). III.
Anteon rugosus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 96. ¢.

Genus HESPERODRYINUS Perkins

Hesperodryinus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4:
40.
Type-species: Hesperodryinus arizonicus Perkins. Orig. desig.
amphiscepae Perkins. Ariz. (Nogales). Host: Acanalonia bivittata (Say) nymph.
Hesperodryinus amphiscepae Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 4: 41. @.

Superfamily BETHYLOIDEA 1243

arizonicus Perkins. Ariz. (Nogales). Host: Mistharnophantia sonorana Kirk. nymph.
Hesperodryinus arizonicus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., ‘Div.
Ent., Bul. 4: 42. @.

audax Perkins Ariz. (Nogales).
Hesperodryinus audax Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,

Bul. 4: 48. @.
Genus BOCCHUS Ashmead

Bocchus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 91.
Type-species: Bocchus flavicollis Ashmead. Orig. desig.
Phorbas Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 90. Preoce.
Type-species: Phorbas laticeps Ashmead. Monotypic.
Eukoebeleia Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1: 59.
Type-species: Hukoebeleia mirabilis Perkins. Orig. desig.
Phorbasia kieffer, 1914. Das Tierreich, Lief. 41, p. 58. N. name.
Neoanteon Fouts, 1922. Philippine Jour. Sci. 20: 633.
Type-species: Neoanteon rubrica Fouts. Orig. desig.

Taxonomy: Fenton, 1918. Ohio Jour. Sci. 18: 261 (key to spp.).

arizonica (Perkins). Ariz. (Nogales). Host: Bruchomorpha sp.?
Eukoebeleia arizonica Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,

Bul. 4: 44 9.

flavicollis Ashmead. Mich. (Marquette).
Bocchus flavicollis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 91. °.

laticeps (Ashmead). Fla.
Phorbas laticeps Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 90. 6.

mirabilis (Perkins). Ohio. Host: Bruchomorpha oculata Newm.
Eukoebeleia mirabilis Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,

Bul. 1: 59. 9.
SUBFAMILY DRYININAE
Genus MESODRYINUS Kieffer
Mesodryinus Kieffer, 1906. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 497.

Type-species: Dryinus niger Kieffer. Orig. desig.

alatus (Cresson). Tex.
Gonatopus (?) alatus Cresson, 1872. Amer. Ent. Soce., Trans. 4: 198. 9.

americanus (Ashmead). Fla.
Mystrophorus americanus Ashmead, 1887. Ent. Amer. 3: 128. 9.

crawfordi Krombein. Md. (Plummers Island).
Mesodryinus crawfordi Krombein, 1962. Biol. Soc. Wash., Proc. 75: 3. 9.
Genus TETRADRYINUS Kieffer
Tetradryinus Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 325.
Type-species: Bocchus flavipes Kieffer. Monotypic.
flavipes (Kieffer). Nev. (Ormsby Co.).
Bocchus flavipes Kieffer, 1906. Berlin. Ent. Ztschr. 50: 237 @.
Genus THAUMATODRYINUS Perkins

Thaumatodryinus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul.

1 (1): 58.
Type-species: Thaumatodryinus koebelei Perkins. Orig. desig.

Taxonomy: Krombein, 1952. Amer. Ent. Soc., Trans. 78: 97-99, 1 pl. (revis. gen. diag. and key
to spp.)

1244. Hymenoptera in America North of Mexico

perkinsi Krombein. Va. (Westmoreland St. Pk.).
Thaumatodryinus perkinsi Krombein, 1952. Amer. Ent. Soc., Trans. 78: 99, figs. 2, 2a, 2b.

OF
Genus PERODRYINUS Perkins

Perodryinus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4: 48.
Type-species: Perodryinus amoenus Perkins. Monotypic.

amoenus Perkins. Ariz. (Nogales). Host: Acanalonia bivittata (Say) nymph.
Perodryinus amoenus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 48. 2, 6.

SUBFAMILY GONATOPODINAE
Genus NEODRYINUS Perkins

Genus NEODRYINUS Subgenus NEODRYINUS Perkins

Neodryinus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1 (1):
50.
Type-species: Neodryinus koebelei Perkins. Orig. desig.
Prodryinus subg. Ctenodryinus Arle, 1935. Rio de Janeiro, Mus. Nac., Bol. 11: 46.
Type-species: Prodryinus (Ctenodryinus) affinis Arle. Monotypic.

The typical subgenus does not occur in North America.
Genus NEODRYINUS Subgenus PRODRYINUS Kieffer

Neodryinus subg. Psilodryinus Kieffer, 1906. In Andre, Spec. Hym., Eur. Alg., v. 9, p. 497.
Type-species: Dryinus acuticollis Kieffer. Orig. desig.

Prodryinus Kieffer, 1906. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 497.
Type-species: Dryinus brachycerus Kieffer. Orig. desig.

Phanerodryinus Roepke, 1916. Tijdschr. v. Ent. 59: 289.
Type-species: Phanerodryinus javanus Roepke. Orig. desig.

Psilodryinus has line priority over Prodryinus, but Richards (1953), the first reviser, placed
the former as a synonym of the latter.

arizonicus Perkins. Ariz. (Nogales). Host: Ormenis septentrionalis (Spin.) nymph.
Neodryinus arizonicus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 38. @.
carinatus (Fouts). Pa. (Carlisle).
Psilodryinus carinatus Fouts, 1924. Ent. Soc. Wash., Proc. 26: 160. ?.
dubiosus Perkins. Ariz. (Nogales).
Neodryinus dubiosus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 39. 2.
typhlocybae (Ashmead). N. Y., D. C., Ohio, IIl., lowa, Ark., Kans., Tex. Host: Metcalfa
pruinosa (Say), Anormenis septentrionalis (Spin.).
Labeo typhlocybae Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 89. d.
Dryinus ormenidis Ashmead, 1903. Ent. News 14: 192. 2.
Dryinus nigrellus Brues, 1904. Canad. Ent. 36: 117. ¢.

Biology: Dean and Bailey, 1961. Jour. Econ. Ent. 54: 1104-1106, 2 figs.

Genus CYRTOGONATOPUS Kieffer

Pachygonatopus subg. Cyrtogonatopus Kieffer, 1907. In Wytsman, Gen. Ins., fase. 54, p.
1g}
Type-species: Pachygonatopus (Cyrtogonatopus) breviforceps (Kieffer). Monotypic.
Taxonomy: Richards, 1969. Roy. Ent. Soc. London, Proc. (B) 38: 80 (genus redescription).

clavicornis Fenton. Tex. (Brownsville).
Cyrtogonatopus clavicornis Fenton, 1927. U. S. Natl. Mus., Proce. 72(8): 3.

Superfamily BETHYLOIDEA 1245

Genus PRISTOGONATOPUS Kieffer

Pristogonatopus Kieffer, 1913. Lab. Zool. Gen. e. Agr. Portici, Bol. 7: 325.
Type-species: Gonatopus dentatiforceps Kieffer. Orig. desig.
sjostedti (Kieffer). Tex.
Gonatopus sjostedti Kieffer, 1904. Arkiv. for Zool. 1: 525. @.

Genus DICONDYLUS Haliday

Dicondylus Haliday, 1829-30. In Curtis, Guide Brit. Ins., col. 110.
Type-species: Dryinus bicolor Haliday. Desig. by Kieffer, 1914.
Labeo Haliday, 1833. Ent. Mag. 1: 273. Preoce.
Type-species: Labeo vitripennis Haliday. Monotypic.
Labeola Haldeman, 1842. Acad. Nat. Sci. Phila., Proc. 1: 192. N. name.
Laberius Kieffer, 1914. Das Tierreich, Lief. 41, p. 59. N. name.
Laberinus(!) Ogloblin, 1932. Rev. de Ent. Sao Paulo 2: 266. Misspelling.
albitarsis (Kieffer). Nev.
Labeo albitarsis Kieffer, 1906. Berlin. Ent. Ztschr. 50: 2.
nasutus (Ashmead). Fla. (Jacksonville).
Embolemus nasutus Ashmead, 1887. Ent. Amer. 3: 75. ¢.
texanus (Ashmead). Tex. Assigned questionably to Dicondylus.
Labeo texanus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 89. ¢.

Genus PSEUDOGONATOPUS Perkins

Pseudogonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul.
1: 34.
Type-species: Pseudogonatopus kurandae Perkins. Orig. desig.
americanus Perkins. Ohio (Columbus).
Pseudogonatopus americanus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 1: 37. @.
arizonicus Perkins. Ariz. (Nogales). Host: Stobaera? sp., nymph and adult.
Pseudogonatopus arizonicus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 36. 2.
autoxenobius Perkins. Ariz. (Nogales). Host: Stobaera? sp. adult.
Pseudogonatopus autoxenobius Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 4: 37. @.
iowensis Fenton. Iowa. Host: Delphacodes lutulenta (Van D.).
Pseudogonatopus iowensis Fenton, 1924. Ohio Jour. Sci. 24: 191. 2.
magnus Brown. Ind. (Wawasee Lake).
Pseudogonatopus magnus Brown, 1940. Ent. News. 51: 11. 9.
stenocrani dubiosus Perkins. Ohio.
Pseudogonatopus stenocrani var. dubiosus Perkins, 1905. Hawaii. Sugar Planters’ Assoc.
Expt. Sta., Div. Ent., Bul. 1: 39. 2.
stenocrani stenocrani Perkins. Ohio.
Pseudogonatopus stenocrani Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 1: 38. @.

Genus AGONATOPOIDES Perkins
Agonatopoides Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4:
33.

Type-species: Agonatopoides synchromus Perkins. Monotypic.
synchromus Perkins. Ariz. (Nogales). Host: Bostaera nasuta Ball, nymph and adult.
Agonatopoides synchromus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 33.

1246 Hymenoptera in America North of Mexico

Genus HAPLOGONATOPUS Perkins

Haplogonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1:
39.
Type-species: Haplogonatopus apicalis perkins. Orig. desig.
americanus Perkins. Va., Ohio, Iowa. Host: Delphacodes campestris (Van D.); D. lutulenta
(Van D.).
Haplogonatopus americanus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt Sta., Div.
Ent., Bul. 1: 40. @

Genus EUCAMPTONYX Perkins
Eucamptonyx Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4:
28.

Type-species: Eucamptonyx testaceus Perkins. Monotypic.
secundus Fenton. Ohio, Fla., Md., Kans.
Eucamptonyx secundus Fenton, 1927. U.S. Natl. Mus., Proc. 72 (8): 5. 2.
testaceus Perkins. Ariz. (Nogales).
Eucamptonyx testaceus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 28. °.

Genus APTERODRYINUS Perkins

Apterodryinus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4:
14.
Type-species: Apterodryinus torvus Perkins. Monotypic.
torvus Perkins. Ariz. (Nogales). Host: Mistharnophantia sonorana Kirk.; Acanalonia
immaculata (Kirk.).
Apterodryinus torvus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 14. 9.

Genus PACHYGONATOPUS Perkins

Pachygonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul.
1: 45.
Type-species: Pachygonatopus melanias Perkins. Orig. desig.
minimus Fenton. S. Dak., Man. Host: Macrosteles fascifrons (Stal).
Pachygonatopus minimus Fenton, 1927. U. S. Natl. Mus., Proce. 72 (8): 6. °.

Taxonomy: Barrett, Westdal and Richardson, 1965. Canad. Ent. 97: 217-219, figs. 1-10 (egg,
larvae, pupa, adults).

Biology: Barrett, Westdal and Richardson, 1965. Canad. Ent. 97: 219-221 (life history).

nearcticus Fenton. Iowa (Sioux City).
Pachygonatopus nearcticus Fenton, 1927. U. S. Natl. Mus., Proc. 72 (8): 6. ?.

Genus CHALCOGONATOPUS Perkins

Chalcogonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul.
1: 46.
Type-species: Chalcogonatopus gigas Perkins. Orig. desig.
Chalcogonatopus subg. Eugonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt.
Sta., Div. Ent., Bul. 1: 46.
Type-species: Chalcogonatopus (Eugonatopus) pseudochromus Perkins. Monotypic.
areolatus Fenton. Va. (Falls Church).
Chalcogonatopus areolatus Fenton, 1927. U. S. Natl. Mus., Proce. 72 (8): 7. °.

argyrias Perkins. Ariz. (Nogales).
Chalcogonatopus argyrias Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 20. °.

californicus (Ashmead). Calif.
Gonatopus californicus Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 85. @.

Superfamily BETHYLOIDEA 1247

echo Perkins. Ariz. (Nogales).
Chalcogonatopus echo Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 23. 2.
euprepes Perkins. Ariz. (Nogales).
Chalcogonatopus euprepes Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 18. °.
Chalcogonatopus euprepes var. a Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 4: 19. 2.
Chalcogonatopus cuprepes (!) Kieffer, 1914. Das Tierreich, Lief. 41, p. 99.
euprepoides Perkins. Ariz. (Nogales).
Chalcogonatopus euprepoides Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 4: 19. 9°.
Chalcogonatopus cuprepoides (!) Kieffer, 1914. Das Tierreich, Lief. 41, p. 99.
flavifrons (Ashmead). N. Y. (Albany).
Gonatopus flavifrons Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 84, pl. 5, fig. 4. °.
frequens Perkins. Ariz. (Nogales).
Chalcogonatopus frequens Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 21. @.
harpax Krombein. W. Va. (Lost River St. Pk.).
Chalcogonatopus harpax Krombein, 1956. Ent. Soc. Wash., Proc. 58: 158, fig. 1. 2.
herbarum Perkins. Ariz. (Nogales).
Chalcogonatopus herbarum Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 23. 2.
koebelei Perkins. Ariz. (Nogales).
Chalcogonatopus koebelei Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 26. 2.
leptias Perkins. Ariz. (Nogales).
Chalcogonatopus leptias Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 17. 2.
niger Fenton. Iowa. Host: Scaphoideus sp., probably immistus Say.
Chalcogonatopus nigrus Fenton, 1924. Ohio Jour. Sci. 24: 193, fig. 3. 2.
paraleptias Perkins. Ariz. (Nogales).
Chalcogonatopus paraleptias Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 4: 18. @.
perdebilis Perkins. Ariz. (Nogales).
Chalcogonatopus perdebilis Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 27. @.
pseudochromus Perkins. Ohio (Columbus).
Chalcogonatopus (Eugonatopus) pseudochromus Perkins, 1905. Hawaii. Sugar Planters’
Assoc. Expt. Sta., Div. Ent., Bul. 1: 48. ¢.
rapax Perkins. Ariz. (Nogales). Host: Cicadellidae nymph on willow.
Chalcogonatopus rapax Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 20. 2.
simulator Perkins. Ariz. (Nogales).
Chalcogonatopus simulator Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 26. 2.
solitarius Perkins. Ariz. (Nogales).
Chalcogonatopus solitarius Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 22. 9°.
unicus Perkins. Ariz. (Nogales).
Chalcogonatopus unicus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 25. °.
xestocephalus Perkins. Ariz. (Nogales).
Chalcogonatopus xestocephalus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 4: 24. @.

1248 Hymenoptera in America North of Mexico

Genus GONATOPUS Ljungh

Gonatopus Ljungh, 1810. Beitr. Naturk., v. 2, p. 161.
Type-species: Gonatopus formicarius Ljungh. Monotypic.

Revision: Richards, 1939. Roy. Ent. Soc. London, Trans. 89: 185-344 (European spp.).
Taxonomy: Fenton, 1918. Ohio Jour. Sci. 18: 265-266 (key to spp.).
Biology: Haupt, 1916. Ent. Gesell. Halle, Mitt. 10: 41-50.

Morphology: Haupt, 1932. Zool. Anz. 99: 1-18.
affinis Fenton. Ohio (Bay View). Host: Psammotettix affinis (G. and B.).
Gonatopus affinis Fenton, 1918. Ohio Jour. Sci. 18: 266. 2, d.
agropyrus Fenton. Iowa (Ames). Host: Psammotettix affinis (G. and B.).
Gonatopus agropyrus Fenton, 1921. Canad. Ent. 58: 71. 2, d.
ashmeadi Kieffer. Ala., Fla., Tex. Host: Delphacodes lutulenta (Van D.)?
Gonatopus bicolor Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 85. 2. Preoce.
Gonatopus ashmeadi Kieffer, 1904. In Andre, Spec. Hym. Eur. Alg., v. 9, p. 108. N. name.
contortulus Patton. Que., Mass., Conn., Ohio. Host: Latalus sayi (Fitch).
Gonatopus contortulus Patton, 1879. Canad. Ent. 11: 65. °.
curriei Krombein. Md. (Plummers Island).
Gonatopus curriei Krombein, 1962. Biol. Soc. Wash., Proc. 75: 4. 2.
cyphonotus Bradley. B. C. (Downie Creek, Big Bend Country).
Gonatopus cyphonotus Bradley, 1906. Canad. Ent. 38: 380. °.
foutsi Whittaker. B. C. (Chilliwack).
Gonatopus foutsi Whittaker, 1928. Ent. Soc. London, Trans. p. 388. ?.
inimicis Fenton. Ohio. Host: Endria inimica (Say).
Gonatopus inimicis Fenton, 1918. Ohio Jour. Sci. 18:.270. 2.
longicornis (Brues). Pa. (Saegerstown). Questionably assigned to Gonatopus.
Phorbas longicornis Brues, 1907. Wis. Nat. Hist. Soe., Bul. 5: 152. ¢.
peculiaris Brues. Tex. (Austin).
Gonatopus peculiaris Brues, 1903. Amer. Ent. Soc., Trans. 29: 125. 9.
punctatus Fenton. Ohio (Columbus). Host: Latalus sayz (Fitch).
Gonatopus punctatus Fenton, 1918. Ohio Jour. Sci. 18: 268. 3, 9.
septentrionalis Whittaker. B. C. (Chilliwack).
Gonatopus septentrionalis Whittaker, 1928. Ent. Soc. London, Trans. p. 388. @.
similis Fenton. lowa (Ames). Host: Psammotettix affinis (G. and B.).
Gonatopus similis Fenton, 1921. Canad. Ent. 53: 71. 2.

Genus NEOGONATOPUS Perkins

Neogonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1:
42.
Type-species: Neogonatopus ombrodes Perkins. Orig. desig.
Allogonatopus Haupt, 1938. Ztschr. f. Naturw. 92: 21.
Type-species: Allogonatopus procerus Haupt. Orig. desig.

Taxonomy: Fenton, 1918. Ohio Jour. Sci. 18: 265-266 (key to spp.).
brunnescens Perkins. Ohio.
Neogonatopus brunnescens Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 1: 44. 2.
erythrodes Perkins. Ohio, Iowa. Host: Endria inimica (Say).
Neogonatopus erythrodes Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent, Bull 12432 9:
longitarsis (Ashmead). Fla., La.
Labeo longitarsis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 88. 6.

Superfamily BETHYLOIDEA 1249

mimoides Perkins. Ariz., Utah.
Neogonatopus mimoides Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 34. 9.
mimus Perkins. Ariz. (Nogales). Host: Cicadellidae.
Neogonatopus mimus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 33. 9.
obscurissimus Perkins. Ohio (Columbus). Host: Deltocephalus spp.
Neogonatopus obscurissimus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta.,
Div. Ent., Bul. 1: 43. °.
ombrodes Perkins. Ohio, Colo., Calif., Conn., S. Dak. Host: Macrosteles divisus (Uhl.).
Neogonatopus ombrodes Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 1: 43. °.

Biology: Ainslie, 1920. Ent. News 31: 169, 187.
pallidiceps Perkins. Calif. (Alameda).
Neogonatopus pallidiceps Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 1: 45. 9.

Genus AGONATOPUS Perkins

Agonatopus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 4: 29.
Type-species: Agonatopus pallidicornis Perkins. Desig. by Richards, 1939.
dubiosus Perkins. Ariz. (Nogales).
Agonatopus pallidicornis var. dubiosus Perkins, 1907. Hawaii. Sugar Planters’ Asoc. Expt.
Sta., Div. Ent., Bul, 4: 30. 9.
ferrugineus Perkins. Ariz. (Nogales).
Agonatopus ferrugineus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 30. 2.
heterothorax Perkins. Ariz. (Nogales). Host: Cicadellidae nymph on grass.
Agonatopus heterothorax Perkins, 1907. Hawaii. Sugar Planters’ Assoc Expt. Sta., Div.
Ent., Bul. 4: 32. 9.
innitidus Perkins. Ariz (Nogales). Host: Cicadellidae nymph on grass.
Agonatopus innitidus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 31. @.
pallidicornis Perkins. Ariz.
Agonatopus pallidicornis Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 29. °.
picescens Perkins. Ariz. (Nogales).
Agonatopus picescens Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 30. ¢.

suturalis Fenton. Ariz. (Tucson).
Agonatopus suturalis Fenton, 1927. U.S. Natl. Mus., Proce. 72 (8): 9. 9.

Genus EPIGONATOPUS Perkins

Epigonatopus Perkins, 1905. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul. 1:
45.
Type-species: Epigonatopus solitarius Perkins. Orig. desig.
americanus Fenton. Iowa (Ames).
Epigonatopus americanus Fenton, 1921. Canad. Ent. 53: 70. °.

plesuis Fenton. Ont., S. Dak. Host: Macrosteles fascifrons (Stal).
Epigonatopus plesuis Fenton, 1927. U. S. Natl. Mus., Proc. 72 (8): 9. °.
Biology: George, 1959. Canad. Ent. 91: 256.

tenuis Fenton. Ind. (La Fayette).
Epigonatopus tenuis Fenton, 1927. U. S. Natl. Mus., Proe. 72 (8): 8. 2.

1250 Hymenoptera in America North of Mexico

Genus DIGONATOPUS Kieffer

Digonatopus Kieffer, 1913. Lab. Zool. Gen. e Agr. Portici, Bol. 7: 325.
Type-species: Discondylus (!) javanus Perkins. Orig. desig-
haplothorax (Perkins). Ariz. (Nogales).
Gonatopus haplothorax Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 35. 9.

SUBFAMILY APHELOPINAE

Genus APHELOPUS Dalman

Dryinus subg. Aphelopus Dalman, 1823. Analecta Ent., p. 8.
Type-species: Aphelopus atratus (Dalman). Desig. by Westwood, 1840.

Taxonomy: Fenton, 1918. Ohio Jour. Sci. 18: 275 (key to spp.).

Morphology: Haupt, 1932. Zool. Anz. 99: 1-18.
affinis Ashmead. Canada.
Aphelopus affinis Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 102. 9.
albopictus Ashmead. Md., D. C., Va.
Aphelopus albopictus Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 100. °.
americanus Ashmead. Fla.
Aphelopus americanus Ashmead, 1887. Ent. Amer. 3: 74. 6.
arizonicus Perkins. Ariz. (Nogales). Host: Dikraneura sp.
Aphelopus arizonicus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Buls4as5osce oe 3
bicolor Fenton. Iowa, Ark., Del. Host: Hymetta trifasciata (Say).
Aphelopus bicolor Fenton, 1924. Ohio Jour. Sci. 24: 192. 2.
comesi Fenton. Del., Ohio, Mich., Tenn.. Iowa, Ark., La. Host: Erythroneura spp.
Aphelopus comesi Fenton, 1918. Ohio Jour. Sci. 18: 277. d.
dikraneuri Fenton. Ohio, Iowa. Host: Forcipata sp., prob. loca DeL. and C.
Aphelopus dikraneuri Fenton, 1918. Ohio Jour. Sci. 18: 276. 2, 3.
microleucus Perkins. Ariz. (Nogales). Host: Typhlocyba sp.
Aphelopus microleucus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent.,
Bul. 4: 55. ¢.
microtomus Whittaker. B. C. (Chilliwack).
Aphelopus microtomus Whittaker, 1928. Ent. Soc. London, Trans. p. 389. °.
pilicornis Whittaker. B. C. (Chilliwack).
Aphelopus pilicornis Whittaker, 1928. Ent. Soc. London, Trans. p. 389. 2, ¢.
pulcherrimus Perkins. Ariz. (Nogales). Host: Erythroneura sp.; Typhlocyba sp.?
Aphelopus pulcherrimus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div.
Ent., Bul. 4: 54. ? 6.
rufiventris Ashmead. Fla. (Jacksonville).
Aphelopus rufiventris Ashmead, 1893. U. S. Natl. Mus., Bul. 45: 100. 2.
theliae Gahan. N. Y. (Cold Spring Harbor, L. I.). Host: Thelia bimaculata (F.).
Aphelopus theliae Gahan, 1918. Canad. Ent. 50: 151. 2, d.

Biology: Kornhauser, 1919. Jour. Morph. 32: 547-554, figs. 1-4.

typhlocybae Muesebeck. N. Y., Md. Host: Typhlocyba pomaria McA.; T. froggatti Baker (in
New Zealand).
Aphelopus typhlocybae Muesebeck, 1936. Ent. Soc. Wash., Proc. 37: 167. 2, 3.

Biology: Steiner, 1936. Jour. Econ. Ent. 29: 632. —Dumbleton, 1937. Jour. Sci. and Technol.
New Zealand 18: 869.

varicornis Brues. Mass. (Woods Hole).
Aphelopus varicornis Brues, 1906. Wis. Nat. Hist. Soc., Bul. 4: 143. 2.

Superfamily BETHYLOIDEA 1251

viduus Perkins. Ariz. (Nogales).
Aphelopus viduus Perkins, 1907. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Div. Ent., Bul.
4: 54. 6.

UNPLACED TAXA OF DRYINIDAE

Dryinus bifasciatus Say, 1828. Contrib. Maclur. Lye. Phila. 1: 81. 2. Ind. Possibly
Deinodryinus Perkins.

Family EMBOLEMIDAE

Genus AMPULICOMORPHA Ashmead

Ampulicomorpha Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 79.
Type-species: Ampulicomorpha confusa Ashmead. Orig. desig.
confusa Ashmead. Ont., Md., Va., N. C., Ga., Wis., Mo., Colo., Sask., B. C., Calif. Host: Nymphs
of Epiptera floridae (Wlkr.) on rotten pine and oak logs, and E. pallida (Say) feeding on
fungi under bark.
Ampulicomorpha confusa Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 80. “3” = 9.

Biology: Bridwell, 1958. Ent. Soc. Wash., Proc. 60: 23-26.

Genus EMBOLEMUS Westwood

Embolemus Westwood, 1833. London and Edinb. Phil. Mag. and Jour. Sci. 2: 444.
Type-species: Embolemus ruddii Westwood. Monotypic.

Myrmecomorphus Westwood, 1833. Mag. Nat. Hist. 6: 496.
Type-species: Myrmecomorphus rufescens Westwood. Monotypic.

Embolimus Agassiz, 1846. Nomencl. Zool., Ind. Univ., p. 137. Emend.

Formila de Romand, 1846. Soc. Ent. France, Ann. (2) 4: Bul., p. XXXII.
Type-species: Formila chevrolatii de Romand. Monotypic.

Pedinomma Foerster, 1856. Hym. Stud., v. 2, p. 94. Unnecessarily proposed as a n. name

for Myrmecomorphus Westwood.

Taxonomy: Richards, 1939. Roy. Ent. Soc. London, Trans. 89: 294.
Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 412, figs. 51-52 (female, male
thorax).

nearcticus (Brues). Mass., N. Y., Va.?
Pedinomma nearcticum Brues, 1922. Psyche 29: 7. “5” = °.

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1253

Superfamily SCOLIOIDEA
By KARL V. KROMBEIN

Taxonomy: Ashmead, 1903-1904. Canad. Ent. 35: 4-8, 39-46, 95-107, 155-158, 199-205, 303-310,
323-332; 36: 5-9 (keys to genera).

Family TIPHIIDAE

So far as is known all of the North American taxa are parasites of larvae of fossorial species.
In the primitive subfamily Tiphiinae, species of Tiphia F. have been reported as parasites of
scarabaeid larvae. A few exotic species have been liberated in North America for control of
economically important pest scarab species. Hosts of the other genera of Tiphiinae are unknown.
In the Myzininae, species of Myzinum Latr. have scarabaeid larvae as hosts; it is anticipated
that our native Pterombrus Sm. parasitize tiger beetle larvae as do some of the Neotropical spe-
cies. Biology is unknown for the Anthoboscinae and Brachycistidinae, but the species probably
have as hosts deserticolous scarabaeid larvae. The Methochinae parasitize tiger beetle larvae,
and the Myrmosinae are parasites of ground-nesting aculeate Hymenoptera.

Revision: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 45-115, 30 figs. (western North

America species). —Allen, 1966. Amer. Ent. Soc., Trans. 92: 231-356, 40 figs., 19 maps
(eastern North America species).

Taxonomy: Ashmead, 1903. Canad. Ent. 35: 4-7, 39-41, 155-156, 201-204. —Krombein, 1940.
Amer. Ent. Soc., Trans. 65: 419. —Pate, 1947. N. Y. Ent. Soc., Jour. 55: 115-143.

SUBFAMILY TIPHIINAE

Taxonomy: Hedicke, 1936. Hym. Cat., Pars. 1, Tiphiidae, pp. 1-32 (catalog of world species).
—Allen, 1962. Ent. Soc. Amer., Trans. 88: 21-75, 30 figs. (redescription of types of
American species in British museum).

Genus TIPHIA Fabricius
Genus TIPHIA Subgenus TIPHIA Fabricius

Tiphia Fabricius, 1775. Systema Ent., p. 553.
Type-species: Tiphia femorata Fabricius. Desig. by Latreille, 1810.

Tiphiana (!) Rafinesque, 1815. Analyse Nat., p. 124. Emend.
So far as is known species of this subgenus parasitize larvae of various Scarabaeidae

(Coleoptera). Detailed liberation records of the Oriental species released in the United States
for control of the introduced Popillia, Anomala, Autoserica, and Serica were published by

1254 Hymenoptera in America North of Mexico

Krombein (1948, Ent. Soe. Amer., Ann. 41: 58-62). Of the liberated species only vernalis Rohwer,
popilliavora Rohwer, and asericae Allen and Jaynes established themselves; sternata Parker is
doubtfully established; biseculata Allen and Jaynes, castaneaevora Parker, chosensis Allen,
frater Parker, matura Allen and Jaynes, notopolita var. alleni Roberts, pullivora Allen and
Jaynes, and totopunctata Allen and Jaynes apparently did not become established.

Tiphia femorata Fabricius and T. morio Fabricius were introduced from Europe and liberated
in New York in 1956 and 1957 for control of Amphimallon majalis (Raz.), the European chafer;
so far as is known neither species became established.

Only the typical subgenus occurs in the New World.

Revision: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 1-24, 13 figs. (eastern species).
—Allen, 1971. Amer. Ent. Soc., Trans. 97: 201-359, 147 figs., 13 maps (species of western
North America).

Taxonomy: Allen, 1961. Amer. Ent. Soc., Trans. 87: 1-20 (key to species north of Mexico).
—Evans, 1965. Ent. Soc. Wash., Proc. 67: 91 (larva).

alamosae Allen. Colo. (Alamosa Co.); Mexico (Chihuahua).
Tiphia alamosae Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 54, figs. 1, 6, 11. 3, @.

Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 264, figs. 63-68, map 5. 6, 9.
andersoni Allen. Ariz. south in Mexico to Michoacan and Morelos.
Tiphia andersoni Allen, 1971. Amer. Ent. Soc., Trans. 97: 288, figs. 100-103, map 8. 6, °.
anguis Allen. Wyo., B. C. to Calif.
Tiphia anguis Allen, 1971. Amer. Ent. Soc., Trans. 97: 278, figs. 94-98, map 7. ¢, 2.

arcuata Allen. Ariz. (Chiricahua Mts.).

Tiphia arcuata Allen, 1971. Amer. Ent. Soc., Trans. 97: 315, figs. 127, 129, 130, map 11. 2.
asensoria Allen. Fla. (Fort Lauderdale).

Tiphia asensoria Allen, 1966. Amer. Ent. Soc., Trans. 92: 317, map 12. 6.
asericae Allen and Jaynes. N. Y., N. J., Pa.; Korea. Introduced from Korea. Host: Awtoserica

castanea (Arrow); Serica peregrina Chapin.
Tiphia asericae Allen and Jaynes, 1930. U.S. Natl. Mus., Proc. 76 (17): 74. 2, 6.

Biology: Clausen, Jaynes, and Gardner, 1933. U. S. Dept. Agr., Tech. Bul. 366: 45-46.
—Gardner and Parker, 1940. U.S. Dept. Agr., Tech. Bul. 738: 21. —Clausen, 1956. U.S.
Dept. Agr., Tech. Bul. 1139: 181 (life history).

atlantis Allen. N. H. to Fla., Mich., Ill., Kans., Ariz. ?
Tiphia atlantis Allen, 1934. Amer. Ent. Soc., Trans. 60: 314. @.

Taxonomy: Allen, 1961. Amer. Ent. Soc., Trans. 87: 16. 6. —Allen, 1966. Amer. Ent. Soc.,
Trans. 92: 336, figs. 28, 32, map 17. 2, 6.

barberi Allen. Mont., Wyo., Ariz.
Tiphia barberi Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 57, figs. 13, 28, 29. ¢.

Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 270, figs. 75-78, map 6. 6, °.

berbereti Allen. Nebr. (Ainsworth). Host: Phyllophaga anxia (LeC.).
Tiphia berbereti Allen, 1970. Ent. Soc. Amer., Ann. 63: 473. 2, d.
Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 274, figs. 82-85, map 6. 2, ¢.

Biology: Berberet and Helms, 1970. Ent. Soc. Amer., Ann. 63: 471-473 (life history).
biseculata Allen and Jaynes. Liberated in N. Y. and N. J. but not established. Introduced from
Japan. Host: Anomala orientalis Waterh.; Popillia japonica Newm.
Tiphia biseculata Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art. 17: 85, figs. 30,
Slory.

Biology: Clausen, Jaynes and Gardner, 1933. U. S. Dept. Agr., Tech. Bul. 366: 38-39 (life
history).
boharti Allen. Oreg. (Bend).
Tiphia boharti Allen, 1971. Amer. Ent. Soc., Trans. 97: 287, map 8. ¢.

Superfamily SCOLIOIDEA 1255

canamexica Rohwer. Mont., S. Dak., Colo., Utah, Calif., Ariz. south in Mexico to Chiapas and
Yucatan.
Tiphia canamexica Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 452. °.
Tiphia dreisbachi Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 70. d, °.
Tiphia canamexicana (!) Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 85. 2.

Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 248, figs. 41-43, map 3. d, °.
castaneaevora Parker. Liberated in N. J. and Pa. but not established. Introduced from Japan.
Host: Autoserica castanea (Arrow).
Tiphia castaneaevora Parker, 1937. N. Y. Ent. Soc., Jour. 45: 288. 2, d.
chosensis Allen. Liberated in Conn. and N. J. but not established. Introduced from Korea.
Host: Anomala orientalis Waterh.
Tiphia (Tiphia) chosensis Allen, 1969. Amer. Ent. Soc., Trans. 95: 359. 2, d.

Taxonomy: Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art. 17: 66, figs. 2, 5, 7, 11, 24.
9, 6. (Misdet. as bicarinata Cameron).

Biology: Clausen, Jaynes and Gardner, 1933. U. S. Dept. Agr., Tech. Bul. 366: 43-44 (life

history, as bicarinata).
cochiseae Allen. N. Mex., Ariz., south in Mexico to Jalisco and Puebla.

Tiphia cochiseae Allen, 1971. Amer. Ent. Soc., Trans. 97: 282, fig. 99, map 7. d.
coloradensis Allen. Colo.

Tiphia coloradensis Allen, 1971. Amer. Ent. Soc., Trans. 97: 314, fig. 126, map 11. 2.
conformis Malloch. Mass., Mich. and Nebr. south to Fla. and Tex., Ariz.

Tiphia conformis Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 22. 9, 6.

Tiphia imitatrix Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 22. °.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 229 (lectotype designation
conformis). —Allen, 1934. Amer. Ent. Soc., Trans. 60: 312. —Allen, 1966. Amer. Ent. Soc.,
Trans. 92: 295, fig. 14, map 10. 6, 2.

convexa Allen. N. H. to N. C., Mich., Kans., Mont.
Tiphia convexa Allen, 1934. Amer. Ent. Soc., Trans. 60: 306. ¢.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 290, map 9. 6, 2.

crassipunctata Allen. N. C.
Tiphia crassipunctata Allen, 1934. Amer. Ent. Soe., Trans. 60: 307. 6.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 254, map 3. d.
dallasae Allen. Tex. (Dallas, Victoria).
Tiphia dallasae Allen, 1966. Amer. Ent. Soc., Trans. 92: 339, fig. 9, map 17. °.
dawsoni Allen. Nebr., Mont., Idaho, Tex.
Tiphia dawsoni Allen, 1971. Amer. Ent. Soc., Trans. 97: 259, fig. 62, map 5. d.

dentonae Allen. Tex., Kans.

Tiphia dentonae Allen, 1966. Amer. Ent. Soc., Trans. 92: 300, map 10. d.
dryophila Krombein. N. J., N. C., Fla., Ohio.

Tiphia dryophila Krombein, 1953. Wasmann Jour. Biol. 10: 301. d, °.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 299, map 10. d, °.
egregia Viereck. N. H. to S. Dak., south to Fla. and Tex.

Tiphia eyregia (!) Viereck, 1906. Ent. News 17: 303. 6.
Tiphia egregia Viereck, 1906. Ent. News 17: 350.
Tiphia tuberculata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 14. 2, ¢. Preoce.
Tiphia aterrima Malloch, 1918. Il. Nat. Hist. Survey, Bul. 13: 19. °.
Tiphia papillata Roberts, 1930. Canad. Ent. 62: 190. N. name.
Tiphia diminuta Roberts, 1933. Kans. Ent. Soe., Jour. 6: 91. 2, d.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 229-280 (lectotype designation
aterrima, tuberculata). —Allen, 1934. Amer. Ent. Soc., Trans. 60: 312. —Allen, 1966.
Amer. Ent. Soc., Trans. 92: 325, fig. 34, map 15. 6, °.

elaka Allen. Fla.
Tiphia elaka Allen, 1966. Amer. Ent. Soc., Trans. 92: 289, fig. 6, map 9. d.

1256 Hymenoptera in America North of Mexico

femorata Fabricius. Liberated in N. Y. but not established. Introduced from Europe. Host:
Amphimallon majalis (Raz.).
Tiphia femorata Fabricius, 1775. Systema Ent., p. 353. ¢.
Sphex palmipes Schrank, 1781. Enum. Ins. Austriae, p. 384.
Bethylus glabrata Fabricius, 1798. Sup. Ent. System., p. 254. ..
Bethylus villosa Fabricius, 1804. Systema Piezatorum, p. 236.
Bethylus pilipennis Klug, 1808. Mag. Gesell. Naturf. Freunde Berlin 2: 49. ¢.
Bethylus ater Giraud, 1856. Zool.-Bot. Ver., Verhandl. 6: 184.
Tiphia laeviceps Tournier, 1889. Soc. Ent. Belg., Ann. 33: 21. 9.
Tiphia austriaca Tournier, 1889. Soc. Ent. Belg., Ann. 33: 24. 2.
fenestrata Klug. N. J., N. C., Fla., Ill., Miss., La.
Bethylus fenestratus Klug, 1810. In Weber, Beitr. z. Naturk. 2: 193. 9.
Tiphia confusa Allen, 1934. Amer. Ent. Soc., Trans. 60: 302. °.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 253. 9. —Allen, 1972. Ent. Soc. Wash.,

Proc. 74: 381-383 (redescription and synonymy).

fisheri Allen. Va. (Cape Henry).
Tiphia fisheri Allen, 1966. Amer. Ent. Soc., Trans. 92: 301, map 10. ¢.

flavipennis Spinola. Calif. (?); Mexico (Guerrero).
Tiphia flavipennis Spinola, 1841. Soe. Ent. France, Ann. 10: 102. 2.
Tiphia elegans Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 240. 2, 3.
Tiphia ochroptera Dalla Torre, 1897. Cat. Hym., v. 8, p. 189. N. name err. proposed for

flavipennis Spinola, not Smith, 1857.

Taxonomy: Turner, 1908. Ann. and Mag. Nat. Hist. (8) 2: 131 (synonymy). —Allen, 1962.
Amer. Ent. Soc., Trans. 88: 38 (redescription elegans female).
floridana flavida Allen. Central Fla.
Tiphia floridana flavida Allen, 1966. Amer. Ent. Soc., Trans. 92: 279, map 7. 9°, 6.

floridana floridana Robertson. N. Y. to Fla.
Tiphia floridana Robertson, 1901. Amer. Ent. Soc., Trans. 27: 195. 2, ¢.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 23. —Allen, 1934. Amer. Ent. Soc.,
Trans. 60: 314. —Krombein, 1953. Wasmann Jour. Biol. 10: 305. —Allen, 1966. Amer. Ent.
Soc., Trans. 92: 276, figs. 5, 33, 38, map 7. 6, @.

fortistriolata Cameron. B. C. south to Calif. and N. Mex.; Mexico (Michoacan, Morelos,
Veracruz).
Tiphia fortistriolata Cameron, 1907. Invertebrata Pacifica 1: 170. 6.
Tiphia bakeri Allen, 1961. Amer. Ent. Soc., Trans. 87: 12. 6.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 50, 74, figs. 4, 15, 27. 2. —Allen,
1969. Amer. Ent. Soc., Trans. 95: 433. ¢. —Allen, 1971. Amer. Ent. Soce., Trans. 97: 229,
figs. 23-27, map 1. 6, @.

frater Parker. Liberated in N. J. but not established. Introduced from China.
Tiphia frater Parker, 1935. N. Y. Ent. Soc., Jour. 43: 397. °, d.

fulvicauda Cameron. Nev. (Ormsby Co.).
Tiphia fulvicauda Cameron, 1907. Invertebrata Pacifica 1: 169. 2.

Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 24 (type redescription).

gehlsbachi Allen. Tex.

Tiphia gehlsbachi Allen, 1971. Amer. Ent. Soc., Trans. 97: 237, figs 32-34, map 2. ¢.
greenei Allen. Md. (Plummers Island).

Tiphia greenei Allen, 1966. Amer. Ent. Soc., Trans. 92: 256, fig. 31, map 3. ¢.
greeleyi Allen. Kans. (Greeley Co.).

Tiphia greeleyi Allen, 1971. Amer. Ent. Soc., Trans. 97: 258, map 5. 3.
illinoensis Robertson. N. H., Ont., Mich., Il]. and Iowa south to Fla. and Tex.

Tiphia illinoensis Robertson, 1901. Amer. Ent. Soce., Trans. 27: 196. 2, ¢.

Tiphia waldonii (!) Viereck, 1906. Ent. News 17: 302. 6.

Tphia brunneicornis Viereck, 1906. Ent. News 17: 303. 6.

Tiphia waldenii Viereck, 1906. Ent. News 17: 350.

Superfamily SCOLIOIDEA 1257

Tiphia robertsoni Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 23. °.
Tiphia waldeni (!) Malloch, 1918. Il. Nat. Hist. Survey, Bul. 13: 23.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 24. ¢. —Frison, 1927. Ill. Nat. Hist.
Survey, Bul. 16: 230 (lectotype designation, robertsoni). —Allen, 1934. Amer. Ent. Soc.,
Trans. 60: 313. —Allen, 1961. Amer. Ent. Soe., Trans. 87: 18. —Allen, 1966. Amer. Ent.
Soe., Trans. 92: 280, figs. 4, 27, 35, map 8. d, °.

inaequalis Malloch. Mass. to Nebr., south to Ga., Ala. and Tex.
Tiphia inaequalis Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 138: 22. d.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 229-230 (lectotype designation).
—Allen, 1966. Amer. Ent. Soc., Trans. 92: 293, fig. 29, map 10. ¢.

incisurata Malloch N. C.
Tiphia incisurata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 17. °.

Taxonomy: Allen, 1966. Amer. Ent.-Soc., Trans. 92: 288, map 9. °.

indistincta Allen. W. Va. (Kanawha Sta.).
Tiphia indistincta Allen, 1961. Amer. Ent. Soc., Trans. 87: 14. 2.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 254, map 3. 2.
infossata Allen. Maine to Mich., Wis., Iowa and S. Dak., south to Fla. and Tex., Alta., Mont.,
Colo., Calif.
Tiphia infossata Allen, 1934. Amer. Ent. Soc., Trans. 60: 301. °.
Tiphia hollowayi Allen, 1934. Amer. Ent. Soc., Trans. 60: 303. d.

Taxonomy: Allen, 1961. Amer. Ent. Soc., Trans. 87: 19 (synonymy). —Allen, 1965. Acad. Nat.
Sci., Phila., Proc. 117: 60. —Allen, 1966. Amer. Ent. Soc., Trans. 92: 248, figs. 8, 16, 21, 36,
map 2. 2, 6. —Allen, 1971. Amer. Ent. Soc., Trans. 97: 275, figs. 86-89, map 6. 2, d.

inornata Say. Ohio, Pa.
Tiphia inornata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 331 °.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 12. —Allen, 1934. Amer. Ent. Soc.,
Trans. 60: 300 (redescription.) — Allen, 1966. Amer. Ent. Soc., Trans. 92: 262. °.
(Redescription.)

intermedia Malloch. Maine and Que. south to Fla., west to N. Dak. and Calif., south to

Nicaragua. Host: Phyllophaga spp.

Tiphia punctata Robertson, 1901. Amer. Ent. Soe., Trans. 27: 196. d. Preoce.
Tiphia clypeolata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 16. 2, 3 (in part?).
Tiphia arida Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 20. 2.
Tiphia punctata var. intermedia Malloch, 1918. Ill. Nat. Hist. Survey Bul. 13: 21. 9.
Tiphia reticulata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 18: 23. 2.
Tiphia intermedia var. exitialis Roberts, 1930. Canad. Ent. 62: 189. N. name.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 230 (lectotype designation, punctata
var intermedia). —Allen, 1934. Amer. Ent. Soc., Trans. 60: 310. —Krombein, 1938. Ent.
News 49: 186. —Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 56. —Allen, 1966. Amer. Ent.
Soc., Trans. 92: 302, figs. 10, 12, 15, 17, 26, 37, 40, map 11. d, 2. —Allen, 1971. Amer. Ent.
Soe., Trans. 97: 268, figs. 69-72, map 6. d, °.

Biology: Davis, 1919. Ill. Nat. Hist. Survey, Bul. 13: 59-68.
irfla Allen. Fla.
Tiphia irfla Allen, 1961. Amer. Ent. Soc., Trans. 87: 10. d, °.
Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 314, map 12.
jaynesi Allen. Que. and Vt. to Fla., La., Okla., Kans., Ill.
Tiphia jaynesi Allen, 1934. Amer. Ent. Soc., Trans. 60: 311. 2, d.
Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 315, map 13. d, °.
krombeini Allen. N. H. and Ont. to N. C., W. Va., Mich., S. Dak., ? Tex.
Tiphia krombeini Allen, 1966. Amer. Ent. Soe., Trans. 92: 321, map 14. 2, d.
letalis Roberts. N. Y. to N. C., west to S. Dak. and Kans., Miss.
Tiphia clypeata Robertson, 1901. Amer. Ent. Soc., Trans. 27: 196. d. Preoce.

1258 | Hymenoptera in America North of Mexico

Tiphia letalis Roberts, 1930. Canad. Ent. 62: 189. N. name.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 11. 2, ¢. —Allen, 1966. Amer. Ent.
Soe., Trans. 92: 257, figs. 1, 3, 11, map 4. d, 9.
matura Allen and Jaynes. Liberated in N. J. but not established. Introduced from India. Host:
Popillia japonica Newm.
Tiphia matura Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art. 17: 81. 2, ¢.
micropunctata Allen. N.S. to Man., Mont. and Calif., south to Fla. and Okla.
Tiphia micropunctata Allen, 1934. Amer. Ent. Soc., Trans. 60: 297. 2, 6.
Tiphia dentata Allen, 1934. Amer. Ent. Soe. Trans. 60: 303. 3.

Taxonomy: Allen, 1961. Amer. Ent. Soc., Trans. 87: 18 (synonymy). —Evans, 1965. Ent. Soe.
Wash., Proc. 67: 90-91, figs. 6-10 (larva). —Allen, 1966. Amer. Ent. Soc., Trans. 92: 243,
figs. 7, 18, 22, 25, map 1. 2, 6.

minor Provancher. B. C. (Vancouver Isl.), ? Wash., ? Oreg.
Tiphia minor Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 254. 6.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 65. 6. —Allen, 1971. Amer. Ent.
Soc., Trans. 97: 294, map 8.
montana Allen. Sask., Alta., Mont., Utah, Ariz; Mexico (Jalisco).
Tiphia montana Allen, 1961. Amer. Ent. Soc., Trans. 87: 13. d.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proce. 117: 58, fig. 22. ¢. —Allen, 1971. Amer.
Ent. Soc., Trans. 97: 234, figs. 30, 31, map 2. d.
morio Fabricius. Liberated in N. Y. but not established. Introduced from Europe. Host:
Amphimallon majalis (Raz.).
Tiphia morio Fabricius, 1787. Mantissa Insectorum, v. 1, p. 280.
nebra Allen. Nebr., Ky.
Tiphia nebra Allen, 1966. Amer. Ent. Soc., Trans. 92: 261, map 3. °.
nevadana Cameron. B. C. south to Calif., Idaho, Wyo., Nev., Colo., Ariz., N. Mex., Tex; Mexico
(Hidalgo).
Tiphia nevadana Cameron, 1906. Invertebrata Pacifica 1: 157. 3.
Tiphia essigi Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 62, figs. 7, 9, 26. 2, 6.

Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 22 (type redescription). — Allen, 1971.
Amer. Ent. Soc., Trans. 97: 298, figs. 115, 116, map 9. 6, 2.
nona Allen. Ariz. south in Mexico to San Luis Potosi and Michoacan.
Tiphia nona Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 61. ¢.
Tiphia hurdi Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 75, fig. 24. d, 2.

Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 248, figs. 46-48, map 4. 6, 2.
notopolita var. alleni Roberts. Liberated in N. Y., but not established. Introduced from Korea
and China. Host: Anomala orientalis Waterh.
Tiphia notopolita var. intermedia Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art.
17: 41. 2. Preoce.
Tiphia notopolita var. alleni Roberts, 1930. Canad. Ent. 62: 190. N. name.
Biology: Clausen, Jaynes and Gardner, 1933. U.S. Dept. Agr., Tech. Bul. 366: 46 (life
history).
occidentata Malloch. Wash., Colo., N. Mex., Ariz. south in Mexico to Oaxaca.
Tiphia occidentata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 17. ¢.
Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 52. 6. —Allen, 1971. Amer. Ent.
Soc., Trans. 97: 241, figs. 38-40, map 3. 6.
odontogaster Viereck. N. Mex., Ariz.; Mexico (Durango).
Tiphia odontogaster Viereck, 1903. Amer. Ent. Soc., Trans. 29: 71. ¢.
Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 13. —Allen, 1965. Acad. Nat. Sci.
Phila., Proc. 117: 51, fig. 21. d. —Allen, 1971. Amer. Ent. Soc., Trans. 97: 232, figs. 28, 29,
map 2.

Superfamily SCOLIOIDEA 1259

pecosae Allen. Tex. (Pecos River at Sheffield).
Tiphia pecosae Allen, 1971. Amer. Ent. Soc., Trans. 97: 311, fig. 123. °.

pennsylvanica Allen. N. J. to Fla., La.
Tiphia pennsylvanica Allen, 1961. Amer. Ent. Soc., Trans. 87: 15. 2.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 286, map 9. 2, d.
popilliavora Rohwer. N. H., Mass., Conn., N. Y., N. J., Pa., Del., Md., Va., Ohio. Introduced
from Japan and Korea. Host: Popillia japonica Newm.
Tiphia popilliavora Rohwer, 1924. Ent. Soc. Wash., Proc. 26: 89. 2, d.
Taxonomy: Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art. 17: 51, figs. 3, 21, 22, 27,
Zone Ol
Biology: Clausen, King, and Teranishi, 1927. U. S. Dept. Agr., Bul. 1429: 33-39. —King and

Holloway, 1930. Jour. Econ. Ent. 23: 266-274. —Holloway, 1931. N. Y. Ent. Soc., Jour. 39:
555-564. —White, 1943. N. Y. Ent. Soc., Jour. 51: 213-218.

portalae Allen. Ariz., Nev.
Tiphia portalae Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 49. d.
Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 226, figs. 17, 18, map 1. 6, @.
pullivora Allen and Jaynes. Liberated in Pa. but not established. Introduced from India. Host:
Popillia japonica Newm.
Tiphia pullivora Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art. 17: 83. 2, d.
Biology: Clausen, Jaynes and Gardner, 1933. U. S. Dept. Agr., Tech. Bul. 366: 39-41 (life
history).
puncticeps Cameron Nev.; ? Mexico (Michoacan).
Tiphia puncticeps Cameron, 1906. Invertebrata Pacifica 1: 158. ¢.
Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 23 (type redescription). — Allen, 1971.
Amer. Ent. Soc., Trans. 97: 246, map 4. ¢.
pygidialis Allen. Conn., Pa., Md., W. Va., Ky., Iowa, Kans., Okla., Miss.
Tiphia pygidialis Allen, 1966. Amer. Ent. Soc., Trans. 92: 310, map 12. 2.
raui Allen. Mo., Miss.
Tiphia raui Allen, 1934. Amer. Ent. Soc., Trans. 60: 308. 2.
Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 269. 2.
relativa Viereck. Maine to Ga., Mich., Iowa, Nebr., Kans. Host: Cyclocephala borealis Arrow;
Lichnathe vulpina (Hentz).
Tiphia relativa Viereck, 1906. Ent. News 17: 304. 6.
Tiphia winnemanae Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 23. d.
Taxonomy: Allen, 1961. Amer. Ent. Soc., Trans. 87: 17 (synonymy). —Allen, 1966. Amer. Ent.
Soe., Trans. 92: 312, map 12. 6, 92.
Biology: Adams, 1949. Jour. Econ. Ent. 42: 626. —Franklin, 1950. Mass. Agr. Expt. Sta., Bul.
445, pts. 2-7: 63, 66, 67.
roda Allen. N. Mex., Ariz.
Tiphia roda Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 56. d.
Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 286, map 8. ¢.
rodeoensis Allen. N. Mex. south in Mexico to Jalisco and Morelos.
Tiphia rodeoensis Allen, 1971. Amer. Ent. Soc., Trans. 97: 336, fig. 148, map 13. @.
roosevelti Allen. Ariz.
Tiphia roosevelti Allen, 1971. Amer. Ent. Soc., Trans. 97: 339, figs. 145-147, map 13. °.
rothi Allen. Ariz. (Cochise Co.).
Tiphia rothi Allen, 1971. Amer. Ent. Soc., Trans. 97: 231, map 1. 6.
rugulosa Malloch. Maine and Que. to Ga., west to Minn., Iowa and Kans.
Tiphia rugulosa Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 15. 9, ¢.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 230 (lectotype designation). — Allen,
1966. Amer. Ent. Soc., Trans. 92: 333, fig. 30, map 16. 6, °.

1260 | Hymenoptera in America North of Mexico

sayi Allen. N. H., Ont., Wis. and Iowa south to Fla. and Miss.
Tiphia sayi Allen, 1966. Amer. Ent. Soc., Trans. 92: 272, fig. 18, map 6. d, °.

schlingeri Allen. Tex., N. Mex., Ariz., Nev., Calif.
Tiphia schlingeri Allen, 1965. Acad. Nat. Sci. Phila., Proce. 117: 64, fig. 12. 6.
Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 311, map 10. d.
sculleni Allen. Ariz. (Cochise and Pima Counties).
Tiphia sculleni Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 60, fig. 25. ¢.
Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 291, figs. 104, 105, map 8.
similis Malloch. Maine and Que. south to Fla., west to Ont., Mich., Ill. and Kans.

° Tiphia similis Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 18. ¢.
Tiphia affinis Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 19. 2, d.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 229-230 (lectotype designations,
similis, affinis). —Allen, 1966. Amer. Ent. Soc., Trans. 92: 318, figs. 20, 28, map 14. 6, @.
sinaloae Allen. Okla., Tex., south to Costa Rica.
Tiphia sinaloae Allen, 1971. Amer. Ent. Soc., Trans. 97: 309, map 10. 6.
sternata Parker. Liberated 1933-35 in N. Y., N. J., Pa. but has not been recovered since 1937.
Introduced from Japan. Host: Serica peregrina Chapin.
Tiphia sternata Parker, 1935. N. Y. Ent. Soc., Jour. 43: 399. 2, ¢.
strangulata Allen. Mont., Ariz.; ? Mexico (Puebla).
Tiphia strangulata Allen, 1971. Amer. Ent. Soc., Trans. 97: 261, fig. 73, map 5. d.
subcarinata Malloch. Mass. to Ga., W. Va., Mich., IIl.
Tiphia subcarinata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 15. ¢.
Tiphia insolita Allen, 1934. Amer. Ent. Soc., Trans. 60: 316. d.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 330, map 16. 6.

tarda Say. Ind.
Tiphia tarda Say, 1836. Boston Jour. Nat. Hist. 1: 300. ¢.

Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 351 (unrecognized).
tegulina Malloch. B. C. and Idaho to Calif. and Nev. Host: Phyllophaga errans LeC.
Tiphia tegulina Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 21. ?.
Tiphia shastensis Krombein, 1942. Pan-Pacific Ent. 18: 139. 2, 3.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 59. —Allen, 1971. Amer. Ent. Soc.,
Trans. 97: 272, figs. 79-81, map 6. °, 6.

Biology: Linsley and Michener, 1942. Pan-Pacifie Ent. 18: 154.
tempeae Allen. Kans., Colo., Wash., Calif., Ariz. and N. Mex. south to Costa Rica.
Tiphia tempeae Allen, 1961. Amer. Ent. Soc., Trans. 87: 11. d.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 53, 80. d. —Allen, 1971. Amer. Ent.
Soc., Trans. 97: 263, fig. 74, map 5. d, 9.

texensis Malloch. Tex., Kans., Mont.
Tiphia texensis Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 20. °.

Taxonomy: Allen, 1961. Amer. Ent. Soc., Trans. 87: 19. —Allen, 1965. Acad. Nat. Sci. Phila.,
Procsli(eiGos ee
toddi Allen. Ariz.; Mexico (Jalisco, Morelos).
Tiphia toddi Allen, 1971. Amer. Ent. Soc., Trans. 97: 252, fig. 50, map 4. d.
totopunctata Allen and Jaynes. Liberated in N. Y. but not established. Introduced from
Korea. Host: Anomala orientalis Waterh.
Tiphia totopunctata Allen and Jaynes, 1930. U.S. Natl. Mus., Proc. 76, Art. 17: 35. 2.

Biology: Clausen, Jaynes and Gardner, 1933. U. S. Dept. Agr., Tech. Bul. 366: 44 (life
history).

townesi Allen. N. Y. (Shokan).
Tiphia townesi Allen, 1966. Amer. Ent. Soc., Trans. 92: 338, map 17. d.

Superfamily SCOLIOIDEA 1261

transversa Say. Vt. to Kans., south to Ga., Ala. and Tex. Host: Phyllophaga spp.
Tiphia transversa Say, 1828. Contrib. Maclur. Lye. Phila. 1: 83 ¢.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 11. —Allen, 1934. Amer. Ent. Soc.,
Trans. 60: 297. —Allen, 1966. Amer. Ent. Soc., Trans. 92: 270, fig. 19, map 6. d, °.
Biology: Davis, 1919 Ill. Nat. Hist. Survey, Bul. 13: 68.

treherni Allen. B. C. to Calif.
Tiphia treherni Allen, 1971. Amer. Ent. Soc., Trans. 97: 329, figs. 35, 36, map 12. °.

truncata Cameron. Nev. (Ormsby Co.).
Tiphia truncata Cameron, 1906. Invertebrata Pacifica 1: 156. d.
Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 22 (type unlocated).
unica Allen. N. C. to Fla., Ala., Tenn., Kans.
Tiphia unica Allen, 1934. Amer. Ent. Soc., Trans. 60: 309. °.
Taxonomy: Allen, 1966. Amer. Ent. Soc., Trans. 92: 267, fig. 2, map 5. 2, d.

uvaldae Allen. Tex. (Uvalde Co.).
Tiphia uvaldae Allen, 1971. Amer. Ent. Soc., Trans. 97: 313, fig. 124, map 11. °.

vallis Allen. Calif. (Kern and El] Dorado Counties).
Tiphia vallis Allen, 1971. Amer. Ent. Soc., Trans. 97: 334, map 12. °.

veracruzae Allen. Southern Ariz. to Panama.
Tiphia veracruzae Allen, 1965. Acad. Nat. Sci. Phila., Proe. 117: 82, fig. 5. d, °.
Tiphia litoris Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 84. d, °.
Taxonomy: Allen, 1971. Amer. Ent. Soc., Trans. 97: 295, figs. 108-114, map 9. d, °.

vernalis Rohwer. Mass., N. H., R. I., Conn., N. Y., N. J., Pa., Del., Md., D. C., Va., N. C., W. Va.,
Ohio; Orient. Introduced from Japan, Korea, and China. Host: Popillia japonica Newm.
Tiphia vernalis Rohwer, 1924. Ent. Soc. Wash., Proc. 26: 91. 2, d.

Taxonomy: Allen and Jaynes, 1930. U. S. Natl. Mus., Proc. 76, Art. 17: 78, figs. 6, 16, 19, 25,
29.

Biology: Clausen, King, and Teranishi, 1927. U. S. Dept. Agr., Bul. 1429: 40-41. —Balock,
1934. Jour. Econ. Ent. 27: 491-496. —White, 1943. N. Y. Ent. Soc., Jour. 51: 213-218.

virotha Allen. Ariz. (Cochise Co.); ? Mexico (Hidalgo).
Tiphia virotha Allen, 1971. Amer. Ent. Soc., Trans. 97: 338, map 13. °.

vulgaris Robertson. IIl., Wis., Ind., lowa, S. Dak., Kans., Mo., Tex. Host: Phyllophaga spp.
Tiphia vulgaris Robertson, 1901. Amer. Ent. Soc., Trans. 27: 195. 2, 6.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 12. —Allen, 1934. Amer. Ent. Soc.,
Trans. 60: 305. —Allen, 1961. Amer. Ent. Soc., Trans. 87: 19. —Allen, 1966. Amer. Ent.
Soc., Trans. 92: 263, fig. 39, map 5. d, °.

Biology: Davis, 1919. Ill. Nat. Hist. Survey, Bul. 13: 69.

yosemitensis Allen. Nebr., Mont., Calif.
Tiphia yosemitensis Allen, 1971. Amer. Ent. Soc., Trans. 97: 330, figs. 187-139, map 12. °.

Genus KROMBEINIA Pate
Neotiphia subg. Krombeinia Pate, 1947. N. Y. Ent. Soc., Jour. 55: 182.
Type-species: Neotiphia chiricahua Pate. Orig. desig.

Revision: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 211-241.

chiricahua (Pate). Ariz. (Chiricahua).
Neotiphia chiricahua Pate, 1939. Ent. News 50: 221. d, °.

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 233. d, °.

cristata (Allen). Ariz., N. Mex.
Neotiphia cristata Allen, 1935. Amer. Ent. Soc., Trans. 61: 59. 2.

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 238, figs. 3, 29. 2.

1262 Hymenoptera in America North of Mexico

jonesi Allen and Krombein. Ariz. (Douglas, Chiricahua, Huachuca Mts., Oracle).
Krombeinia jonesi Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 239, figs. 6, 65.
Onna
robusta (Allen). Tex.
Neotiphia ocellata Allen, 1935. Amer. Ent. Soc., Trans. 61: 56. 9°.
Neotiphia robusta Allen, 1935. Amer. Ent. Soc., Trans. 61: 64. 3.

Taxonomy: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 739 (synonymy). —Allen and
Krombein, 1964. Amer. Ent. Soc., Trans. 89: 229, figs. 2, 30. 6, 2.

Genus NEOTIPHIA Malloch

Neotiphia Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 9.
Type-species: Neotiphia acuta Malloch. Orig. desig.

Revision: Allen, 1935. Amer. Ent. Soc., Trans. 61: 53-65. —Allen and Krombein, 1964. Amer.
Ent. Soc., Trans. 89: 211-214, 241-270.
acuta Malloch. Tex.
Neotiphia acuta Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 9. 2, d.

Taxonomy: Frison, 1927. Ill. Nat. Hist. Survey, Bul. 16: 229 (lectotype designation). — Allen

and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 259, figs. 5, 46, 67. 3.
anima Allen and Krombein. N. Mex., Ariz.

Neotiphia anima Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 262. °.
barbata Allen and Krombein. Ariz. (Continental, Tucson, Douglas).

Neotiphia barbata Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 264. d.
cockerelli Allen. Tex., N. Mex.

Neotiphia cockerelli Allen, 1935. Amer. Ent. Soc., Trans. 61: 62. 3.

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 255, fig. 50. d, 9.
crawfordi Allen. Tex.
Neotiphia crawfordi Allen, 1935. Amer. Ent. Soc., Trans. 61: 60. 9.
Neotiphia conspicua Allen, 1935. Amer. Ent. Soc., Trans. 61: 61. 3.

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 257, figs. 28, 41, 42. ?, d.
flavipennis Allen and Krombein. Ariz., N. Mex. south in Mexico to Mexico and Hidalgo.
Neotiphia flavipennis Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 263. 9.

Neotiphia parma Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 267, figs. 7, 54. ¢.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 87 (synonymy).

luteipennis (Cresson). Colo.
Tiphia luteipennis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 445. 9.

Taxonomy: Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 10. —Allen and Krombein, 1964.
Amer. Ent. Soc., Trans. 89: 260. °.
novomexicana Allen. N. Mex., Ariz.
Neotiphia novomexicana Allen, 1935. Amer. Ent. Soc., Trans. 61: 63. 9°.
Neotiphia carinata Krombein, 1938. Ent. Soc. Amer., Ann. 31: 59. d.

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 251, figs. 4, 9, 11, 18, 21,
25, 38, 49, 55, 60, 68. 2, d.
rioverdei Allen and Krombein. Ariz. (Del Rio Verde).
Neotiphia rioverdei Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 263. °.
rostrata Allen. Ariz.
Neotiphia rostrata Allen 1935. Amer. Ent. Soc., Trans. 61: 64. 3.

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 265, figs. 14, 53. 3.

sulcata (Roberts). Mich., lowa and S. Dak. south to Tex. and Ariz.; Mexico (Jalisco).
Tiphia canaliculata Malloch, 1918. Ill. Nat. Hist. Survey, Bul. 13: 10. 6. Preoce.
Tiphia sulcata Roberts, 1930. Canad. Ent. 62: 190. N. name.
Tiphia mexicana Allen, 1935. Amer. Ent. Soc., Trans. 61: 57. 9.

Superfamily SCOLIOIDEA 1263

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soc., Trans. 89: 245, figs. 1, 8, 12, 17, 20,
23, 24, 27, 34, 44, 48, 56, 58, 59, 66. d, °.
waltoni Allen. Kans. and Colo. to Tex., N. Mex. and Ariz.; Mexico (Chihuahua, Durango).
Neotiphia waltoni Allen 1935. Amer. Ent. Soc., Trans. 61: 62. °.
Neotiphia pima Pate, 1939. Ent. News 50: 245. 3, (2 misdet.).

Taxonomy: Allen and Krombein, 1964. Amer. Ent. Soe., Trans. 89: 248, 254, fig. 15, 36, 37, 47,
62. 2, 3 (treat waltoni and pima as discrete species). —Allen, 1965. Acad. Nat. Sci. Phila.,
Proce. 117: 87 (synonymy).

Genus PARATIPHIA Sichel

Epomidiopteron subg. Paratiphia Sichel, 1864. In Saussure and Sichel, Cat. Spec. Gen.
Scol., p. 269.
Type-species: Epomidiopteron Sumichrasti Sichel. Monotypic.

Revision: Allen, 1968. Amer. Ent. Soc., Trans. 94: 25-109, 41 figs., 10 maps.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 89-107 (species of western U. S.).
aequalis atrata Allen. Calif., Ariz., Tex. south to El. Salvador. Typical aequalis Fox occurs in
Baja California.
Paratiphia aequalis atrata Allen, 1963. Amer. Ent. Soc., Trans. 88: 228, figs. 5, 10, 15, 16.
Onmee

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 90, figs. 3, 20, 30. d, 2. —Allen,
1968. Amer. Ent. Soc., Trans. 94: 33, figs. 1-3, map 1. d, °.
albilabris (Spinola). Calif.
Tiphia albilabris Spinola, 1841. Soc. Ent. France, Ann. 10: 102. d.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 107 (type not located).

asotinae Allen. Wash. and Nebr. south to Calif., Kans. and N. Mex.
Paratiphia asotinae Allen, 1963. Ent. News 74: 222. d, °.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 57, map 4. d, °.
belfragei Allen. Nebr. and Idaho to Tex. and Calif., south in Mexico to Hidalgo and Jalisco.
Paratiphia belfragei Allen, 1963. Ent. Soc. Amer., Ann. 56: 575, figs. 1, 2 ¢, daGrase
Paratiphia insueta Allen, 1963. Ent. Soc. Amer., Ann. 56: 576, figs. 1,2 e. d.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 66, figs. 21, 23, 24, map 6. d, °.

bridwelli Allen. Kans., Tex., Ariz.; Mexico (Tamaulipas).
Paratiphia bridwelli Allen, 1963. Amer. Ent. Soc., Trans. 88: 231, figs. 1, 6, 11, 19. d.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 44, map 2. 6.

cincta Allen. Calif., Nev., Ariz.; Mexico (Baja California).
Paratiphia cincta Allen, 1962. Ent. News 73: 262. d.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 98, 106, fig. 2. ¢. —Allen, 1968.
Amer. Ent. Soc., Trans. 94: 78, figs. 13, 34, map 5. d, °.
bradleyi Allen. Tex. to Ariz., south in Mexico to Baja California and Veracruz.
Paratiphia bradleyi Allen, 1963. Amer. Ent. Soc., Trans. 88: 231, figs. 2, 7, 12. d.
Paratiphia pachucae Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 101, fig. 17. ¢.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 49, map 3. 6, 2. —Betrem and Allen,

1970. Ent. Soc. Wash., Proc. 72: 71.

claripennis Cameron. B. C., Idaho, Nev., Calif. to Tex., south in Mexico to Baja California,

Sinaloa and Puebla.

Paratiphia claripennis Cameron, 1905. Invertebrata Pacifica 1: 108. d.
Paratiphia varipunctata Cameron, 1905. Invertebrata Pacifica 1: 106. 2.
Paratiphia fuscinerva Cameron, 1905. Invertebrata Pacifica 1: 107. °.
Paratiphia bakeri Cameron, 1905. Invertebrata Pacifica 1: 109. 6.
Paratiphia ormsbyensis Cameron, 1907. Invertebrata Pacifica 1: 175. d.

1264 Hymenoptera in America North of Mexico

Taxonomy: Cameron, 1907. Invertebrata Pacifica 1: 172. 2. —Allen, 1962. Amer. Ent. Soc.,
Trans. 88: 29-30. —Allen, 1965. Acad. Nat. Sci. Phila., Proe. 117: 96. ¢. —Allen, 1968. Amer.
Ent. Soe., Trans. 94: 75, fig. 31, map 7. d.

dreisbachi Allen. Colo., N. Mex.
Paratiphia dreisbachi Allen, 1968. Amer. Ent. Soc., Trans. 94: 83, fig. 32, map 6. °.
ephippiata Allen. Kans. and Wyo. to Tex. and Ariz. south in Mexico to Zacatecas and San Luis
Potosi.
Paratiphia ephippiata Allen, 1963. Ent. Soe. Amer., Ann. 56: 574, figs. 1, 2 a, b, f. d, 2.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 93, fig. 19. ¢, 2. —Allen, 1968.

Amer. Ent. Soc., Trans. 94: 50, figs. 6-10, map 3. 3, @.
fossata Allen. N. Mex. and Ariz. south in Mexico to Veracruz and Oaxaca.
Paratiphia fossata Allen, 1962. Ent. News 73: 259. 6.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 98, 103. ¢. —Allen, 1968. Amer. Ent.

Soc., Trans. 94: 64, figs. 18-20, map 4. ¢, 9.
halli Allen. Calif. (Riverside Co.).

Paratiphia halli Allen, 1968. Amer. Ent. Soc., Trans. 94: 68, map 6. 6.
huachucae Allen. Ariz. (Huachuca Mts.).

Paratiphia huachucae Allen, 1968. Amer. Ent. Soc., Trans. 94: 47, map 2. 6.
irwini Allen. Ariz. (Chiricahua Mts. ).

Paratiphia irwini Allen, 1968. Amer. Ent. Soc., Trans. 94: 74, map 6. ¢.
knowltoni Allen. Utah., Nev.

Paratiphia knowltoni Allen, 1968. Amer. Ent. Soc., Trans. 94: 82, fig. 35, map 6. °.
knulli Allen. Tex.

Paratiphia knulli Allen, 1968. Amer. Ent. Soc., Trans. 94: 60, fig. 16, map 5. ¢.
lagosae Allen. Tex. and Calif. south in Mexico to Michoacan and Hidalgo.

Paratiphia lagosae Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 101. 3.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 37, map 1. 6, °.

magna Roberts. Idaho (Pocatello).
Paratiphia magna Roberts, 1929. Psyche 36: 358. 9.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 84, map 6. °.

mitchelli Allen. Tex. to Ariz. south in Mexico to Puebla.
Paratiphia mitchelli Allen, 1963. Amer. Ent. Soc., Trans. 88: 232, figs. 3, 8, 13, 18. d.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 46, map 2. d, °.

neomexicana Cameron. Oreg. to Nebr. south to Calif. and Tex.; Mexico (Sonora, Chihuahua).
Paratiphia neomexicana Cameron, 1907. Invertebrata Pacifica 1: 175. ¢.

Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 31 (type redescription). —Allen, 1965.
Acad. Nat. Sci. Phila., Proc. 117: 94. d, 2. —Allen, 1968. Amer. Ent. Soc., Trans. 94: 54,
figs. 11, 12, 14, map 4. d, 9.

nevadensis Cameron. B. C. to Calif. and Tex., south in Mexico to Baja California and Puebla.
Paratiphia nevadensis Cameron, 1905. Invertebrata Pacifica 1: 108. 3.
Paratiphia parvula Cameron, 1907. Invertebrata Pacifica 1: 176. d.
Paratiphia intermedia Cameron, 1907. Invertebrata Pacifica 1: 176. ¢.

Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 27, 32 (synonymy, type redescription).
—Allen, 1963. Amer. Ent. Soc., Trans. 88: 229, figs. 4, 9, 14, 17 (redescription). —Allen,
1965. Acad. Nat. Sci. Phila., Proe. 117: 91, 95. d, 2. —Allen, 1968. Amer. Ent. Soc., Trans.
94: 39, fig. 5, map 2. d, 9.

occidentalis Cameron. Nev. (Ormsby Co.).
Paratiphia occidentalis Cameron, 1907. Invertebrata Pacifica 1: 173. ¢.

Taxonomy: Allen, 1968. Amer. Ent. Soc., Trans. 94: 108 (type not located).

robusta Cameron. B. C., Idaho and Nebr. to Calif. and Tex.; Mexico (Baja California, Nayarit).
Paratiphia robusta Cameron, 1905. Invertebrata Pacifica 1: 106. 2.
Paratiphia fortistriolata Cameron, 1907. Invertebrata Pacifica 1: 172. ¢.

Superfamily SCOLIOIDEA 1265

Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 33 (type redescribed, synonymy). —Allen,

1965. Acad. Nat. Sci. Phila., Proc. 117: 92. 9, 6. —Allen, 1968. Amer. Ent. Soc., Trans. 94:
70, figs. 22, 25-30, 41, map 6. 9, d.

sculleni Allen. Oreg., Utah, Ariz., N. Mex.
Paratiphia sculleni Allen, 1968. Amer. Ent. Soc., Trans. 94: 52, map 3. d, °.

texana Cameron. Ont. and Mass. to Mont. south to Fla. and Tex.
Paratiphia tecana Cameron, 1907. Invertebrata Pacifica 1: 174. 2, d.
Paratiphia algonguina Viereck, 1909. Ent. Soc. Wash., Proce. 11: 45. 2, d.
Paratiphia algonquina (!) Smith, 1910. N. J. State Mus., Ann. Rpt. for 1909, p. 666.

Taxonomy: Allen, 1962. Amer. Ent. Soc., Trans. 88: 25 (redescription texana). —Allen, 1965.
Acad. Nat. Sci. Phila., Proc. 117: 97. 2, 6. —Allen, 1968. Amer. Ent. Soc., Trans. 94: 62,
figs. 15, 17, map 5. d, 9.

verna Allen. Calif., Nev., Ariz.; Mexico (Baja California).
Paratiphia verna Allen, 1963. Ent. News 74: 219. 6, °.

Taxonomy: Allen, 1965. Acad. Nat. Sci. Phila., Proc. 117: 99. ¢. —Allen, 1968. Amer. Ent.
Soc., Trans. 94: 80, fig. 36, map 7. d, °.

UNPLACED TAXON OF PARATIPHIA SICHEL

fuscipennis Cameron. No locality given (title says n. spp. from “Nevada and Southern

California”).
Paratiphia fuscipennis Cameron. 1905. Invertebrata Pacifica 1: 106. 2 (in key).

Genus EPOMIDIOPTERON Romand

Epomidiopteron Romand, 1835. Soc. Ent. France, Ann. 4: 653.
Type-species: Epomidiopteron Julii Romand. Monotypic.

Scoliphia Banks, 1912. Canad. Ent. 44: 201.
Type-species: Scoliphia spilota Banks. Orig. desig.

Epimodiopteron (!) Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 119.

julii Romand. Southern Ariz. to Peru and Argentina.

Epomidiopteron Julii Romand, 1836. Soc. Ent. France, Ann. 4: 653. °.

Paratiphia 12-maculata Cameron, 1904. Amer. Ent. Soc., Trans. 30: 94. ¢.

Scoliphia spilota Banks, 1912. Canad. Ent. 44: 201. 9, d.

Epimodiopteron (!) spilota Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 119.

Taxonomy: Allen, 1966. Ent. News 77: 206, 2 figs. 2, ¢ (synonymy and redescription).
—Allen, 1972. Smithsn. Contrib. Zool. 113: 3.

SUBFAMILY MYZININAE
Revision: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 227-292, 12 figs.
Genus MYZINUM Latreille

Myzinum Latreille, 1803. Nouv. Dict. Hist. Nat., v. 15, p. 326.
Type-species: Tiphia maculata Fabricius. Monotypic.
Elis Fabricius, 1805. Systema Piezatorum, p. 248.
Type-species: Elis sexcincta Fabricius. Desig. by Bingham, 1897.
Plesia Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 150.
Type-species: Tiphia namea F'abricius. Desig. by Ashmead, 1903.
Myzina (!) Rafinesque, 1815. Analyse Nat., p. 124.

In describing Myzinum as new, Latreille gave a vernacular as well as a scientific name, thus,
myzine, Myzinum. Later, he and most subsequent authors incorrectly used this vernacular
name rather than the properly formed scientific name.

Two North American species have been recorded as parasitizing scarabaeid larvae in the soil,
a host relationship which has been recorded also for several Neotropical species.

Taxonomy: Pate, 1935. Ent. News 46: 265-267 (nomenclature). —Krombein, 1937. Ent. Soc.
Amer., Ann. 30: 26-30 (key to world genera).

1266 Hymenoptera in America North of Mexico

berlyi berlyi (Brimley). S. C., Ill., Ky., Wis., Kans., Miss., Tex.
Elis berlyi Brimley, 1927. Ent. News 38: 238. 2.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 249-251. 2, ¢.

berlyi parksi Krombein. La., Tex.

Myzine (!) berlyi parksi Krombein, 1938. Amer. Ent. Soc., Trans. 64: 251. 2, d.
berlyi patei Krombein. N. J., N. C., Ga., Fla.

Myzine (!) berlyi patei Krombein, 1938. Amer. Ent. Soc., Trans. 64: 254. 2, d.
carolinianum carolinianum (Panzer). Va. to Fla., west to Tex.

Tiphia caroliniana Panzer, 1806. Krit. Revis. Insektenf. Deutschlands, v. 2, p. i, pl. 1, figs.

a-c. 2.
Myzine (!) flavipes Olivier, 1811. Eneycl. Meth., v. 8, p. 136. 2.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 260-262, fig. 7. 2, ¢.
carolinianum collare (Say). Ohio west to S. Dak., south to La. and N. Mex.
Meria collaris Say, 1837. Boston Jour. Nat. Hist. 1: 362. °.
Myzine (!) thoracica Fox, 1893. Canad. Ent. 35: 113. 2. Preoce.
Myzine (!) illinoisensis Dalla Torre, 1897. Cat. Hym., v. 8, p. 124. N. name.
Elis atriventris Gahan, 1913. U. S. Natl. Mus., Proc. 40: 431. @, 6.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 262-265. 9, 6.

confluens Cresson. Utah, Ariz., south in western Mexico to Oaxaca.
Myzine (!) conflwens Cresson, 1865. Ent. Soc. Phila., Proc. 4: 443. °.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 287-288. ?.
dubiosum Cresson. D. C. south to Fla., Ala., Ill. south to La., Colo., Ariz., south in Mexico to
Veracruz, Sonora and Baja California.
Myzine (!) dubiosa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 200. 9.
Myzine (!) tecana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 200. ?.
Plesia fulvinervis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 239. 9.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 281-284, figs. 1, 10. 9, d.
frontale Cresson. Kans., Tex., N. Mex., Colo., Ariz., Nev., Calif., south in Mexico to Baja
California, Sonora and Chihuahua.
Plesia marginata Say, 1823. West. Quart. Rptr. 2: 75. 2. Preoce.
Myzine (!) frontalis Cresson, 1875. Rpt. Geog. Geol. Expl. and Survey west of 100th
Meridian, p. 711. 9.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 285-287, fig. 9. 2, d.
maculatum (Fabricius). Mass. to Fla., west to Nev. and Ariz., Calif. south to Guatemala and El
Salvador.
Tiphia maculata Fabricius. 1793. Ent. System., p. 224. 9.
Sapyga majorta Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. iv, pl. 2, figs. d,
e. 3.
Tiphia interrupta Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 322. 9.
Myzine (!) hamatus Say, 1836. Boston Jour. Nat. Hist. 1: 300. ¢.
Meria costata Say, 1837. Boston Jour. Nat. Hist. 1: 360. 2.
Myzine (!) hyalina Cresson, 1865. Ent. Soc. Phila., Proc. 4: 442. ¢.
Myzine (!) maiorta (!) Cresson, 1887. Amer. Ent. Soc., Trans., Sup. Vol., p. 268.
Elis floridanus Rohwer, 1920. Ent. Soc. Wash., Proce. 22: 54. 2, ¢.
Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 274-281, fig. 11. 2, ¢. —Krombein,
1949. U. S. Natl. Mus., Proc. 100: 55-56, figs. 1-3 (gynandromorph).
marginatum (Klug). Ga.
Tiphia marginata Klug, 1810. In Weber, Beitr. z. Naturk. 2: 184. ¢.
namea fulviceps Cameron. South. Ariz. to Guatemala.
Myzine (!) fulviceps Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 248, pl. 12, fig. 20. °.

namea namea (Fabricius). Va., N. C., S. C., Fla., Ala., Tex.
Tiphia namea Fabricius, 1805. Systema Piezatorum, p. 233. °.
Elis propodealis Rohwer, 1920. Ent. Soc. Wash., Proc. 22: 56. 2.

Superfamily SCOLIOIDEA 1267

Myzine (!) magna Krombein, 1938. Amer. Ent. Soc., Trans. 64: 256. d.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 256-259, fi F001). e-

navajo Krombein. Ariz. south to E] Salvador.

Myzine (!) navajo Krombein, 1938. Amer. Ent. Soc., Trans. 64: 272. °, OE
obscurum (Fabricius). N. Y. to Fla., west to Ill. and Tex., ? Ariz. Host: Dyscinetus

trachypygus (Burm.). Parasite: Dasymutilla mutata (B1.).

Tiphia obscura Fabricius, 1805. Systema Piezatorum, p. 233. 2.

Plesia nigripes Guerin, 1838. Rev. Zool. 1: 57. 9.

Plesia fuliginosa Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 574. d.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 246-249, fig. 3. 2, d.
quinquecinctum (Fabricius). Que. to Fla., west to Wyo. and N. Mex., Calif.; Mexico (Coahuila).
Host: Phyllophaga spp. Parasite: Dasymutilla quadriguttata (Say).
Tiphia 5-cincta Fabricius, 1775. Systema Ent., p. 358. °.
Tiphia cingulata Klug, 1810. In Weber, Beitr. z. Naturk. 2: 185. ¢.
Sapyga subulata Say, 1823. West. Quart. Rptr. 2: 75. d.
Myzine (!) proxima Guerin, 1837. Dict. Pitt. Hist. Nat., p. 581. d.

Taxonomy: Krombein, 1938. Amer. Ent. Soe., Trans. 64: 265-272, fig. 6. 2, d. —Evans, 1965.
Ent. Soc. Wash., Proce. 67: 92, figs. 11-14 (larva).

Biology: Flint and Sanders, 1912. Jour. Econ. Ent. 3: 490. —Davis, 1919. Ill. Nat. Hist.
Survey, Bul. 13: 73.

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99, no. 14, p. 44, pl. 14, figs. M-S
(male genitalia).
serenum (Fabricius). S. C.
Tiphia serena Fabricius, 1805. Systema Piezatorum, p. 234. °.
spilonotum (Cameron). Tex., N. Mex., Colo., Utah, Ariz., south in western Mexico to Oaxaca.
Plesia spilonota Cameron, 1908. Amer. Ent. Soc., Trans. 34: 240. °.

Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 288-290, fig. 12. 2, d.

Genus PTEROMBRUS Smith

Pterombrus Smith, 1869. Ent. Soc. London, Trans., p. 302.
Type-species: Pterombrus aenigmaticus Smith. Monotypic.
Engycystis Fox, 1895. Calif. Acad. Sci., Proc. 4: 262.
Type-species: Myzine (!) rufiventris Cresson. Monotypic.
Huberia Ducke, 1907. Rev. d’Ent. 26: 5.
Type-species: Huberia glabricollis Ducke. Desig. by Krombein, 1937.
Pterombus (!) Williams, 1928. Hawaii. Sugar Planters’ Assoc. Expt. Sta., Bul. Ent. Ser. 19:
144.

Nothing is known of the habits of our sole North American species, but some of the South
American species are parasitic on larvae of two genera of Cicindelidae (Coleoptera).
rufiventris hyalinatus Krombein. West. Tex., Ariz., Calif.
Pterombrus rufiventris hyalinatus Krombein, 1949. Pan-Pacifie Ent. 25: 88. 2, d.
rufiventris rufiventris (Cresson). Va. south to Ga. to east. Tex.
Myzine rufiventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 201. d.
Myzine cressonii Dalla Torre, 1897. Cat. Hym., v. 8, p. 122. N. name for Myzine rufiventris
Cresson, not Meria rufiventris Klug.
Taxonomy: Krombein, 1938. Amer. Ent. Soc., Trans. 64: 236-238, fig. 2. 2, 6. —Krombein,
1949. Pan-Pacifie Ent. 25: 88.

SUBFAMILY ANTHOBOSCINAE

Lalapa Pate, 1947. N. Y. Ent. Soc., Jour. 55: 126.
Type-species: Lalapa lusa Pate. Orig. desig.

1268 | Hymenoptera in America North of Mexico

lusa Pate. Idaho, Wash. to Calif.
Lalapa lusa Pate, 1947. N. Y. Ent. Soc., Jour. 55: 128. 2, 6.

SUBFAMILY BRACHYCISTIDINAE

Brachycistidinae exhibit very pronounced sexual dimorphism, males being fully winged and
females wingless and with modified thorax. Positive sex associations have been established only
in Brachycistis Fox. Eventually there will be considerable generic synonymy because the
remaining males and females are assigned to separate genera.

Revision: Bradley, 1917. Amer. Ent. Soc., Trans. 43: 252-283, pls. 20-25 (males only).
—Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 1-28, pls. 1-4 (males only). —Mickel and
Krombein, 1942. Amer. Midland Nat. 28: 648-679, pls. 1-3 (females only).

Taxonomy: Wasbauer, 1966. Calif. Univ. Pubs. Ent. 43: 15 (key to male genera). —Wasbauer,
1968. Pan-Pacific Ent. 44: 196-197 (key to male genera).

Biology: Rozen, 1952. Pan-Pacifiec Ent. 23: 91-92 (collecting techniques for females).

Genus BRACHYCISTIS Fox

Brachycistis Fox, 1893. Calif. Acad. Sci., Proc. 4 (ser. 2): 7.
Type-species: Brachycistis petiolatus Fox. Orig. desig.
Brachycistus (!) Cockerell and Casad, 1894. Ent. News 5: 295.
Glyptometopa Ashmead, 1898. Psyche 8: 251.
Type-species: Glyptometopa americana Ashmead. Orig. desig.
Glytometopa (!) Fox, 1899. Amer. Ent. Soc., Trans. 25: 289.
Brachycystis (!) Viereck, 1906. Amer. Ent. Soc., Trans. 32: 190.
Stilbopogon Mickel and Krombein, 1942. Amer. Midland Nat. 28: 658.
Type-species: Stilbopogon alutacea Mickel and Krombein. Orig. desig.
Astigmometopa Mickel and Krombein, 1942. Amer. Midland Nat. 28: 668.
Type-species: Astigmometopa emarginata Mickel and Krombein. Orig. desig.

Revision: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 1-96, 144 figs., 21 maps (males).

Taxonomy: Wasbauer, 1969 (1968). Pan-Pacific Ent. 44: 297-298 (sex association).
—Wasbauer, 1971. Pan-Pacifie Ent. 47: 211-212 (sex association).

SPECIES GROUP NITIDA

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 20.
arenivaga Bradley. Sonoran desert of Calif. and Ariz.; Mexico (Baja California, Sonora).
Brachycistis (Brachycistis) arenivaga Bradley, 1917. Amer. Ent. Soc., Trans. 43: 269. ¢.
Brachycistis (Brachycistis) eriopis Bradley, 1917. Amer. Ent. Soc., Trans. 43: 266. ¢.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 20-21, figs. 23, 40, 95, 96, map 1. d.
linsleyi Wasbauer. Mojave, Sonoran and Chihuahuan deserts, Calif., Nev., Ariz., N. Mex. and

Tex.; Mexico (Baja California, Chihuahua).
Brachycistis linsleyi Wasbauer, 1966. Univ. Calif. Pubs. Ent. 48: 21, figs. 16, 17, 41, 97, 98,

map 2. 6.
nitida (Cresson). Great Basin and Rocky Mt. grasslands, Nebr., Kans., Tex., Mont., Utah, N.
Mex., Ariz.
Agama nitida Cresson, 1875. Rpt. Geog. Geol. Expl. and Survey west of 100th Meridian, p.
TO iG):

Photopsis sobrinus Blake, 1886. Amer. Ent. Soc., Trans. 13: 268. ¢.
Photopsis lepidus Blake, 1886. Amer. Ent. Soc., Trans. 13: 269. ¢.
Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 23-25, figs. 43, 44, 99, 100, map 3. ¢.
texana Malloch. Chihuahuan desert of Tex.; Mexico (Coahuila, Nuevo Leon).
Brachycistis (Brachycistis) terana Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 12. 3.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 25-27, figs. 18, 19, 45, 46, 91, 101, 102,
map 5 (not 4). 6.

Superfamily SCOLIOIDEA 1269

verticalis Malloch. Sonoran desert of Ariz.; Mexico (Sonora).
Brachycistis (Brachycistis) verticalis Malloch, 1926. U. S. Natl. Mus., Proce. 68 (3): 19. d.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 27-30, map 4 (not 5). d.

SPECIES GROUP PETIOLATA

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 30-31.
elegantula Cockerell and Casad. Southern Mojave, Sonoran and Chihuahuan deserts of Calif.,

Ariz., N. Mex. and Tex.; Mexico (Baja California, Sonora).
Brachycistus (!) elegantulus Cockerell and Casad, 1894. Ent. News 5: 295. 6.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 31-33, figs. 35, 47, 48, 103, 104, map 6.
3.
imitans Malloch. Central Valley, Mojave and Sonoran deserts of Calif., Ariz.; Mexico (Baja

California).
Brachycistis (Brachycistis) imitans Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 23. ¢.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 33-35, figs. 39, 49, 50, 92, 105, 106,
map 7. 6. —Wasbauer, 1969 (1968). Pan-Pacific Ent. 44: 298-299. 2.
juncea Wasbauer. Calif. (San Bernardino Co.).
Brachycistis juncea Wasbauer, 1966. Univ. Calif. Pubs., Ent. 43: 35, figs. 36, 51, 52, 107,
108. do.
lacustris hurdi Wasbauer. Sonoran desert of Calif., Ariz., ? Tex.
Brachycistis lacustris hurdi Wasbauer, 1966. Univ. Calif. Pubs., Ent. 43: 38, map 8. d.
lacustris lacustris Malloch. Mojave desert of Calif., Nev.
Brachycistis (Brachycistis) lacustris Malloch, 1926. U. S. Natl. Mus., Proc. 68 (8): 24. d.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 37, map 8. 6.
longula Wasbauer. Calif. (Riverside Co.). '
Brachycistis longula Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 39, figs. 37, 55, 56, 111,
12a
petiolata Fox. Mojave and Sonoran deserts of Calif., Nev., Ariz., also Utah, Idaho; Mexico
(Baja California, Sonora).
Brachycistis petiolata Fox, 1893. Calif. Acad. Sci., Proce. (2) 4: 8. d.
Brachycistis gaudii Cockerell, 1901. Canad. Ent. 33: 340. ¢.
Brachycistis (Brachycistis) parva Malloch, 1926. U.S. Natl. Mus., Proc. 68 (3): 14. ¢.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 40-42, figs. 8, 27, 57, 58, 118, 114, map
9. 6.

SPECIES GROUP ATRATA

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 43.
alcanor (Blake). Alta., Mont. and S. Dak. south to Ariz. and Tex.; Mexico (Chihuahua, Coahuila,
Durango, Jalisco, Mexico, Puebla, Zacatecas).
Agama Alcanor Blake, 1871. Amer. Ent. Soc., Trans. 3: 264. d.
Brachycistis cremastogaster Melander, 1903. Amer. Ent. Soc., Trans. 29: 329. ¢.
Brachycistis crematogaster (!) Krombein, 1951. U. S. Dept. Agr., Monog. 2: 743.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 46-48, 75, figs. 30, 61, 62, 117, 118,
map 11. 6.
ampla (Blake). Colo., Ariz., Nev., Calif., Wash.
Agama ampla Blake, 1879. Amer. Ent. Soc., Trans. 7: 252. ¢.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 438: 48, figs. 63, 64, 119, 120, map 12. d.
atrata (Blake). B. C., Wash. and Idaho south to Calif. and Ariz.; Mexico (Baja California,

Sonora).

Agama atrata Blake, 1879. Amer. Ent. Soc., Trans. 7: 253. ¢.

Mutilla agama Dalla Torre, 1897. Cat. Hym., v. 8, p. 7. N. name for Agama atrata Blake,
not Mutilla atrata Linnaeus.

Brachycistis nudus Fox, 1899. Amer. Ent. Soc., Trans. 25: 281. ¢.

Brachycistis nigritus Fox, 1899. Amer. Ent. Soc., Trans. 25: 282. ¢.

1270 Hymenoptera in America North of Mexico

Brachycistis contiguus Fox, 1899. Amer. Ent. Soe., Trans. 25: 282. d.
Glyptometopa francisca Mickel and Krombein, 1942. Amer. Midland Nat. 28: 656. °.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 43-46, figs. 6, 21, 34, 59, 60, 89, 115,
116, map 10. ¢. —Wasbauer, 1969 (1968). Pan-Pacific Ent. 44: 297-298 (female synonymy).
carinata Fox. South, Calif. coastal mts.; Mexico (Baja California).
Brachycistis carinatus Fox, 1899. Amer. Ent. Soc., Trans. 25: 283. d.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 49-51, figs. 20, 65, 121, 122, map 18.
36. —Wasbauer, 1971. Pan-Pacifie Ent. 47: 212. °.
convexa Wasbauer. Calif. (Imperial Co.).
Brachycistis convexa Wasbauer, 1966. Univ. Calif. Pubs. Ent. 48: 51, figs. 31, 66, 67, 94,
123, 124. ¢.
cruenta Wasbauer. Tex. (Nueces Co.); Mexico (Tamaulipas).
Brachycistis cruenta Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 53, figs. 68, 69, 125, 126.
3.
curvata Malloch. Sonoran desert of Calif., Ariz.; Mexico (Baja California).
Brachycistis (Brachycistis) curvata Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 16. ¢.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 54-56, figs. 70, 71, 127, 128, map 14.
3.
davidi Wasbauer. Tex.; Mexico (Coahuila).
Brachycistis davidi Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 56, figs. 22, 32, 72, 73, 129,
130. d.
glabrella (Cresson). Alta., Oreg., Idaho, Wyo. and Nebr. to Calif. and Tex., south in Mexico to
Oaxaca.
Mutilla glabrella Cresson, 1865. Ent. Soc. Phila., Proc. 4: 441. ¢.
Brachycistis (Brachycistis) dentata Bradley, 1917. Amer. Ent. Soc., Trans. 43: 263. ¢.
Brachycistis (Brachycistis) micrommata Bradley, 1917. Amer. Ent. Soc., Trans. 43: 265. d.
Brachycistis (Brachycistis) utahensis Malloch, 1926. U. S. Natl. Mus., Proc. 68 (8): 17. ¢.
Brachycistis (Brachycistis) melanocephala Malloch, 1926. U.S. Natl. Mus., Proc. 68 (3): 19.

é.
Brachycistis (Brachycistis) dakotensis Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 20. 3.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 57-59, figs. 25, 33, 74, 75, 131, 132,
map 15. 6.
inaequalis Fox. South. Calif., Nev., Ariz. and Tex.; Mexico (Baja California).
Brachycistis inaequalis Fox, 1899. Amer. Ent. Soc., Trans. 25: 282. ¢.
Brachycistis nevadensis Fox, 1899. Amer. Ent. Soc., Trans. 25: 283. ¢.
Brachycistis nocticola Bradley, 1917. Amer. Ent. Soc., Trans. 43: 289, figs. 19, 39, 40. d.

Taxonomy: Krombein, 1954. Ent. Soc. Wash., Proc. 56: 85-86 (synonymy of nocticola).
—Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 59-61, figs. 29, 76, 77, 133, 134, map 16. ¢.
indiscreta Fox. Nebr. to Tex., N. Mex., Ariz.; Mexico (Chihuahua).
Brachycistis indiscreta Fox, 1899. Amer. Ent. Soc., Trans. 25: 284. d.
Brachycystis (!) stictinotus Viereck, 1906. Amer. Ent. Soc., Trans. 32: 190. d.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 61-63, figs. 26, 38, 78, 135, 136, map
enor
ioachinensis Bradley. Wash. and Mont. to Calif. and Tex. south in Mexico to Baja California
and Sonora.
Brachycistis (Brachycistis) ioachinensis Bradley, 1917. Amer. Ent. Soc., Trans. 43: 267. d.
Brachycistis (Brachycistis) rugosa Malloch, 1926. U. S. Natl. Mus., Proc. 68 (8): 11. 3.
Brachycistis (Brachycistis) subcarinata Malloch, 1926. U. S. Natl. Mus., Proc. 68(3): 11. ¢.
Brachycistis (Brachycistis) nigrifrons Malloch, 1926. U.S. Natl. Mus., Proce. 68 (3): 16. d.
Brachycistis (Brachycistis) washingtona Malloch, 1926. U.S. Natl. Mus., Proce. 68 (3): 21.
3.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 48: 65-68, figs. 1, 12, 13, 79, 80, 137, 138,
map 18. 6.

Superfamily SCOLIOIDEA 1271

semiatra Malloch. Wash. to northeast. Calif., to Idaho and Utah.
Brachycistis (Brachycistis) semiatra Malloch, 1926. U. 8S. Natl. Mus., Proce. 68 (8): 24. 6.
Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 68-70, figs. 81, 82, 139, 140, map 19.
3.
subquadrata Fox. Calif. (San Diego Co.).
Brachycistis subquadratus Fox, 1899. Amer. Ent. Soc., Trans. 25: 282. 6.
Brachycistis (Brachycistis) intermedia Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 18. 6.
Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 64-65. 3.
timberlakei Wasbauer. Calif., Nev., Utah, Ariz., N. Mex., Tex.
Brachycistis timberlakei Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 70, figs. 90, 141, 142,
map 20. 6.
triangularis Fox. South. Calif., Nev., Ariz., N. Mex., Tex.; Mexico (Baja California, Sonora,
Chihuahua, Durango).
Brachycistis triangularis Fox, 1899. Amer. Ent. Soc., Trans. 25: 284. 3.
Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 72-78, figs. 24, 83, 84, 93, 143, 144,
map 21. 6.
UNPLACED TAXA OF BRACHYCISTIS Fox
alutacea (Mickel and Krombein). Tex., N. Mex.
Stilbopogon alutacea Mickel and Krombein, 1942. Amer. Midland Nat. 28: 659. 9.

americana (Ashmead). Calif. (Alameda Co.).
Glyptometopa americana Ashmead, 1898. Psyche 8: 251. °.

convergens (Mickel and Krombein). Calif. (Santa Monica).
Glyptometopa convergens Mickel and Krombein, 1942. Amer. Midland Nat. 28: 657. °.

emarginata (Mickel and Krombein). Tex. (Valentine).
Astigmometopa emarginata Mickel and Krombein, 1942. Amer. Midland Nat. 28: 668. °.
Taxonomy: Wasbauer, 1971. Pan-Pacifie Ent. 47: 212.

protracta Bradley. Ariz. (Phoenix).
Brachycistis (Brachycistis) protracta Bradley, 1917. Amer. Ent. Soc., Trans. 43: 270. 3.
Brachycistis (Brachycistis) protractor (!) Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 7.

Taxonomy: Wasbauer, 1966. Univ. Calif. Pubs. Ent. 43: 74. d.
Genus COLOCISTIS Krombein

Colocistis Krombein, 1942. Ent. Soc. Wash., Proc. 44: 65.
Type-species: Colocistis pilosa Krombein. Orig. desig.
Taxonomy: Wasbauer, 1968. Pan-Pacifie Ent. 44: 197, figs. 21, 23.

brevis (Fox). Ariz., Nev., Calif.
Brachycistis brevis Fox, 1899. Amer. Ent. Soe., Trans. 25: 285. 3.

castanea (Cresson). Tex., N. Mex., Colo., Ariz., Nev., Calif.
Mutilla castanea Cresson, 1865. Ent. Soc. Phila., Proc. 4: 388. 3.

crassa (Bradley). Tex., Colo., Ariz., Nev., Calif.
Brachycistis (Brachycistis) crassa Bradley, 1917. Amer. Ent. Soc., Trans. 43: 277. 3.

eremi (Bradley). Ariz., Nev., Calif.
Brachycistis (Brachycistis) eremi Bradley, 1917. Amer. Ent. Soc., Trans. 43: 279. 6.

pilosa Krombein. Calif. (San Diego).
Colocistis pilosa Krombein, 1942. Ent. Soc. Wash., Proc. 44: 66. d.

stygia (Bradley). Ariz. (Nogales).
Brachycistis (Brachycistis) stygia Bradley, 1917. Amer. Ent. Soc., Trans. 43: 276. 3.

thermarum (Bradley). Ariz.
Brachycistis (Brachycistis) thermarum Bradley, 1917. Amer. Ent. Soc., Trans. 43: 274. ¢.

Genus ACANTHETROPIS Wasbauer
Acanthetropis Wasbauer, 1958. Pan-Pacifie Ent. 34: 139.

1272 Hymenoptera in America North of Mexico

Type-species: Acanthetropis lamellatus Wasbauer. Orig. desig.

Taxonomy: Wasbauer, 1968. Pan-Pacifie Ent. 44: 197, figs. 20, 24.

aequalis (Fox). Nebr., Kans., Colo., Idaho, Nev., Ariz.
Brachycistis aequalis Fox, 1899. Amer. Ent. Soc., Trans. 25: 284. d.

idiotes (Cockerell). N. Mex., Calif.
Brachycistis idiotes Cockerell, 1895. Ent. News. 6: 63. ¢.

noctivaga (Bradley). Utah, Nev., Ariz., Calif.; Mexico (Sonora, Baja California).
Brachycistis (Brachycistis) noctivaga Bradley, 1917. Amer. Ent. Soc., Trans. 43: 272. d.

normalis (Malloch). S. Dak., Colo., Ariz., Calif.
Brachycistis normalis Malloch, 1926. U. S. Natl. Mus., Proc. 68 (8): 8. ¢.

Genus BRACHYCISTINA Malloch

Brachycistis subg. Brachycistina Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 5, 25.
Type-species: Brachycistis (Brachycistina) acuta Malloch. Orig. desig.
Taxonomy: Wasbauer, 1968. Pan-Pacific Ent. 44: 197, fig. 25.

acuta (Malloch). Ariz., Nev., Calif.
Brachycistis (Brachycistina) acuta Malloch, 1926. U. S. Natl. Mus., Proc. 68 (3): 25. ¢.

Genus HADROCISTIS Wasbauer

Hadrocistis Wasbauer, 1968. Pan-Pacifiec Ent. 44: 185.
Type-species: Hadrocistis bicolor Wasbauer. Orig. desig.

bicolor Wasbauer. Calif. (Imperial Co.).
Hadrocistis bicolor Wasbauer, 1968. Pan-Pacific Ent. 44: 187, figs. 4-6, 14, 18. ¢.

slanskyae Wasbauer. Calif. (Imperial Co.).
Hadrocistis slanskyae Wasbauer, 1968. Pan-Pacifie Ent. 44: 188, figs. 1-3, 13, 17. ¢.

Genus BRACHYCISTELLUS Baker

Brachycistellus Baker, 1907. Invertebrata Pacifica 1: 177.
Type-species: Brachycistellus figitiformis Baker. Monotypic.
Taxonomy: Wasbauer, 1968. Pan-Pacific Ent. 44: 193-195.
figitiformis Baker. Calif. (Colusa Co., Claremont).
Brachycistellus figitiformis Baker, 1907. Invertebrata Pacifica 1: 178. 6.

Taxonomy: Wasbauer, 1968. Pan-Pacific Ent. 44: 195-196, figs. 10-12, 16, 19. d.
Genus QUEMAYA Pate

Quemaya Pate, 1947. N. Y. Ent. Soe., Jour. 55: 139.
Type-species: Brachycistis (Brachycistellus) paupercula Bradley. Orig. desig.
Taxonomy: Wasbauer, 1968. Pan-Pacific Ent. 44: 196, fig. 22.

arenicola Wasbauer. Calif. (Imperial Co.).
Quemaya arenicola Wasbauer, 1967. Biol. Soc. Wash., Proc. 80: 169, figs. 2-6. 3.

inermis (Malloch). Ariz. (Higley).
Brachycistus (!) (Brachycistellus) inermis Malloch, 1924. Brooklyn Ent. Soce., Bul. 19: 23.
3.

marcida (Bradley). Calif. (Calexico).
Brachycistis (Brachycistellus) marcida Bradley, 1917. Amer. Ent. Soc., Trans. 43: 283. d.

paupercula (Bradley). Calif., Nev.
Brachycistis (Brachycistellus) pawpercula Bradley, 1917. Amer. Ent. Soc., Trans. 43: 282.

3.
perpunctata (Cockerell). N. Mex., Calif.
Brachycistis perpunctatus Cockerell, 1896. Amer. Ent. Soc., Trans. 22: 291. 6.

Taxonomy: Wasbauer, 1968. Biol. Soc. Wash., Proc. 80: 170, fig. 1.

Superfamily SCOLIOIDEA 1273

Genus GLYPTACROS Mickel and Krombein

Glyptacros Mickel and Krombein, 1942. Amer. Midland Nat. 28: 660.
Type-species: Glyptacros angustior Mickel and Krombein. Orig. desig.

angustior Mickel and Krombein. Ariz.
Glyptacros angustior Mickel and Krombein, 1942. Amer. Midland Nat. 28: 660. °.

ashmeadi Mickel and Krombein. No locality given.
Glyptacros ashmeadi Mickel and Krombein, 1942. Amer. Midland Nat. 28: 661. °.

Genus XEROGLYPTA Mickel and Krombein

Xeroglypta Mickel and Krombein, 1942. Amer. Midland Nat. 28: 663.
Type-species: Xeroglypta egregia Mickel and Krombein. Orig. desig.
egregia Mickel and Krombein. Calif. (Palm Springs).
Xeroglypta egregia Mickel and Krombein, 1942. Amer. Midland Nat. 28: 663. °.

Genus AULACROS Mickel and Krombein

Aulacros Mickel and Krombein, 1942. Amer. Midland Nat. 28: 664.
Type-species: Aulacros latior Mickel and Krombein. Orig. desig.

latior Mickel and Krombein. Calif. (Palm Springs).
Aulacros latior Mickel and Krombein, 1942. Amer. Midland Nat. 28: 665. 2.

Genus BRUESIELLA Mann

Bruesiella Mann, 1914. Psyche 21: 182.
Type-species: Bruesiella formicaria Mann. Orig. desig.
Eurycros Mickel and Krombein, 1942. Amer. Midland Nat. 28: 666.
Type-species: Eurycros furtivus Mickel and Krombein. Orig. desig.
Taxonomy: Krombein, 1967. U. S. Dept. Agr., Monog. 2, Sup. 2: 325 (synonymy).

furtivus (Mickel and Krombein). Ariz. (Tucson, Avondale Ranch).
Eurycros furtivus Mickel and Krombein, 1942. Amer. Midland Nat. 28: 667. 2.

Genus AGLYPTACROS Mickel and Krombein

Aglyptacros Mickel and Krombein, 1942. Amer. Midland Nat. 28: 669.
Type-species: Glyptometopa eureka Banks. Orig. desig.

eureka (Banks). Ariz. (Palmerlee).

Glyptometopa eureka Banks, 1912. Canad. Ent. 44: 202. °.
paxillatus Mickel and Krombein. Colo. (La Junta).

Aglyptacros paxillatus Mickel and Krombein, 1942. Amer. Midland Nat. 28: 673. 2.
segredentatus Mickel and Krombein. Ariz. (Benson).

Aglyptacros segredentatus Mickel and Krombein, 1942. Amer. Midland Nat. 28: 671. °.

sulcatus Mickel and Krombein. Tex. (Valentine).
Aglyptacros sulcatus Mickel and Krombein, 1942. Amer. Midland Nat. 28: 674. °.

SUBFAMILY METHOCHINAE
Genus METHOCHA Latreille

Genus METHOCHA Subgenus METHOCHA Latreille

Methocha Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, p. 179.
Type-species: Mutilla articulata Latreille. Monotypic.
Methoca (!) Latreille, 1804. Hist. Nat. Crust. Ins., v. 18, p. 268.
Tengyra Latreille, 1809. Gen. Crust. Ins., v. 4, p. 115.
Type-species: Tengyra Sanvitali Latreille. Monotypic.
Spinolia Costa, 1858. Fauna Napoli Scol., p. 21.
Type-species: Spinolia italica Costa. Monotypic.

1274. Hymenoptera in America North of Mexico

Two of our native species have been reared from larvae of tiger beetles, the known host of
several extralimital species.
Only the typical subgenus occurs in North America.
Morphology: Reid, 1941. Roy. Ent. Soe. London, Trans. 91: 389-390, figs. 26-28 (female, male
thorax).
californica Westwood. Calif., Nev., Wash. Host: Cicindela senilis Horn ?; cicindelid sp.; reared
in laboratory on Omus californicus Esch.
Methoca (!) californica Westwood, 1881. Ent. Soe. London, Trans., p. 133. @.
Methoca (!) nigrior Fox, 1899. Amer. Ent. Soc., Trans. 25: 288. ¢.

Biology: Bridwell, 1912. Pacific Coast Ent. Soe., Proc. 1: 46-48 (host). —Burdick and
Wasbauer, 1959. Wasmann Jour. Biol. 17: 75-88, 6 figs. (prey capture, oviposition, life
history, egg, larva).

formosa Krombein. Fla. (Arcadia).
Methocha (Methocha) formosa Krombein, 1954. Amer. Ent. Soc., Trans. 80: 2, figs. 1, 3. 2.

impolita Krombein. W. Va., D. C., N. Y.
Methocha (Methocha) impolita Krombein, 1958. Ent. Soc. Wash., Proc. 60: 58. 2.

stygia (Say). General in south. Canada and U. S. Host: Cicindelid larvae.
Mutilla (Methoca (!)) pacalis Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool.
Mass., p. 587. Nom. nud.
Tengyra stygia Say, 1836. Boston Jour. Nat. Hist. 1: 299. 6.
Methoca (!) bicolor Say, 1836. Boston Jour. Nat. Hist. 1: 299. °.
Methoca (!) canadensis Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 67. 3.

Taxonomy: Evans, 1965. Ent. Soc. Wash., Proc. 67: 88-90, figs. 1-5 (larva).
Biology: Williams, 1916. Psyche 23: 121-125, pl. 12.

SUBFAMILY MYRMOSINAE

The Myrmosinae have as hosts the larvae of small ground-nesting wasps and bees.
Brothers (1975) transferred this subfamily to the Mutillidae.

Revision: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 415-465, pl. 24.
Taxonomy: Wasbauer, 1974 (1973). Pan-Pacific Ent. 49: 335 (key to female genera).

Genus MYRMOSA Latreille

Genus M-YRMOSA Subgenus MYRMOSA Latreille

Myrmosa Latreille, 1796. Precis Caract. Gen. Ins., p. 118.
Type-species: Myrmosa atra Panzer. First included species.
Ischioceras Provancher, 1882. Nat. Canad. 13: 8.
Type-species: Ischioceras rugosa Provancher. First included species.
Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 384-385, figs. 11-14 (female, male
thorax).
blakei Bradley. N. Y., Va.
Myrmosa (Myrmosa) blakei Bradley, 1917. Amer. Ent. Soc., Trans. 43: 251. @.
Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 438. °.
bradleyi Roberts. Wash., Calif.
Myrmosa bradleyi Roberts, 1929. Psyche 36: 362. ¢.
Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 436-438, fig. 2. d, 2.

Biology: Linsley, 1960. Pan-Pacific Ent. 36: 36 (mating behavior).
peculiaris Krombein. Kans. (Onaga).
Myrmosa (Myrmosa) peculiaris Krombein, 1940. Amer. Ent. Soc., Trans. 65: 438. 2.
unicolor Say. N. B. south to N. C., west to B. C. and Ariz. Host: Tiphia sp.; Lindenius
columbianus errans (Fox) ?; Dialictus pruinosus (Robt.); D. inconspicuus (Sm.).
Myrmosa unicolor Say, 1824. In Keating, Narr. Long’s 2nd. Exped., v. 2, p. 331. ¢.

Superfamily SCOLIOIDEA 1275

Ischioceras rugosa Provancher, 1882. Nat. Canad. 13: 8. 2, (d misdet.).

Mutilla thoracica Blake, 1886. Amer. Ent. Soc., Trans. 13: 204. Preoce.

Mutilla erythronota Dalla Torre, 1897. Cat. Hym., v. 8, p. 36. N. name.

Myrmosa (Myrmosa) banksi Bradley, 1917. Amer. Ent. Soc., Trans. 43: 249. 2, d.
Myrmosa dakotensis Weber, 1934. Psyche 41: 57. 2.

Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 430-436, fig. 6.

Biology: Melander and Brues, 1903. Biol. Bul. 5: 7. —Krombein, 1940. Amer. Ent. Soc., Trans.
65: 435-436. —Krombein, 1956. Ent. Soe. Wash., Proc. 58: 154 (mating behavior).

Genus MYRMOSA Subgenus MYRMOSINA Krombein

Myrmosa subg. Myrmosina Krombein, 1940. Amer. Ent. Soc., Trans. 65: 452.
Type-species: Myrmosa (Myrmosina) texana Krombein. Orig. desig.
nocturna nocturna Krombein. Md., La., Tex., Kans.
Myrmosa (Myrmosina) nocturna nocturna Krombein, 1940. Amer. Ent. Soe., Trans. 65:
454. 6.
nocturna rufigastra Krombein. Tex., Calif.
Myrmosa (Myrmosina) nocturna rufigastra Krombein, 1940. Amer. Ent. Soc., Trans. 65:
455. 6.
texana Krombein. Dak., Tex.
Myrmosa (Myrmosina) tecana Krombein, 1940. Amer. Ent. Soc., Trans. 65: 453. ¢.

Genus LEIOMYRMOSA Wasbauer

Leiomyrmosa Wasbauer, 1974 (1973). Pan-Pacifie Ent. 49: 325.
Type-species: Leiomyrmosa spilota Wasbauer. Orig. desig.
spilota Wasbauer. Calif. (Blythe in Riverside Co.).
Leiomyrmosa spilota Wasbauer, 1974 (1973). Pan-Pacifie Ent. 49: 326, figs. 1, 2, 19, 23. °.

Genus MYRMOSULA Bradley

Myrmosa subg. Myrmosula Bradley, 1917. Amer. Ent. Soc., Trans. 43: 249.
Type-species: Myrmosa parvula Fox. Desig. by Bridwell, 1920.

Taxonomy: Mickel, 1940. Pan-Pacific Ent. 16: 133-134 (key to females). —Wasbauer, 1974
(1973). Pan-Pacific Ent. 49: 335-336 (key to females).
boharti Wasbauer. Calif. (Thousand Palms in Riverside Co.).
Myrmosula boharti Wasbauer, 1974 (1973). Pan-Pacific Ent. 49: 331, figs. 18, 14. 2.

exaggerata (Krombein). Calif.
Myrmosa (Myrmosula) exaggerata Krombein, 1940. Amer. Ent. Soc., Trans. 65: 460. °.

Taxonomy: Mickel, 1940. Pan-Pacific Ent. 16: 133.
nasuta Wasbauer. Southern Calif. and Ariz. Ecology: Visits mat Euphorbia and has been
collected in a kangaroo rat burrow.
Myrmosula nasuta Wasbauer, 1974 (1973). Pan-Pacifie Ent. 49: 329, figs. 7, 8, 20. 2.
pacifica (Mickel). Calif. (Antioch). Possibly a synonym of exaggerata Krom.
Myrmosa (Myrmosula) pacifica Mickel, 1940. Pan-Pacific Ent. 16: 134. °.
parvula (Fox). D. C. south to Ala., west to Mont. and Tex. Host: Dialictus imitatus (Sm.), D.
zephyrus (Sm.); Augochlorella striata (Prov.)(?), A. persimilis (Vier.)(?).
Myrmosa parvula Fox, 1893. N. Y. Ent. Soc., Jour. 1: 53. d.
Mutilla antisemitica Dalla Torre, 1897. Cat. Hym., v. 8, p. 10. N. name for Myrmosa
parvula fox, not Mutilla parvula Fabricius.
Brachycistis bimaculatus Fox, 1899. Amer. Ent. Soc., Trans. 25: 285. °.

Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 457-459, figs. 5, 10. d, °.

Biology: Michener and Wille, 1961. Univ. Kans. Sci. Bul. 42: 1130 (host record). —Ordway,
1964. Kans. Ent. Soc., Jour. 37: 149 (host records). —Batra, 1965. Kans. Ent. Soe., Jour. 38:
386-387 (behavior in host nest). —Brothers, 1972. Univ. Kans. Sci. Bul. 50: 23 (host record).

1276 Hymenoptera in America North of Mexico

peregrinatrix (Krombein). Alta., Ariz., Tex.
Myrmosa (Myrmosula) peregrinatrix Krombein, 1946. Ent. Soc. Wash., Proc. 48: 247. 9.
rufiventris Blake. Nev., Oreg.
Myrmosa rufiventris Blake, 1879. Amer. Ent. Soc., Trans. 7: 254. d.
Mutilla erythrogastra Dalla Torre, 1897. Cat. Hym., v. 8, p. 36. N. name for Myrmosa
rufiventris Blake, not Mutilla rufiventris Klug nor M. rufiventris Smith.

Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 459-460. d.

rutilans (Blake). Calif. Host: Nomadopsis scutellaris (Fwlr.)
Mutilla rutilans Blake, 1879. Amer. Ent. Soc., Trans. 7: 248. °.

Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 461-462, fig. 12. 2.
Family SIEROLOMORPHIDAE

This is a primitive scolioid group whose closest relationship appears to be with the Tiphiidae.
The family is known from the single genus Sierolomorpha which has six New World species
ranging from North America south to Panama, one possibly adventive species in Hawaii, and
two Old World species from North China and Tadzhikistan.

There is no information on host relationships, but the lack of a tarsal pecten in Sierolomorpha
females suggests that they are not fossorial in habit.

Revision: Evans, 1961. Breviora, No. 140, 12 pp., 6 figs.

Taxonomy: Melander and Prues, 1932. Mus. Compar. Zool. Bul. 73: 500. —Schuster, 1949.
Ent. Amer. (n. s.) 29: 124-125.

Genus SIEROLOMORPHA Ashmead

Sierolomorpha Ashmead, 1903. Canad. Ent. 35: 42.
Type-species: Sierola (?) ambigua Ashmead. Orig. desig.

apache Evans. Ariz.

Sierolomorpha apache Evans, 1961. Breviora, No. 140, p. 5, fig. 6. dé.
bicolor Evans. Ariz.

Sierolomorpha bicolor Evans, 1961. Breviora, No. 140, p. 4, fig. 5. 2, 3.
brevicornis Evans. S. C. (Greenville).

Sierolomorpha brevicornis Evans, 1961. Breviora, No. 140, p. 8, fig. 3. ¢.
canadensis (Provancher). Alta., Ont. and Mass. south to Ariz., Tex. and Fla.

Photopsis Canadensis Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 410. 6.

Secondary homonym in Mutilla.

Sierola? ambigua Ashmead, 1893. U.S. Natl. Mus., Bul. 45: 56.

Mutilla tertia Dalla Torre. 1897. Cat. Hym., v. 8, p. 91. N. name.
nigrescens Evans. Yukon, Alta. and Sask. south to Calif., Ariz. and Colo.

Sierolomorpha nigrescens Evans, 1961. Breviora, No. 140, p. 9, fig. 1. d, @.
similis Evans. Conn. south to Ga., W. Va., Kans.

Sierolomorpha similis Evans, 1961. Breviora, No. 140, p. 6, fig. 4. 5, 9.

Family MUTILLIDAE

Females are entirely wingless and have a greatly modified thorax. Males are normally fully
winged, but are brachypterous or apterous in a few species. Some females are called cowkillers
or mulekillers because of their size and painful sting, or velvet ants because of the appearance of
the dense pile covering the body.

Hosts are known for very few of the North American species. The stages of host attacked are
limited to diapausing larvae or pupae. Most recorded North American hosts are wasps or bees
which nest in the soil, in borings in wood or which build mud or resin cells or paper nests. How-
ever, there is one authenticated record of coleopterous pupae as a host. Extralimital genera have
been recorded as parasitizing Lepidoptera, Coleoptera and Diptera, as well as aculeate
Hymenoptera.

Brothers’ important contribution on phylogeny of the Mutillidae was received too late to adopt

Superfamily SCOLIOIDEA 1277

his classification but it will be used in the next edition. He places the Myrmosinae here rather
than in the Tiphiidae. He raises the exotic Bradynobaeninae to family rank and transfers to it as
subfamilies the Typhoctinae and Chyphotinae (here considered a tribe of Apterogyninae). He as-
signs the other North American genera as follows: Sphaeropthalminae, Pseudomethocina, M yr-
milloides, Pseudomethoca; Sphaeropthalminae, Sphaeropthalmina, Acanthophotopsis,
Acrophotopsis, Dasymutilla, Dilophotopsis, Lomachaeta, Morsyma, Odontophotopsis,
Photomorphus, Protophotopsis, Smicromutilla, Sphaeropthalma; Mutillinae, Mutillini, Smicro-
myrmina, Timulla; and Mutillinae, Ephutini, FE phuta.

Taxonomy: Blake, 1886. Amer. Ent.. Soc., Trans. 13: 179-286, 21 figs. (N. A. spp.).
—Ashmead, 1899. N. Y. Ent. Soc., Jour. 7: 52-60 (gen. of world). —Fox, 1899. Amer. Ent.
Soc., Trans. 25: 219-292 (N. A. spp.). —Andre, 1903. In Wytsman, Gen. Ins., fase. 11, pp.
1-77, 3 pls. (gen. of world). —Melander, 1903. Amer. Ent. Soc., Trans. 29: 291-330, pl. 4
(taxonomic notes and some revised keys). —Bradley, 1916. Amer. Ent. Soc., Trans. 42:
309-336 (spp. of east. U. S.). —Schuster, 1946. Ent. Soc. Amer., Ann. 39: 700-703 (subfam.
of New World). —Schuster, 1949. Ent. Amer. (n. s.) 29: 61-64 (key to Neotrop. subfam.).
—Mickel, 1970. Minn. Univ., Agr. Expt. Sta., Tech. Bul. 271: 1-77 (annotated bibliography
of world literature). —Brothers, 1975. Kans. Univ. Sci. Bul. 50: 483-648, 101 figs., 7 tabs.
(phylogeny).

Morphology: Reid, 1941. Roy. Ent. Soe. London, Trans. 91: 375-387, figs. 1-10, 15-20 (female,
male thorax). —Hinton, Gibbs and Silberglied, 1969. Jour. Ins. Physiol. 15: 549-552, 10 figs.
(stridulatory file). —Sheldon, 1970. Ent. News 81: 57-61 (male adaptations for female
carriage). —Debolt, 1973. Ent. Soc. Amer., Ann. 66: 100-108, 15 figs. (felt line and felt line
organ).

SUBFAMILY APTEROGYNINAE
The North American species are all nocturnal.
TRIBE CHYPHOTINI

Genus CHYPHOTES Blake

Revision: Buzicky, 1941. Ent. Amer. (n. s.) 21: 201-243. —Mickel, 1967. Amer. Ent. Soc.,
Trans. 93: 125-234, 40 figs., 26 maps.

Genus CHYPHOTES Subgenus CHYPHOTES Blake

Chyphotes Blake, 1886. Amer. Ent. Soc., Trans. 13: 276.
Type-species: Chyphotes elevatus Blake. Orig. desig.
Blaketa Pate, 1948. Ent. News 59: 41. N. name for Chyphotes Blake, not Cyphotes
Burmeister 1835.

aenigmus Mickel. N. Mex. to Calif.; Mexico (Sonora).
Chyphotes (Chyphotes) aenigmus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 147, fig. 2, map
Bild

atriceps Mickel. Wash., Idaho, Oreg., Calif., Nev., Utah, Ariz., N. Mex., west. Tex.; Mexico

(Sonora, Chihuahua).
Chyphotes (Chyphotes) atriceps Mickel, 1967. Amer. Ent. Soc., Trans. 93: 134, fig. 1, map 1.
é.

auripilus Buzicky. Calif., Ariz.
Chyphotes auripilus Buzicky, 1941. Ent. Amer. (n. s.) 21: 232. 9°.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 174.
belfragei (Blake). Wyo., Nebr., Colo., Utah, Ariz., N. Mex.; Mexico (Baja California, Chihuahua
to Nuevo Leon, south to Jalisco and Puebla).

Agama Belfragei Blake, 1871. Amer. Ent. Soc., Trans. 3: 263. ¢.
Sphaerophthalma(!) frugala Cameron, 1896. Biol. Cent.-Amer., Hym. 2: 394. d.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 151, fig. 12, map 7. d, °.
Morphology: Reid, 1941. Roy. Ent. Soe. London, Trans. 91: 386-387, figs. 19-20 (male thorax).

1278 Hymenoptera in America North of Mexico

bicolor Schuster. Calif. (San Diego).
Chyphotes bicolor Schuster, 1945. Pan-Pacifie Ent. 21: 89. °.

calexicensis Bradley. Ariz., Calif.; Mexico (Baja California, Sonora).
Chyphotes calexicensis Bradley, 1917. Amer. Ent. Soc., Trans. 43: 284. 6.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 144, fig. 7, map 4.

californicus Baker. Ariz., Nev., Calif.
Chyphotes californicus Baker, 1905. Invertebrata Pacifica 1: 117. d.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 171, fig. 23, map 14.
capitatus Mickel. Ariz.
Chyphotes (Chyphotes) capitatus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 166, fig. 19, map
tee
cooki Mickel. Calif.
Chyphotes (Chyphotes) cooki Mickel, 1967. Amer. Ent. Soc., Trans. 93: 159, fig. 17, map 9.
3.

dubius Mickel. Calif., Ariz.; Mexico (Chihuahua).
Chyphotes (Chyphotes) dubius Mickel, 1967. Amer. Ent. Soc., Trans. 93: 175. 2.

elevatus Blake. N. Mex., Ariz.; Mexico (Sonora).
Chyphotes elevatus Blake, 1886. Amer. Ent. Soc., Trans. 13: 276. 2.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 155, fig. 11, map 8. 6, °.

incredulus Mickel. Southwest Tex., south. N. Mex.; Mexico (Chihuahua, Nuevo Leon,
Durango).
Chyphotes (Chyphotes) incredulus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 149, fig. 15,
map 6. 6.
knulli Mickel. Tex. (Davis Mts.).
Chyphotes (Chyphotes) knulli Mickel, 1967. Amer. Ent. Soc., Trans. 93: 181. 2.
mandibularis Mickel. Calif.
Chyphotes (Chyphotes) mandibularis Mickel, 1967. Amer. Ent. Soc., Trans. 93: 139, fig. 3,
map 2. 6, @.
melaniceps (Blake). Utah, Nev., Ariz., Calif.; Mexico (Baja California).
Photopsis melaniceps Blake, 1886. Amer. Ent. Soc., Trans. 13: 264. ¢.
Chyphotes piceiceps Baker, 1905. Invertebrata Pacifica 1: 116. ¢.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 141, fig. 6, map 3.
mexicanus Mickel. West. Tex., Ariz.; Mexico (Chihuahua, Sinaloa, Nayarit).
Chyphotes (Chyphotes) mexicanus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 168, fig. 20,
map 12. 6.
minusculus Mickel. Ariz., Nev., Calif.; Mexico (Baja California).
Chyphotes (Chyphotes) minusculus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 163, fig. 18,
map 10. 6.
peninsularis Fox. Calif.; Mexico (Baja California).
Chyphotes peninsularis Fox, 1899. Amer. Ent. Soc., Trans. 25: 277. 6.
Chyphotes jugatus Buzicky, 1941. Ent. Amer. (n. s.) 21: 222. ¢.
Chyphotes pilosus Buzicky, 1941. Ent. Amer. (n. s.) 21: 233. °.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 170, figs. 4, 21, map 13.
petiolatus Fox. Tex., Ariz., Nev., Calif.
Chyphotes petiolatus Fox, 1899. Amer. Ent. Soc., Trans. 25: 277. 2.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 174.
pimus Mickel. Ariz. (Pima Co.).
Chyphotes (Chyphotes) pimus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 181, fig. 5. °.
rugosus Mickel. Calif.
Chyphotes (Chyphotes) rugosus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 177. 2.
rugulosus Mickel. Ariz. (Pima Co.).
Chyphotes (Chyphotes) rugulosus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 173. 9.

Superfamily SCOLIOIDEA 1279

testaceipes Fox. Ariz.
Chyphotes testaceipes Fox, 1899. Amer. Ent. Soc., Trans. 25: 277. 2.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 173.

Morphology: Reid, 1941. Roy. Ent. Soe. London, Trans. 91: 386-387, figs. 17-18 (thorax).

wasbaueri Mickel. Calif. (San Bernardino Co.).
Chyphotes (Chyphotes) wasbaueri Mickel, 1967. Amer. Ent. Soc., Trans. 93: 180. 2.

Genus CHYPHOTES Subgenus PITANTA Pate

Pitanta Pate, 1947. Notulae Nat. 192: 1.
Type-species: Pitanta mojave Pate. Orig. desig.

SPECIES GROUP SIMILIS

similis Baker. Calif.; Mexico (Baja California).
Chyphotes similis Baker, 1905. Invertebrata Pacifica 1: 117. ¢.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 188, fig. 8, map 15.

SPECIES GROUP NUBECULUS

fergusoni Mickel. Oreg., Idaho, Utah, Nev., Calif.
Chyphotes (Pitanta) fergusoni Mickel, 1967. Amer. Ent. Soc., Trans. 93: 193, fig. 24, map
err
nubeculus (Cresson). Mont., Wyo., Nebr., Kans., Colo., Tex., N. Mex., Ariz.; Mexico (Sonora).
Mutilla nubecula Cresson, 1865. Ent. Soe. Phila., Proc. 4: 440. 3.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 191, fig. 22, map 16.
SPECIES GROUP HEATHII

bruscus Buzicky. Ariz., Calif.
Chyphotes bruscus Buzicky, 1941. Ent. Amer. (n. s.) 21: 214. d.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 195, fig. 26, map 19.
heathii Melander. Calif. (Pacific Grove).
Chyphotes Heathii Melander, 1903. Amer. Ent. Soc., Trans. 29: 326. ¢.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 194, fig. 25, map 18.
SPECIES GROUP PALLIDUS
buzickyi Mickel. Calif.; Mexico (Baja California).
Chyphotes (Pitanta) buzickyi Mickel, 1967. Amer. Ent. Soc., Trans. 93: 200, fig. 14. d, 2.

marginalis Mickel. Calif. (Imperial Co.).
Chyphotes (Pitanta) marginalis Mickel, 1967. Amer. Ent. Soc., Trans. 93: 196, fig. 27. ¢.

minimus Mickel. Ariz.; Mexico (Sonora).
Chyphotes minimus Mickel, 1963. Pan-Pacifie Ent. 39: 186. ¢.

mojave (Pate). Calif.
Pitanta mojave Pate, 1947. Notulae Nat. 192: 3. d.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 202, fig. 29. d, 2.
nitidus Mickel. Calif. (Winterhaven, Glamis).
Chyphotes nitidus Mickel, 1963. Pan-Pacific Ent. 39: 188. °.
Taxonomy: Mickel, 1974. Ent. Soc. Amer., Ann. 67: 461, fig. 1. ¢.
pallidus Buzicky. Ariz., Calif.
Chyphotes pallidus Buzicky, 1941. Ent. Amer. (n. s.) 21: 226. ¢.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 198, fig. 28, map 20.
SPECIES GROUP ALBIPES

albipes (Cresson). Alta., B. C., Idaho, Wash., Oreg., Calif., Nev., Utah, Ariz.
Agama albipes Cresson, 1874. Amer. Ent. Soc., Trans. 5: 99. d.
Chyphotes nevadensis Baker, 1905. Invertebrata Pacifica 1: 118. 6.

1280 Hymenoptera in America North of Mexico

Taxonomy: Buzicky, 1941. Ent. Amer. (n. s.) 21: 215-217. 2. —Mickel, 1967. Amer. Ent. Soc.,
Trans. 93: 206, fig. 34, map 21.
mickeli inyoensis Mickel. Calif. (Inyo Co.).
Chyphotes (Pitanta) mickeli inyoensis Mickel, 1967. Amer. Ent. Soc., Trans. 93: 213, fig.
3a. 6.
mickeli mickeli Buzicky. N. Mex., Ariz., Calif.; Mexico (Baja California).
Chyphotes mickeli mickeli Buzicky, 1941. Ent. Amer. (n. s.) 21: 217. 6.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 209, fig. 32, map 22. d, °.
mickeli polingi Buzicky. Tex., N. Mex.
Chyphotes mickeli polingi Buzicky, 1941. Ent. Amer. (n. s.) 21: 219. ¢.
Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 214.
SPECIES GROUP STRAMINEUS
stramineus Mickel. Ariz., Nev., Calif.; Mexico (Baja California).
Chyphotes (Pitanta) stramineus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 215, fig. 30, map
Z2au1Or

SPECIES GROUP MELLIPES

epedaphus Buzicky. Ariz., Calif.
Chyphotes epedaphus Buzicky, 1941. Ent. Amer. (n. s.) 21: 231. ?.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 232.
evansi Mickel. N. Mex.
Chyphotes (Pitanta) evansi Mickel, 1967. Amer. Ent. Soc., Trans. 93: 231. 9.

gracilis Mickel. Tex.; Mexico (Nuevo Leon).
Chyphotes (Pitanta) gracilis Mickel, 1967. Amer. Ent. Soc., Trans. 93: 225, fig. 37, map 26.
é.

mellipes (Blake). Wash., Oreg., Idaho, Calif., Nev., Utah, Colo., west. Kans., Tex., N. Mex.,

Ariz.; Mexico (Chihuahua, Durango, Zacatecas, San Luis Potosi, Jalisco).

Agama attenuata Blake, 1872. Amer. Ent. Soc., Trans. 4: 76. ¢.

Photopsis mellipes Blake, 1886. Amer. Ent. Soc., Trans. 13: 262. d.

Photopsis picus Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 289. d.

Mutilla picas (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 73.

Mutilla tenula Dalla Torre, 1897. Cat. Hym., v. 8, p. 91. N. name for Agama attenuata
Blake, not Mutilla attenwata Spinola.

Chyphotes punctatus Fox, 1899. Amer. Ent. Soc., Trans. 25: 276. 2.

Chyphotes striatus Buzicky, 1941. Ent. Amer. (n. s.) 21: 228. °.

Chyphotes segregatus Buzicky, 1941. Ent. Amer. (n. s.) 21: 229. °.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 216, fig. 31, map 24.

pixus Buzicky. Ariz. (Gila Bend).
Chyphotes pixus Buzicky, 1941. Ent. Amer. (n. s.) 21: 227. 2.

sonorus Mickel. Ariz.; Mexico (Sonora).
Chyphotes (Pitanta) sonorus Mickel, 1967. Amer. Ent. Soc., Trans. 93: 228, fig. 35, map 25.
3.

subulatus Buzicky. Ariz.
Chyphotes subulatus Buzicky, 1941. Ent. Amer. (n. s.) 21: 224. 6.

Taxonomy: Mickel, 1967. Amer. Ent. Soc., Trans. 93: 226, fig. 39.
SUBFAMILY TYPHOCTINAE
Revision: Krombein and Schuster, 1957. Ent. Soe. Wash., Proc. 59: 209-232, 2 pls.
All known species are diurnal insects.
Genus TYPHOCTES Ashmead

Typhoctes Ashmead, 1899. N. Y. Ent. Soc., Jour. 7: 53.
Type-species: Mutilla peculiaris Cresson. Orig. desig.

Superfamily SCOLIOIDEA 128]

Anommutilla Mickel, 1936. Ent. Soe. Amer., Ann. 29: 295.
Type-species: Anommutilla difficilis Mickel. Orig. desig.

Taxonomy: Brothers, 1970. Kans. Ent. Soc., Jour. 43: 305 (generic redescription, female).
Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 385-386, figs. 15-16 (female
thorax).

peculiaris mirabilis (Cockerell). Kans., Colo., Tex., N. Mex., Ariz.; Mexico (Chihuahua).
Chyphotes mirabilis Cockerell, 1896. Canad. Ent. 28: 285. 9.

peculiaris peculiaris (Cresson). Wash., Idaho, Utah, Ariz., Calif.
Mutilla peculiaris Cresson, 1875. Amer. Ent. Soc., Trans. 5: 119. @.
Anommiutilla difficilis Mickel, 1936. Ent. Soc. Amer., Ann. 29: 295. d.
Morphology: Hermann, 1975. Ent. Soc. Amer., Ann. 68: 882-884, 5 figs. (sting).

striolatus Krombein and Schuster. Ariz. to western Tex.
Typhoctes striolatus Krombein and Schuster, 1957. Ent. Soc. Wash., Proc. 59: 223, fig. 2. °,
3.

williamsi Krombein and Schuster. Southern Calif.
Typhoctes williamsi Krombein and Schuster, 1957. Ent. Soc. Wash., Proc. 59: 220, figs. 1, 4.
One

SUBFAMILY MUTILLINAE

TRIBE MUTILLINI

Blake (1886. Amer. Ent. Soc., Trans. 13) described three species of Mutilla, mislabeled as com-
ing from Florida, from the Harris collection. Fox, 1898 (Ent. News 9: 14) stated that M. ajax was
a synonym of the African Psammotherma flabellata (F.), M. floridana a synonym of the Eu-
ropean Dasylabris maura (L.), and M. trisignata a synonym of the European D. arenaria (F.).

Genus TIMULLA Ashmead

Genus TIMULLA Subgenus TIMULLA Ashmead

Timulla Ashmead, 1899. N. Y. Ent. Soce., Jour. 7: 55.
Type-species: Mutilla dubitata Smith. Orig. desig.

Revision: Mickel, 1937. Ent. Amer. (n. s.) 17: 1-119 (N. Amer. spp.).

Only the typical subgenus occurs in the New World. Most species are diurnal, but a few spe-
cies with enlarged ocelli, such as ocellaria Mick., are crepuscular or nocturnal.

barbata (Fox). Mo., La.
Mutilla barbata Fox, 1899. Amer. Ent. Soc., Trans. 25: 272. 6.

barbigera barbigera (Bradley). Va. south to Fla., west to S. Dak. and Tex. in U. and L. Austr.
Zones.

Mutilla (Timulla) barbigera Bradley, 1916. Amer. Ent. Soc., Trans. 42: 206. ¢.
barbigera rohweri Mickel. Kans., Okla. in U. Sonor. Zone.

Timulla (Timulla) barbigera rohweri Mickel, 1937. Ent. Amer. (n. s.) 17: 58. 6.
compressicornis Mickel. N. C. to Tex. in L. Austr. Zone.

Timulla (Timulla) compressicornis Mickel, 1937. Ent. Amer (n. s.) 17: 59. 6.
contigua Mickel. Tex. (Brownsville).

Timulla (Timulla) contigua Mickel, 1937. Ent. Amer. (n. s.) 17: 107. 9.
dubitata dubitata (Smith). N. Y. south to Fla., west to Okla. and Tex. in L. Austr. Zone.

Mutilla dubitata Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 60. °.

Mutilla dubiatta (!) Blake, 1886. Amer. Ent. Soc., Trans. 13: 201. @.

Biology: Sheldon, 1970. Ent. News 81: 59-60 (mating behavior).
dubitata fugitiva Mickel. N. J. south to Ga., west to Nebr. and Tex. in U. Austr. Zone.
Timulla (Timulla) dubitata fugitiva Mickel, 1937. Ent. Amer. (n. s.) 17: 39. d.
dubitatiformis Mickel. Mass. south to Fla., west to Mont. and Tex.
Timulla (Timulla) dubitatiformis Mickel, 1937. Ent. Amer. (n. s.) 17: 102. 9.

1282 Hymenoptera in America North of Mexico

euphrosyne Mickel. Fla., Tex.
Timulla (Timulla) euphrosyne Mickel, 1937. Ent. Amer. (n. s.) 17: 64. @.
euterpe (Blake). N. C., Fla., La., Tex.
Mutilla Euterpe Blake, 1879. Amer. Ent. Soc., Trans. 7: 249. °.
ferrugata (Fabricius). N. J. south to Fla., west to Okla. and Tex. in U. and L. Austr. Zones.
Host: Eumenes fraternus Say.
Mutilla ferrugata Fabricius, 1805. Systema Piezatorum, p. 438. 2.
Mutilla rufa Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 631. ¢.
Mutilla Promethea Blake, 1871. Amer. Ent. Soc., Trans. 3: 229. 3.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 3 (host).
floridensis (Blake). S. C. to Tex. in L. Austr. Zone.
Mutilla floridensis Blake, 1879. Amer. Ent. Soc., Trans. 7: 249. 6.
grotei (Blake). Colo., Utah, Tex., N. Mex., Ariz.; Mexico (Jalisco, Guanajuato).
Mutilla Grotei Blake, 1871. Amer. Ent. Soc., Trans. 3: 228. 6.
Mutilla fulviventris Gerstaecker, 1874. Arch. Naturgesch. 40: 323. ¢.

Taxonomy: Mickel, 1937. Ent. Amer. (n. s.) 17: 78.

hollensis hollensis (Melander). Mass., N. Y., Md.

Mutilla Sayi var. hollensis Melander, 1903. Amer. Ent. Soc., Trans. 29: 324. ¢.
hollensis melanderi Mickel. N. J., Pa., Md., Va.

Timulla (Timulla) hollensis melanderi Mickel, 1937. Ent. Amer. (n. s.) 17: 101. 6.
huntleyensis Mickel. Mont. (Huntley).

Timulla (Timulla) huntleyensis Mickel, 1937. Ent. Amer. (n. s.) 17: 82. 6.
kansana Mickel. Kans.

Timulla (Timulla) kansana Mickel, 1937. Ent. Amer. (n. s.) 17: 94. 3.
leona (Blake). Wis. south to Fla., west to Nebr. and Tex. in U. and L. Austr. Zones. Host:

Bembix troglodytes Handl.?
Mutilla Leona Blake, 1871. Amer. Ent. Soc., Trans. 3: 230. 9.

Biology: Evans, 1957. Studies in Compar. Ethology Bembix, p. 134, fig. 36 (entering burrows
of B. troglodytes).
navasota coahuila Krombein. Tex. west of 100th Meridian, N. Mex., Ariz.
Timulla (Timulla) navasota nebulosa Mickel, 1937. Ent. Amer. (n. s.) 17: 29. 3d. Preoce. by
Mickel, 1935.
Timulla (Timulla) navasota coahuila Krombein, 1951. U. S. Dept. Agr., Monog. 2: 771. N.
name.
navasota navasota (Bradley). Tex. east of 100th Meridian.
Mutilla (Timulla) navasota Bradley, 1916. Amer. Ent. Soc., Trans. 42: 213. 6.
neobule Mickel. Ariz.
Timulla (Timulla) neobule Mickel, 1937. Ent. Amer. (n. s.) 17: 44. ¢.
nicholi Mickel. Ariz.
Timulla (Timulla) nicholi Mickel, 1937. Ent. Amer. (n. s.) 17: 47. 2.
nitela Mickel. Ariz. (Douglas).
Timulla (Timulla) nitela Mickel, 1937. Ent. Amer. (n. s.) 17: 43. ¢.
oajaca (Blake). Tex., N. Mex., Ariz.; Mexico (Morelos, Jalisco, Colima).
Mutilla oajaca Blake, 1871. Amer. Ent. Soc., Trans. 3: 228. ¢ (2 misdet.).
Mutilla ardens Gerstaecker, 1874. Arch. Naturgesch. 40: 323. ¢.
Mutilla mazatlanae Cameron, 1894. Biol. Cent.-Amer., Hym., v. 2, p. 294. ¢.
Mutilla nestor Fox, 1899. Amer. Ent. Soc., Trans. 25: 271. ¢.
Mutilla ornata Howard, 1901. Insect Book, pl. 8, fig. 21. 9.

Taxonomy: Mickel, 1938. Roy. Ent. Soc. London, Trans. 87: 641.

Biology: Linsley, 1960. Pan-Pacific Ent. 36: 36 (mating behavior).

ocellaria ocellaria Mickel. Ohio, Ind., Ky., Tenn., Ga. Host: Ligyrus gibbosus (Deg.), pupae.
Timulla (Timulla) ocellaria ocellaria Mickel, 1937. Ent. Amer. (n. s.) 17: 91. 6.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 5 (host record).

Superfamily SCOLIOIDEA 1283

ocellaria rufidorsa Mickel. Miss., Ark., Mo., Kans., Okla., Tex.

Timulla (Timulla) ocellaria rufidorsa Mickel, 1937. Ent. Amer. (n. s.) 17: 93. 6.
ornatipennis (Bradley). N. J. south to Fla. west to Tenn. and Miss.

Mutilla (Timulla) ornatipennis Bradley, 1916. Amer. Ent. Soc., Trans. 42: 205. °, d.
rufosignata (Bradley). N. C., Ga., Fla. Host: Ground-nesting eumenid wasp.

Mutilla (Timulla) rufosignata Bradley, 1916. Amer. Ent. Soc., Trans. 42: 212. ¢.

Taxonomy: Krombein, 1953. Ent. Soe. Wash., Proc. 55: 128. 9.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 6 (ground-nesting host misdet. as Odynerus
erinnys (!) Lep.).
sayi (Blake). Tex. (Plano).
Mutilla Sayi Blake, 1871. Amer. Ent. Soe., Trans. 3: 229. 3.
subhyalina Mickel. Ill., lowa, Minn., N. Dak., S. Dak., Nebr., Kans., Mont., Oreg., B. C.
Timulla (Timulla) subhyalina Mickel, 1937. Ent. Amer. (n. s.) 17: 97. 3.
suspensa jonesi Mickel. Tex., Ariz.
Timulla (Timulla) suspensa jonesi Mickel, 1937. Ent. Amer. (n. s.) 17: 90. d.
suspensa sonora Mickel. Kans. and Colo. south to Tex. and Ariz.
Timulla (Timulla) suspensa sonora Mickel, 1937. Ent. Amer. (n. s.) 17: 84. 2, 6.
suspensa suspensa (Gerstaecker). Ariz.
Mutilla suspensa Gerstaecker, 1874. Arch. f. Naturgesch. 40: 299. °.
tolerata Mickel. Mo., Ga.
Timulla (Timulla) tolerata Mickel, 1937. Ent. Amer. (n. s.) 17: 109. 3.
tyro Mickel. Ariz., Calif.
Timulla (Timulla) tyro Mickel, 1937. Ent. Amer. (n. s.) 17: 25. 9, 6.
vagans rufinota Mickel. D. C., S. C., Ga., Fla.
Timulla (Timulla) vagans rufinota Mickel, 1937. Ent. Amer. (n. s.) 17: 78. 2, 6.
vagans vagans (Fabricius). Vt. and Ont. south to Ala., west to B. C. and N. Mex.
Mutilla vagans Fabricius, 1798. Sup. Ent. System., p. 282. 9.
Mutilla (Mutilla) hexagona Say, 1836. Boston Jour. Nat. Hist. 1: 295. 3.
Mutilla ornativentris Cresson, 1865. Ent. Soc. Phila., Proc. 4: 438. 9.
Mutilla Briaxus Blake, 1871. Amer. Ent. Soe., Trans. 3: 227. 6.
Mutilla Canadensis Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 250. ¢.
Preoce.
Mutilla secunda Dalla Torre, 1897. Cat. Hym., v. 8, p. 84. N. name for Mutilla Canadensis
Provancher, not Mutilla canadensis Blake.

Biology: Fattig, 1936. Ent. News 47: 51-52 (mating behavior). —Shappirio, 1948. Brooklyn
Ent. Soc., Bul. 42: 163 (mating behavior).

wileyae Mickel. Ark., La., Okla., Tex.
Timulla (Timulla) wileyae Mickel, 1937. Ent. Amer. (n. s.) 17: 41. @.

TRIBE EPHUTINI

Genus EPHUTA Say

Revision: Bradley, 1916. Amer. Ent. Soc., Trans. 42: 187-198. —Schuster, 1951. N. Y. Ent.
Soc., Jour. 59: 1-43, 5 pls. (Part I containing only gen. diag. and key to spp.). —Schuster,
1957. N. Y. Ent. Soc., Jour. 64: 7-84 (Part II of revis. treating Species Group Grisea).

All of the species are diurnal.
Genus EPHUTA Subgenus EPHUTA Say

Ephuta Say, 1836. Boston Jour. Nat. Hist. 1: 297.
Type-species: Mutilla (Ephuta) scrupea Say. Desig. by Ashmead, 1899.
Rhoptromutilla Andre, 1903. In Wytsman, Gen. Ins., fase. 11, p. 43.
Type-species: Mutilla chrysodora Perty. Orig. desig.
Ephutopsis Ashmead, 1904. Canad. Ent. 36: 6.
Type-species: Mutilla odontophora Cameron. Desig. by Mickel, 1928.

1284 Hymenoptera in America North of Mexico

The few authenticated host records indicate that species of this subgenus have various species
of Pompilidae as hosts. The spider wasp hosts have diverse nesting habits. Some nest in borings
in wood (Dipogon), some construct mud cells beneath stones (Phanagenia), and some nest in the
ground (? Anoplius fraternus (Bks.)).
albiceps Schuster. Tex. (Tyler).

Ephuta albiceps Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 31 (in key). 2.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 75 (female description).
argenticeps Schuster. Calif.

Ephuta californica Schuster, 1951. N. Y. Ent. Soc., Jour. 59: footnote on p. 25. d (On p. 15
Schuster states that there is only one sp. in Calif., and in key to females, argenticeps is
the only sp. stated to be from Calif.).

Ephuta argenticeps Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 26, 32 (in key). 6, @.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 16 (male, female description).
auricapitis Schuster. Tex. (Edinburgh). Possibly the opposite sex of ecarinata ecarinata

Schuster.
Ephuta auricapitis Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 31 (in key). 9.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 76 (female description).
baboquivari Schuster. Ariz. (Baboquivari Mts.)
Ephuta baboquivari Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 31 (in key), figs. 39, 41. @.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 79 (female description).
battlei battlei Bradley. Coastal N. C., Ga., Fla.
Ephuta battlei Bradley, 1916. Amer. Ent. Soc., Trans. 42: 195. ¢.
Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 29, figs. 5, 44 (in key; reduced to subsp.
rank).

battlei confusa Schuster. Piedmont and west. coastal plain, N. C., Ga., Ala.
Ephuta battlei confusa Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 29 (in key), fig. 44. 3.

battlei microcellaria Schuster. N. J. (Cape May).
Ephuta battlei microcellaria Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 29 (in key), fig. 44.
3.

battlei transitionalis Schuster. Transit. and U. Austr. Zones, Mass. to Va., Ohio.
Ephuta battlei transitionalis Schuster, 1951. N. Y. Ent. Soe., Jour. 59: 28 (in key), fig. 44.

3.
cephalotes Schuster. N. Dak., S. Dak., Wyo., Ariz., west. Tex. The opposite sex is possibly
minuta Schuster.

E phuta cephalotes Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 24 (in key), figs. 10, 20, 46. 3.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 68 (male description).
coloradella Schuster. Colo. Possibly the opposite sex of grisea Bradley.
Ephuta coloradella Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 32 (in key). 2.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 73 (female description).
conchate Mickel. N. Y., Mich., IIll., lowa, Minn., S. Dak., Nebr., Kans.
Ephuta conchate Mickel, 1928. Minn. State Ent., Rpt. 19: 111. ¢.
Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 32 (in key).

copano (Blake). Tex.; Mexico.
Mutilla Copano Blake, 1871. Amer. Ent. Soc., Trans. 3: 282. 6.
Mutilla susura Melander, 1903. Amer. Ent. Soc., Trans. 29: 324. 6.

Taxonomy: Schuster, 1958. Jn Krombein, U. S. Dept. Agr., Monog. 2, Sup. 1: 105 (synonymy).
ecarinata ecarinata Schuster. Tex.; Mexico. The opposite sex is possibly swdatrix (Melander)

or auricapitis Schuster.
Ephuta ecarinata ecarinata Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 26 (in key). 6.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 49 (male description).

Superfamily SCOLIOIDEA 1285

ecarinata neomexicana Schuster. N. Mex.
Ephuta ecarinata neomexicana Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 26 (in key). d.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 52 (male description).
ecarinata pima Schuster. Ariz.
Ephuta ecarinata pima Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 26 (in key). d.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 51 (male description).
eurygnathus Schuster. N. C. (Southern Pines).
Ephuta eurygnathus Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 29 (in key), fig. 11. 6.
floridana dietrichi Schuster. Ala., Miss.
Ephuta floridana dietrichi Schuster, 1951. N. Y. Ent. Soe., Jour. 59: 32 (in key), figs. 35,
Ose

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 40 (female description).

floridana floridana Schuster. Fla.
Ephuta floridana floridana Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 25, 32 (in key). 3, @.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 34 (male. female description).
grisea fuscosericea Schuster. Alta., Mont., Utah, N. Dak.
Ephuta grisea fuscosericea Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 27 (in key), figs. 14,
15, 42. 6.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 14 (male description).

grisea grisea Bradley. Colo. The opposite sex is possibly coloradella Schuster.
Ephuta grisea Bradley, 1916. Amer. Ent. Soc., Trans. 42: 194. ¢.

Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 27, fig. 21 (reduced to subsp. rank).
—Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 13 (male description).
margueritae margueritae Schuster. Fla.
Ephuta margueritae margueritae Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 25, 30 (in key),
figs. 17, 46. 6, @.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 56 (male, female description).

margueritae xanthocephala Schuster. Pa., N. C., Tenn. Host: “cocoons under stones”,
presumably mud cells of Phanagenia bombycina (Cr.) or of a species of Auplopus.
Ephuta margueritae xanthocephala Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 25, 30 (in
key), figs. 25, 46. d, 2.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 63 (male, female description).

Biology: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 14 (rearing note).
minuta minuta Schuster. Tex. (Brownsville, Fedor, College Sta.). Possibly the opposite sex of
cephalotes Schuster.
Ephuta minuta minuta Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 33 (in key). 9.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 71 (tentative sex association).
minuta modesta Schuster. Ariz. (Tempe).
Ephuta minuta modesta Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 32 (in key), figs. 31, 32.
OF
pauxilla pauxilla Bradley. Minn. to Ala., east to Maine and Fla. Host: Dipogon s. sayi Bks.

Almost certainly the male of puteola (Blake).
Ephuta pauxilla Bradley, 1916. Amer. Ent. Soc., Trans. 42: 197. d.

Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 29, figs. 1, 12, 37, 43 (in key; reduced to
subsp. rank).
Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 104 (host record).

pauxilla texanella Schuster. Tex., La., Miss.
Ephuta pauxilla texanella Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 29 (in key), figs. 13,
43. 6.
psephenophila Schuster. Ga. (Stone Mt.) The opposite sex is possibly tentativa Schuster.
Ephuta psephenophila Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 25 (in key). ¢.

1286 Hymenoptera in America North of Mexico

Ephuta ocellaria Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 36, fig. 18. MS name changed to
psephenophila (teste R. M. Schuster).

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 44 (male description; tentative sex
association).
puteola (Blake). Mass. south to Fla., west to Iowa, Tex. Possibly the opposite sex of p.
pauxilla Brad.
Mutilla puteola Blake, 1879. Amer. Ent. Soe., Trans. 7: 252. 9.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 40 (female description in key; putative
sex association).
rufisquamis (Andre). Ariz., south Calif. The opposite sex is possibly tumacacori Schuster.
Rhoptromutilla rufisquamis Andre, 1905. Ztschr. System. Hym. Dipt. 5: 366. 3.

Taxonomy: Schuster, 1951. N. Y. Ent. Soe., Jour. 59: 24 (in key). —Schuster, 1957. N. Y. Ent.
Soc., Jour. 64: 65 (male description; tentative sex association).
sabaliana fattigi Schuster. Ga. (Tifton).
Ephuta sabaliana fattigi Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 28 (in key). 6.

sabaliana sabaliana Schuster. Fla. Host: Anoplius fraternus (Bks.)?
Ephuta sabaliana sabaliana Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 28 (in key). 3.

Biology: Krombein, 1955. Ent. Soc. Wash. Proc. 57: 225-226.
scrupea (Say). N. J. south to Ga. and Ala., Mich., Wis., Ohio, W. Va., Mo., Ark., Tex. Host:
Phanagenia bombycina (Cr.).
Mutilla (Ephuta) scrupea Say, 1836. Boston Jour. Nat. Hist. 1: 297. ¢.
Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 25, 34, figs. 2-4, 27-30, 42 (male, female
in key). —Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 26 (male, female description).
slossonae monochroa Schuster. Tex. (Rock Island).
Ephuta slossonae monochroa Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 34 (in key). °.
slossonae slossonae (Fox). Fla.
Mutilla Slossonae Fox, 1899. Amer. Ent. Soc., Trans. 25: 273. 6.
Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 34 (in key; reduced to subsp. rank).
spinifera Schuster. N. J. to Ga.
Ephuta spinifera Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 27, 34 (in key). 6, 9.
stenognatha psephenophora Schuster. North. Fla.
Ephuta stenognatha psephenophora Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 28. 6.
stenognatha stenognatha Schuster. Cent. and south. Fla.
Ephuta stenognatha stenognatha Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 28 (in key), figs.
7,8. 6.
sudatrix (Melander). Tex. (Fedor). Possibly the opposite sex of ecarinata ecarinata Schuster.
Mutilla sudatrix Melander, 1903. Amer. Ent. Soc., Trans. 29: 325. °.
Taxonomy: Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 30 (in key). —Schuster, 1957. N. Y. Ent.
Soc., Jour. 64: 54 (female description; tentative sex association).

tegulicia Bradley. Tex., Ariz. Possibly a syn. or subsp. of idiasta (Cameron) from Mexico.
Ephuta tegulicia Bradley, 1916. Amer. Ent. Soc., Trans. 42: 193. 3.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 47 (male description).

tentativa Schuster. Ga. (Atlanta, Ochlochnee). Possibly the opposite sex of psephenophila
Schuster.
Ephuta tentativa Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 34 (in key). @.

Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 42 (female description; tentative sex
association).
tumacacori Schuster. Ariz. (Tumacacori Mts.). Possibly the opposite sex of rufisquamis
(Andre).
Ephuta tumacacori Schuster, 1951. N. Y. Ent. Soc., Jour. 59: 30 (in key). @.
Taxonomy: Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 82 (female description; tentative sex
association).

Superfamily SCOLIOIDEA 1287

Genus EPHUTA Subgenus XENOCHILE Schuster

Ephuta subg. Xenochile Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 8.
Type-species: Ephuta (Xenochile) krombeini Schuster. Orig. desig.

krombeini Schuster. Ariz. (Ramsey Canyon in Huachuca Mts.).
Ephuta (Xenochile) krombeini Schuster, 1957. N. Y. Ent. Soc., Jour. 64: 9. d.

SUBFAMILY SPHAEROPTHALMINAE

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul. Biol. Ser. 8, no. 4: 1-26, 7 figs.
(males of Nevada Test Site).

TRIBE SPHAEROPTHALMINI

Taxonomy: Schuster, 1958. Ent. Amer. (n. s.) 37: 1-130, 7 pls. (Part II of a tribal revision of
males containing keys to genera and subgenera and revisions of Acrophotopsis Schuster,
Dilophotopsis Schuster and Acanthophotopsis Schuster).

The majority of species in this tribe are nocturnal, but all members of Morsyma Fox,
Sphaeropthalma subg. Sphaeropthalma Blake, and Protophotopsis Schuster, and most species of
Photomorphus subg. Photomorphus Viereck, are diurnal.

Genus MORSYMA Fox

Morsyma Fox, 1899. Amer. Ent. Soc., Trans. 25: 287.
Type-species: Morsyma Ashmeadii Fox. Orig. desig.

ashmeadii Fox. Calif.
Morsyma Ashmeadii Fox, 1899. Amer. Ent. Soe., Trans. 25: 287. “2” = 6.

Taxonomy: Krombein, 1940. Amer. Ent. Soc., Trans. 65: 420.
Genus ACROPHOTOPSIS Schuster

Acrophotopsis Schuster, 1958. Ent. Amer. (n. s.) 37: 4 (in key), 61.
Type-species: Acrophotopsis eurygnathus Schuster. Orig. desig.
campylognathus Schuster. Calif. (Riverside Co.); Mexico (Baja California).
Acrophotopsis campylognathus Schuster, 1958. Ent. Amer. (n. s.) 37: 11 (in key), 69. d.
eurygnathus Schuster. Ariz., Nev.
Acrophotopsis eurygnathus Schuster, 1958. Ent. Amer. (n. s.) 37: 10 (in key), 65, pl. 1, figs.
M25 Or

Genus DILOPHOTOPSIS Schuster

Dilophotopsis Schuster, 1958. Ent. Amer. (n. s.) 37: 5 (in key), 71.
Type-species: Mutilla concolor Cresson. Orig. desig.
concolor concolor (Cresson). Kans. to Colo. and Wyo., Tex., N. Mex., Ariz.; Mexico (Zacatecas,

Durango, Jalisco).
Mutilla concolor Cresson, 1865. Ent. Soe. Phila., Proc. 4: 390. 6.
Odontophotopsis alamonis Viereck, 1904. Amer. Ent. Soc., Trans. 30: 87. ¢.

Taxonomy: Schuster, 1958. Ent. Amer. (n. s.) 37: 84 (reduction to subsp. rank and syn.).

concolor crassa (Viereck). B. C., Mont., southwest. Idaho, west. Colo., Nev., Utah, Ariz., Calif.,

Wash.
Odontophotopsis crassus Viereck, 1924. Canad. Ent. 56: 112. d.
Dilophotopsis concolor utahensis Schuster, 1958. Ent. Amer. (n. s.) 37: 84 (in key), 87. d.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 8 (synonymy).

concolor laredo Schuster. Tex. (Winterhaven, Laredo, Cotulla).
Dilophotopsis concolor laredo Schuster, 1958. Ent. Amer. (n. s.) 37: 84 (in key), 86. d.
concolor paron (Cameron). Nev., Ariz., Calif.; Mexico (Sonora, Baja California).
Sphaerophthalma (!) paron Cameron, 1896. Biol. Cent.-Amer., Hym., v. 2, p. 381. 3.
Dilophotopsis concolor sonorensis Schuster, 1958. Ent. Amer. (n. s.) 37: 84 (in key), 88. 6.

1288 Hymenoptera in America North of Mexico

Taxonomy: Mickel, 1965. Ent. Soc. Wash., Proe. 67: 1 (synonymy).

stenognatha Schuster. Ariz. (Tucson). :
Dilophotopsis stenognatha Schuster, 1958. Ent. Amer. (n. s.) 37: 11 (in key), 74. d.

Taxonomy: Mickel, 1963. Pan-Pacific Ent. 39: 184 (female description).
Genus ACANTHOPHOTOPSIS Schuster

Acanthophotopsis Schuster, 1958. Ent. Amer. (n. s.) 37: 5 (in key), 88.
Type-species: Acanthophotopsis falciformis Schuster. Orig. desig.
bequaertii Schuster. Ariz. (Hereford).
Acanthophotopsis bequaertii Schuster, 1958. Ent. Amer. (n. s.) 37: 12 (in key), 101. ¢.
bifurca Schuster. Okla., Tex., N. Mex.
Acanthophotopsis bifurca Schuster, 1958. Ent. Amer. (n. s.) 37: 18 (in key), 98, pl. 3, fig. 3.
3.
dorophora Schuster. Ariz. (Tucson, Yuma).
Acanthophotopsis dorophora Schuster, 1958. Ent. Amer. (n. s.) 37: 13 (in key), 104, pl. 2,
fig. 1, pl. 3, fig. 2. 6.
evansii Schuster. Tex. (Alpine and Chisos Mts.); Mexico (Durango).
Acanthophotopsis evansii Schuster, 1958. Ent. Amer. (n. s.) 37: 12 (in key), 93, pl. 3, fig. 1.
3.

falciformis falciformis Schuster. Nev., Calif.; Mexico (Zacatecas).
Acanthophotopsis falciformis falciformis Schuster, 1958. Ent. Amer. (n. s.) 37: 13 (in key),
108, pl. 2, figs. 2, 3. 3.
falciformis furcisterna Schuster. Ariz. (Tucson, Phoenix, Ajo).
Acanthophotopsis falciformis furcisterna Schuster, 1958. Ent. Amer. (n. s.) 37: 14 (in key),
ese

Genus SPHAEROPTHALMA Blake

In addition to the host records cited under several species in this genus, Parker and Bohart,
1968 (Pan-Pacifiec Ent. 44: 2) record an unidentified species of Sphaeropthalma from a twig nest
of Leptochilus washo Parker, and Hurd and Powell, 1958 (Pan-Pacific Ent. 34: 152-153) record an
unidentified female of this species as leaving a burrow of Colletes stepheni Timb.

Genus SPHAEROPTHALMA Subgenus MICROMUTILLA Ashmead

Micromutilla Ashmead, 1899. N. Y. Ent. Soc., Jour. 7: 59.
Type-species: Photopsis nanus Ashmead. Orig. desig.
acontia (Fox). N. Mex., Ariz., Nev.
Photopsis nanus Ashmead, 1896. Amer. Ent. Soc., Trans. 23: 181. ¢. Preoce.
Mutilla acontius Fox, 1899. Amer. Ent. Soc., Trans. 25: 266. d.
Mutilla Ashmeadii Fox, 1899. Amer. Ent. Soc., Trans. 25: 289. N. name.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 9 (synonymy).
becki Ferguson. Nev., Calif.
Sphaeropthalma (Micromutilla) becki Ferguson, 1967. Brigham Young Univ. Sci. Bul.,
Biol. Ser. 8, no. 4: 9, fig. 5. ¢.
bellerophon (Fox). Tex., N. Mex., Ariz.
Mutilla bellerophon Fox, 1899. Amer. Ent. Soc., Trans. 25: 254. ¢.
brachyptera Schuster. Calif., Nev., Ariz.; Mexico (Sonora).
Photopsis brachyptera Schuster, 1945. Pan-Pacific Ent. 21: 149. 6.
Sphaeropthalma (Micromutilla) yavapai Schuster, 1958. Ent. Amer. (n. s.) 37: 19. d.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 11 (synonymy).
difficilis (Baker). Calif., Nev., Ariz., N. Mex.; Mexico (Baja California). Host: Auplopus
architectus metallicus (Bks.).
Photopsis difficilis Baker, 1905. Invertebrata Pacifica 1: 114. ¢.
Sphaeropthalma (Micromutilla) maricopella purismella Schuster, 1958. Ent. Amer. (n. s.)
Senor

Superfamily SCOLIOIDEA 1289

Sphaeropthalma (Microm utilla) maricopella maricopella Schuster, 1958. Ent. Amer. (n. s.)
eee ities (Micromutilla) maricopella castanea Schuster, 1958. Ent. Amer. (n. s.)

Seine (Micromutilla) californiense californiense Schuster, 1958. Ent. Amer. (n.
ee nie (Micromutilla) californiense fuscatella Schuster, 1958. Ent. Amer. (n. s.)
Sphacrthaine (Micromutilla) quijotoa quijotoa Schuster, 1958. Ent. Amer. (n. s.) 37: 18.

Sphaeropthalma (Mcromutilla) quijotoa parrasia Schuster, 1958. Ent. Amer. (n. s.) Bie Iles,
Or

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 11 (synonymy).

Biology: Ferguson, 1962. Univ. Calif. Pubs. Ent. 27: tab. 10 (host record).
hyalina (Blake). Tex.
Agama hyalina Blake, 1871. Amer. Ent. Soc., Trans. 3: 263. d.
Agama minuta Blake, 1872. Amer. Ent. Soc., Trans. 4: 76. d.
macswaini Ferguson. Nev., Calif.
Sphaeropthalma (Micromutilla) macswaini Ferguson, 1967. Brigham Young Univ. Sci.
Bul., Biol. Ser. 8, no. 4: 12, fig. 6. d.
mesillensis (Cockerell). N. Mex. (Mesilla).
Photopsis mesillensis Cockerell, 1897. Entomologist 30: 137. d.
pallida (Blake). Tex., N. Mex., Ariz., Nev.
Agama pallida Blake, 1871. Amer. Ent. Soc., Trans. 3: 263. d.
Sphaeropthalma (Micromutilla) arizonae Schuster, 1958. Ent. Amer. (n. s.) 37: 16. do.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 13 (synonymy).
parapenalis Ferguson. Wash., Oreg., Idaho, Calif., Nev., Utah, Ariz., Tex.; Mexico (Chihuahua,
Coahuila, Durango, Zacatecas).
Sphaeropthalma (Micromutilla) parapenalis Ferguson, 1967. Brigham Young Univ. Sci.
Bul., Biol. Ser. 8, no. 4: 14, fig. 7. d.
pateli Schuster. Ariz. (Hereford).
Sphaeropthalma (Micromutilla) pateli Schuster, 1958. Ent. Amer. (n. s.) 37: 15. 6.
reducta Schuster. Ariz.
Sphaeropthalma (Micromutilla) reducta Schuster, 1958. Ent. Amer. (n. s.) 37: 18. d.
sabino Schuster. Ariz.
Sphaeropthalma (Micromutilla) sabino Schuster, 1958. Ent. Amer. (n. s.) Shel o er)
sonora Schuster. Ariz., Nev., Calif.
Sphaeropthalma (Micromutilla) sonora Schuster, 1958. Ent. Amer. (n. s.) 37: 16. 6.

sublobata Schuster. Idaho.
Sphaeropthalma (Micromutilla) sublobata Schuster, 1958. Ent. Amer. (n. s.) 37: 16. 6.

uvaldella Schuster. Tex.
Sphaeropthalma (Micromutilla) uvaldella Schuster, 1958. Ent. Amer. (n. s.) 37: 19. d.

yumaella Schuster. Nev., Ariz., Calif.; Mexico (Baja California).
Sphaeropthalma (Micromutilla) ywmaella Schuster, 1958. Ent. Amer. (n. s.) 37: 19. d.

Genus SPHAEROPTHALMA Subgenus PHYSETAPSIS Schuster
Sphaeropthalma subg. Physetapsis Schuster, 1958. Ent. Amer. (n. s.) 37: 9 (in key), 20.

Type-species: Sphaeropthalma (Physetapsis) papaga Schuster. Orig. desig.

borealis Schuster. Sask.
Sphaeropthalma (Physetapsis) borealis Schuster, 1958. Ent. Amer. (n. s.) 37: 20. d.

ecarinata Schuster. Calif.
Sphaeropthalma (Physetapsis) ecarinata Schuster, 1958. Ent. Amer. (n. s.) 37: 20. ¢.

1290 Hymenoptera in America North of Mexico

papaga ephysetos Schuster. Southwestern Ariz., southern Calif.
Sphaeropthalma (Physetapsis) papaga ephysetos Schuster, 1958. Ent. Amer. (n. s.) 37: 20.
3

papaga papaga Schuster. Southeastern Ariz., Calif. (Imperial Co.).
Sphaeropthalma (Physetapsis) papaga papaga Schuster, 1958. Ent. Amer. (n. s.) 37: 20, pl.
Guhionowor
subcarinata Schuster. N. Mex.
Sphaeropthalma (Physetapsis) subcarinata Schuster, 1958. Ent. Amer. (n. s.) 37: 20. 3.

Genus SPHAEROPTHALMA Subgenus PHOTOPSIS Blake

Agama Blake, 1871. Amer. Ent. Soc., Trans. 3: 258. Preoce.

Type-species: Agama imperialis Blake. Automatic desig. by Ashmead, 1899.
Photopsis Blake, 1886. Amer. Ent. Soc., Trans. 13: 179. N. name.

Type-species: Agama imperialis Blake. Desig. by Ashmead, 1899.
Pyrrhomutilla Ashmead, 1899. N. Y. Ent. Soe., Jour. 7: 56.

Type-species: Sphaerophthalma (!) anthophorae Ashmead. Orig. desig.
Neophotopsis Ashmead, 1903. Canad. Ent. 35: 306.

Type-species: Mutilla pluto Fox. Orig. desig.

In addition to the host records cited under several species in this subgenus, Parker and
Bohart, 1968 (Pan-Pacific Ent. 44: 2, 3 and 5) record several unidentified species of
Sphaeropthalma (Photopsis) from the following hosts from nests in twigs: Ancistrocerus cat-
skill (Sauss.); Odynerus erythrogaster Bohart; Pisonopsis birkmanni Rohwer; Anthocopa
hypostomalis Michener; Hoplitis fulgida (Cresson); H. sambuci Titus; and Proteriades bullifa-
cies (Michener). Ferguson, 1962 (Univ. Calif. Pubs. Ent. 27: tab. 9) states that Davidson reared
an unidentified species of Sphaeropthalma (Photopsis) from a nest of Hoplitis producta
(Cresson) in a hollow plant stem.

Biology: Ferguson, 1962. Univ. Calif. Pubs. Ent. 27: 1-92, 7 pls., 2 text figs. (life history,

hosts).
albopilosa (Blake). Tex. (Comal Co.).

Mutilla (Sphaeropthalma) albopilosa Blake, 1872. Amer. Ent. Soc., Trans. 4: 74. °.
anaspasia (Cockerell and Rohwer). Colo. (Boulder).

Photopsis anaspasia Cockerell and Rohwer, 1908. Psyche 15: 5. 2.
angulifera Schuster. Calif., Nev.

Sphaeropthalma (Photopsis) angulifera Schuster, 1958. Ent. Amer. (n. s.) 37: 32. 6.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 17
(distribution).
arenicola Schuster. Ariz.
Sphaeropthalma (Photopsis) arenicola Schuster, 1958, Ent. Amer. (n. s.) 37: 25. 3.
arota (Cresson). Calif. (San Diego).
Mutilla Arota Cresson, 1875. Amer. Ent. Soc., Trans. 5: 120. 2.
baboquivari Schuster. Ariz.
Sphaeropthalma (Photopsis) baboquivari Schuster, 1958. Ent. Amer. (n. s.) 37: 24, pl. 4,
figs. 1-6. 3.
bisetosa Schuster. Tex., N. Mex., Ariz.
Sphaeropthalma (Photopsis) bisetosa Schuster, 1958. Ent. Amer. (n. s.) 37: 36. ¢.
blakeii (Fox). Nev., Ariz., N. Mex., Calif.; Mexico (Baja California). Host: Diadasia vallicola
Timb.
Photopsis Blakewi Fox, 18938. Calif. Acad. Sci., Proce. (2) 4: 6. d.
Mutilla Gautschii Dalla Torre, 1897. Cat. Hym. 8: 48. N. name erroneously proposed for
blakevi Fox, thought to be preoce. by Cameron, 1894.
Mutilla ceyx Fox, 1899. Amer. Ent. Soc., Trans. 25: 262. 6.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 17 (synonymy).

Biology: Ferguson, 1962. Univ. Calif. Pubs. Ent. 27: 10-15, pls. 2, 3, 4, 5, 7, tab. 9 (life
history).

Superfamily SCOLIOIDEA 1291

capricornis (Rohwer). Colo. (Boulder).
Photopsis capricornis Rohwer, 1909. Amer. Ent. Soe., Trans. 35: 131. °.
ceres (Fox). Ariz., Colo., Utah.
Mutilla ceres Fox, 1899. Amer. Ent. Soc., Trans. 25: 257. °.
ceyxoides Schuster. Ariz.
Sphaeropthalma (Photopsis) ceyxoides Schuster, 1958. Ent. Amer. (n. s.) 37: PASS Che
clara clara (Cresson). N. Mex., Colo. to Wyo. and Idaho. This subsp. is transitional to
Sphaeropthalma subg. Physetapsis Schuster.
Mutilla clara Cresson, 1865. Ent. Soc. Phila., Proc. 4: 439. 6.

Taxonomy: Schuster, 1958. Ent. Amer. (n. s.) 37: 27.
clara cleomella Schuster. Mont. This subsp. is transitional to Sphaeropthalma subg.
Physetapsis Schuster.

Sphaeropthalma (Photopsis) clara cleomella Schuster, 1958. Ent. Amer. (n. s.) 37: 27. d.

coaequalis Cameron. Mont. and Oreg. to N. Mex. and north. Ariz., Kans., Tex.; north. Mexico.
Sphaerophthalma (!) coaequalis Cameron, 1896. Biol. Cent.-Amer., Hym., v. 2, p. 379. 6.
Mutilla albicincta Fox, 1899. Amer. Ent. Soc., Trans. 25: 255. d.
Sphaeropthalma (Photopsis) albicincta piceoceps Schuster, 1958. Ent. Amer. (n. s.) 37: 35.

é.
Sphaeropthalma (Photopsis) albicincta flavipes Schuster, 1958. Ent. Amer. (n. s.) 37: 36.
é.

Taxonomy: Mickel, 1965. Ent. Soc. Wash., Proc. 67: 1 (synonymy).
danaus (Blake). Kans., Tex., N. Mex.; Mexico.
Agama Danaus Blake, 1871. Amer. Ent. Soc., Trans. 3: 261. d.
dentifera Schuster. S. Dak., Minn.
Sphaeropthalma (Photopsis) dentifera Schuster, 1958. Ent. Amer. (n. s.) 37: 24. d.
diomeda (Fox). Tex.
Mutilla diomeda Fox, 1899. Amer. Ent. Soc., Trans. 25: 257. 9.
dirce (Fox). Ariz., Utah, Tex.
Mutilla dirce Fox, 1899. Amer. Ent. Soc., Trans. 25: 258. 2.
edwardsii edwardsii (Cresson). Oreg., Idaho.
Mutilla Edwardsii Cresson, 1875. Amer. Ent. Soc., Trans. 5: 119. 6.
edwardsii flammifera Schuster. Calif.
Sphaeropthalma (Photopsis) edwardsii flammifera Schuster, 1958. Ent. Amer. (n. s.) 37:
36. 3.
erato (Blake). Tex.
Mutilla Erato Blake, 1879. Amer. Ent. Soc., Trans. 7: 251. °.
erigone (Fox). Colo.
Mutilla erigone Fox, 1899. Amer. Ent. Soc., Trans. 25: 268. °.
facilis (Cameron). N. Mex., Ariz.; Mexico (Durango).
Sphaerophthalma (!) facilis Cameron, 1896. Biol. Cent.-Amer., Hym., v. 2, p. 392. d.
Mutilla Hubbardii Fox, 1899. Amer. Ent. Soc., Trans. 25: 256. 6.
Sphaeropthalma (Photopsis) piceogaster Schuster, 1958. Ent. Amer. (n. s.) 37: 27. d.
ferruginea (Blake). Nev., Calif. Host: Isodontia elegans (Smith).
Agama ferruginea Blake, 1879. Amer. Ent. Soc., Trans. 7: 254. d.
Mutilla ferruginosa Dalla Torre, 1897. Cat. Hym., v. 8, p. 40. N. name for Agama
ferruginea Blake, not Mutilla ferruginea Smith.

Biology: Davidson, 1899. Ent. News 10: 180 (host).

ferruginopsis Schuster. Ariz., Calif.

Sphaeropthalma (Photopsis) ferruginopsis Schuster, 1958 Ent. Amer. (n. s.) OlerZo nor
fuscipes Schuster. Ariz.; Mexico (Durango). This sp. is transitional to Sphaeropthalma subg.

Physetapsis Schuster.

Sphaeropthalma (Photopsis) fuscipes Schuster, 1958. Ent. Amer. (n. s.) 37: 21. d.
halcyone (Fox). Kans. (Hamilton Co.).

Mutilla halcyone Fox, 1899. Amer. Ent. Soc., Trans. 25: 257. °.

1292 Hymenoptera in America North of Mexico

helicaon (Fox). Nev., Ariz., Calif.; Mexico (Baja California, Coahuila).
Mutilla helicaon Fox. 1899. Amer. Ent. Soe., Trans. 25: 254. 6.
Photopsis lingulatus Viereck, 1903. Acad. Nat. Sci. Phila., Proc. 54: 737. d.
Sphaeropthalma (Photopsis) carinata Schuster, 1958. Ent. Amer. (n. s.) 37: 34. d.
Sphaeropthalma (Photopsis) helicaon coahuilae Schuster, 1958. Ent. Amer. (n. s.) 37: 34.
3.
Sphaeropthalma (Photopsis) helicaon diegueno Schuster, 1958. Ent. Amer. (n. s.) 37: 35. d.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., Biol. Ser. 8, no. 4: 17 (synonymy).
hypermnestra (Fox). Calif. (Poway).
Mutilla hypermnestra Fox, 1899. Amer. Ent. Soc., Trans. 25: 268. 2.
ignacio Schuster. Southern Calif.; Mexico (Baja California).
Sphaeropthalma (Photopsioides) amphion ignacio Schuster, 1958. Ent. Amer. (n. s.) 37: 38.
é.

imperialiformis imperialiformis (Viereck). Tex. to Kans. and Colo.
Mutilla (Photopsis) imperialiformis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 189. ¢.
imperialiformis maricopae Schuster. Ariz. to Wash.
Sphaeropthalma (Photopsis) imperialiformis maricopae Schuster, 1958. Ent. Amer. (n. s.)
St 34e de
imperialis (Blake). Tex.
Agama imperialis Blake, 1871. Amer. Ent. Soc., Trans. 3: 260. 6.
insignis (Baker). Calif. (Claremont); Mexico (Baja California).
Photopsis insignis Baker, 1905. Invertebrata Pacifica 1: 115. ¢.
jacala Schuster. Ariz.; Mexico (Jacala).
Sphaeropthalma (Photopsis) jacala Schuster, 1958. Ent. Amer. (n. s.) 387: 21. ¢.
juxta (Blake). Iowa, Kans., Tex., N. Mex., Ariz.
Agama juxta Blake, 1872. Amer. Ent. Soc., Trans. 4: 76. 3.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci. Bul., ser. 8, no. 4: 15.
laodamia (Fox). Ariz.
Mutilla laodamia Fox, 1899. Amer. Ent. Soc., Trans. 25: 258. °.
luiseno Schuster. Calif.
Sphaeropthalma (Photopsis) luiseno Schuster, 1958. Ent. Amer. (n. s.) 37: 23. 3.
marpesia (Blake). Kans., Utah.
Mutilla Marpesia Blake, 1879. Amer. Ent. Soc., Trans. 7: 247. °.
Sphaerophthalma (!) luteola Blake, 1886. Amer. Ent. Soc., Trans. 13: 235. °.
megagnathos aurifera Schuster. Calif., Ariz.
Sphaeropthalma (Photopsis) megagnathos auriferus Schuster, 1958. Ent. Amer. (n. s.) 37:
36. 3.
megagnathos megagnathos Schuster. Calif., Ariz.
Sphaeropthalma (Photopsis) megagnathos megagnathos Schuster, 1958. Ent. Amer. (n. s.)
37: 36. 3.
militaris Schuster. Ariz., Calif.
Sphaeropthalma (Photopsis) militaris Schuster, 1958. Ent. Amer. (n. s.) 37: 27, pl. 5, fig. 2.
3

minutella (Mickel). Calif. (Lassen Co.).
Photopsis minutella Mickel, 1938. Pan-Pacific Ent. 14: 183. @.

nanula (Dalla Torre). Tex., Colo., Nev., Idaho. Host: Tachysphex sp.
Mutilla pygmea Blake, 1879. Amer. Ent. Soc., Trans. 7: 250. 2. Preoce.
Mutilla nanula Dalla Torre, 1897. Cat. Hym., v. 8, p. 65. N. name.
Mutilla pygmaea (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 65.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 102 (host).
neomexicana Schuster. N. Mex.
Sphaeropthalma (Photopsis) neomexicana Schuster, 1958. Ent. Amer. (n. s.) 37: 30. 6.
noctivaga (Melander). Nebr. and Wyo. south to Tex. and Ariz.
Mutilla noctivaga Melander, 1903. Amer. Ent. Soc., Trans. 29: 318. 3.

Superfamily SCOLIOIDEA 1293

Sphaeropthalma (Photopsis) noctivaga infuscata Schuster, 1958. Ent. Amer. (n. s.) 37: 32.
3

nokomis nokomis (Blake). Ariz.
Agama Nokomis Blake, 1871. Amer. Ent. Soc., Trans. 3: 260. 3.
nokomis ruficeps Schuster. Ariz.
Sphaeropthalma (Photopsis) nokomis ruficeps Schuster, 1958. Ent. Amer. (n. s.) 37: 35. d.
ordae Schuster. Tex.
Sphaeropthalma (Photopsis) ordae Schuster, 1958. Ent. Amer. (n. s.) 37: 23. d.
orestes (Fox). Wash. to Calif., Nev.; Mexico (Baja California). Host: Anthidium c. collectum
Huard; Euodynerus a auranus (Cam.); Tachysphex tenuipunctus Fox.
Mutilla orestes Fox, 1899. Amer. Ent. Soc., Trans. 25: 256. ¢.
Mutilla Pattersonae Melander, 1903. Amer. Ent. Soc., Trans. 29: 309. ¢.
Photopsis indigens Baker, 1905. Invertebrata Pacifica 1: 112. ¢.
Photopsis uniformis Baker, 1905. Invertebrata Pacifica 1: 113. d.
Photopsis pedatus Baker, 1905. Invertebrata Pacifica 1: 115. d.
Photopsis ingenuus Baker, 1905. Invertebrata Pacifica 1: 116. ¢.
Photopsis salmani Mickel, 1938. Pan-Pacifie Ent. 14: 178. 2, d.
Sphaeropthalma (Photopsis) salmani fresnoensis Schuster, 1958. Ent. Amer. (n. s.) 37: 30.
3

Sphaeropthalma (Photopsis) salmani oregana Schuster, 1958. Ent. Amer. (n. s.) 37: 31. 6.

Taxonomy: Schuster, 1958. Ent. Amer. (n. s.) 37: 30 (synonymy). —Ferguson, 1962. Univ.
Calif. Pubs. Ent. 27: 10 (synonymy).

Biology: Ferguson, 1962. Univ. Calif. Pubs. Ent. 27: 6-10, pls. 1, 6, tabs. 1, 9 (life history).

pallidipes Schuster. Ariz.
Sphaeropthalma (Photopsis) pallidipes pallidipes Schuster, 1958. Ent. Amer. (n. s.) 37: 27.
3

Sphaeropthalma (Photopsis) pallidipes gila Schuster, 1958. Ent. Amer. (n. s.) 37: 28. d.

parkeri Schuster. Ariz., Calif.
Sphaeropthalma (Photopsis) parkeri Schuster, 1958. Ent. Amer. (n. s.) 37: 28. d.

pervaga (Melander). Tex. (Fedor).
Mutilla pervaga Melander, 1903. Amer. Ent. Soc., Trans. 29: 321. °.
pinalea Schuster. N. Dak. to Tex., Ariz.; Mexico (Coahuila, Durango).
Sphaeropthalma (Photopsis) pinalea pinalea Schuster, 1958. Ent. Amer. (n. s.) 37: 32. d.
Sphaeropthalma (Photopsis) pinalea texanella Schuster, 1957. Ent. Amer. (n. s.) 37: 32. 6.
pinales Schuster. Ariz.
Sphaeropthalma (Photopsis) pinales Schuster. 1958. Ent. Amer. (n. s.) 37: 36. d.
pluto (Fox). Tex., N. Mex., Ariz.
Mutilla pluto Fox, 1899. Amer. Ent. Soc., Trans. 25: 255. d.
Mutilla palamedes Fox, 1899. Amer. Ent. Soc., Trans. 25: 264. ¢.
rubriventris Schuster. Tex., N. Mex., Ariz.; Mexico (Tamaulipas).
Sphaeropthalma (Photopsis) facilis rubriventris Schuster, 1958. Ent. Amer. (n. s.) 37: 26.
3

sanctaefeae (Cockerell). N. Mex. (Santa Fe).

Mutilla sanctae-feae Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila., Proc. 49:

1372

scudderi Schuster. Ariz.

Sphaeropthalma (Photopsis) scudderi Schuster, 1958. Ent. Amer. (n. s.) 37: 28. d.
seminanula Rohwer. Colo. (Boulder).

Photopsis seminanula Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 180. °.
similis Schuster. Calif.

Sphaeropthalma (Photopsis) similis Schuster, 1958. Ent. Amer. (n. s.) 37: 30. d.
spinifera Schuster. Kans., Okla., Colo.

Sphaeropthalma (Photopsis) spinifera Schuster, 1958. Ent. Amer. (n. s.) 37: 23. d.
tetracuspis Schuster. Calif. (Riverside Co.); Mexico (Baja California).

Sphaeropthalma (Photopsis) tetracuspis Schuster, 1958. Ent. Amer. (n. s.) 37: 31. d.

1294 Hymenoptera in America North of Mexico

triangularis (Blake). Tex., N. Mex., Ariz., Nev., Calif; Mexico (Baja California).
Agama triangularis Blake, 1871. Amer. Ent. Soc., Trans. 3: 262. 6.
tuberculifera Schuster. Calif. (?).
Sphaeropthalma (Photopsis) tuberculifera Schuster, 1958. Ent. Amer. (n. s.) 37: 25. 6.
unicolor (Cresson). U. S. west of Rocky Mts.; Mexico (Baja California del Norte). Host:
Isodontia elegans (Sm.); Anthidium c. collectum Huard, Callanthidium illustre (Cr.),
Ashmeadiella c. californica (Ashm.); Anthophora abruptella Ckll., A. bomboides
neomexicana Ckll., A. linsleyi Timb., A. montana Cr.?, A. occidentalis Cr., Diadasia
bituberculata (Cr.), Melissodes robustior Ckll., Xeromelecta californica (Ashm.).
Mutilla unicolor Cresson, 1865. Ent. Soc. Phila., Proc. 4: 389. ¢.
Agama mendica Blake, 1871. Amer. Ent. Soc., Trans. 3: 259. 6.
Mutilla auraria Blake, 1879. Amer. Ent. Soc., Trans. 7: 248. °.
Mutilla Aspasia Blake, 1879. Amer. Ent. Soc., Trans. 7: 250. 9.
Mutilla Phaedra Blake, 1879. Amer. Ent. Soc., Trans. 7: 251. 2.
Agama rustica Blake, 1879. Amer. Ent. Soc., Trans. 7: 252. 3.
Photopsis nebulosus Blake, 1886. Amer. Ent. Soc., Trans. 13: 275. d.
Sphaerophthalmia (!) anthophora Ashmead, 1897. In Davidson, South. Calif. Acad. Sci.,
jena 19 by Oe
Mutilla monochroa Dalla Torre, 1897. Cat. Hym., v. 8, p. 63. N. name for Mutilla unicolor
Cresson, not Myrmosa unicolor Say.
Dasymutilla sumneriella Cockerell, 1915. Entomologist 48: 249. ?.
Sphaeropthalma (Photopsis) rustica ocellaria Schuster, 1958. Ent. Amer. (n. s.) 37: 32. 3.

Biology: Davidson, 1894. Ent. News 5: 170 (host records). —Davidson, 1897. South. Calif.
Acad. Sci., Proc. 1: 4-5 (host records). —Davidson, 1899. Ent. News 10: 180 (host record).
—Linsley and MacSwain, 1942. Amer. Midland Nat. 27: 409 (host records). —Ferguson,
1962. Univ. Calif. Pubs. Ent. 27: 15-22, pls. 3, 5, 7, tab. 9 (life history, host records).
virguncula (Blake). N. Mex.

Sphaerophthalma (!) virguncula Blake, 1886. Amer. Ent. Soc., Trans. 13: 253. 9.
zenobia (Blake). Calif., ? Tex.

Mutilla zenobia Blake, 1879. Amer. Ent. Soc., Trans. 7: 250. °.
zephyritis (Fox). Calif. (Los Angeles).

Mutilla zephyritis Fox, 1899. Amer. Ent. Soc., Trans. 25: 269. °.

NOMEN NUDUM IN SPHAEROPTHALMA SUBGENUS PHOTOPSIS BLAKE

Sphaeropthalma (Photopsis) nixonensis Parker and Bohart, 1966. Pan-Pacific Ent. 42: 98. Nev.
Host: Proteriades sp.

Genus SPHAEROPTHALMA Subgenus PHOTOPSIOIDES Schuster

Sphaeropthalma subg. Photopsioides Schuster, 1958. Ent. Amer. (n. s.) 37: 10 (in key), 36.
Type-species: Agama wro Blake. Orig. desig.

In addition to the host records cited below under wro (Blake), Ferguson, 1962 (Univ. Calif.
Pubs. Ent. 27: tab. 10) records Sceliphron caementarium (Dru.) as the host of an unidentified
species of this subgenus.

abdominalis (Blake). Kans., Tex., Colo., Ariz., Utah. Host: Trypargilum t. tridentatum (Pack.).
Photopsis abdominalis Blake, 1886. Amer. Ent. Soc., Trans. 13: 275. 6.
Mutilla coloradensis Dalla Torre, 1897. Cat. Hym., v. 8, p. 25. N. name for Photopsis
abdominalis Blake, not Mutilla abdominalis Westwood.
amphion (Fox). Ariz., Nev., Calif., Oreg.; Mexico (Baja California). Host: Sapyga aculeata Cr.;
Leptochilus chiricahua Prkr., Ancistrocerus c. catskill (Sauss.); Trypargilum t.
tridentatum (Pack.); Hoplitis bullifacies Mich., H. f. fulgida (Cr.), H. g. grinnelli Ckll.,
H. hypostomalis Mich., H. sambuci Titus, Anthocopa copelandica (Ckll.), Ashmeadiella
m. meliloti (Ckll.), Stelis spp.
Mutilla amphion Fox, 1899. Amer. Ent. Soc., Trans. 25: 263. ¢.
Photopsis abstrusa Baker, 1905. Invertebrata Pacifica 1: 113. 6.
Photopsis nudata Baker, 1905. Invertebrata Pacifica 1: 114. 6.

Superfamily SCOLIOIDEA 1295

Taxonomy: Ferguson, 1967. Brigham Young Univ., Sci. Bul., ser. 8, no. 4: 20 (synonymy).

Biology: Parker, 1975. Pan-Pacific Ent. 51: 116 (host).
contracta (Blake). Nev., Oreg.
Agama contracta Blake, 1879. Amer. Ent. Soc., Trans. 7: 253. d.
Mutilla contrahenda Dalla Torre, 1897. Cat. Hym., v. 7, p. 27. N. name for Agama
contracta Blake, not Mutilla contracta Say.
uro (Blake). Kans., Tex., N. Mex., Ariz. Host: Pachodynerus astraeus (Cam.), eumenid sp.;
Trypargilum t. tridentatum (Pack.); Dianthidium curvatum sayi CkIl.
Agama uro Blake, 1879. Amer. Ent. Soce., Trans. 7: 253. ¢.
Photopsis melanderi Baker, 1905. Invertebrata Pacifica 1: 112. d.
Sphaeropthalma (Photopsioides) uro stenognatha Schuster, 1958. Ent. Amer. (n. s.) 37: 38.
é.

Biology: Fischer, 1951. Kans. Ent. Soc., Jour. 24: 49 (host record). —Ferguson, 1962. Univ.
Calif. Pubs. Ent. 27: tab. 10 (host records). —Parker and Bohart, 1966. Pan-Pacific Ent. 42:
93 (host record). —Krombein, 1967. Trap-nesting wasps and bees, pp. 478-479 (life history).

Genus SPHAEROPTHALMA Subgenus SPHAEROPTHALMA Blake

Sphaeropthalma Blake, 1871. Amer. Ent. Soc., Trans. 3: 2382.
Type-species: Mutilla (Sphaeropthalma) scaeva Blake. Automatic desig. by
Ashmead, 1899 (=Sphaeropthalma pensylvanica scaeva (Blake)).
Sphaerophthalma (!) Blake, 1886. Amer. Ent. Soc., Trans. 13: 179. Emend.
Type-species: Mutilla (Sphaeropthalma) scaeva Blake. Desig. by Ashmead, 1899
(=Sphaeropthalma pensylvanica scaeva (Blake)).

Revision: Schuster, 1944. Brooklyn Ent. Soc., Bul. 39: 141-147.
auripilis auripilis (Blake). Kans., Okla., Tex. Host: Chalybion californicum (Sauss.),
Sceliphron caementarium (Drury), Trypoxylon sp.; solitary eumenid in old clay cell of
Sceliphron.
Mutilla (Sphaeropthalma) auripilis Blake, 1871. Amer. Ent. Soc., Trans. 3: 233. 6.
Sphaerophthalma (!) albiplumosa Schuster, 1944. Brooklyn Ent. Soe., Bul. 39: 145. 2.

Biology: Rau, 1940. Ent. Soc. Amer., Ann. 33: 594 (host).

auripilis fasciventris Schuster. Tex. (Edinburg).
Sphaeropthalma (Sphaeropthalma) auripilis fasciventris Schuster, 1958. Ent. Amer. (n. s.)
37: 39. 3.

Taxonomy: Mickel, 1974. Ent. Soc. Amer., Ann. 67: 466. °.
boweri Schuster. Tex. (College Station).
Sphaerophthalma (!) boweri Schuster, 1944. Brooklyn Ent. Soc., Bul. 39: 143. 6.
pensylvanica floridensis Schuster. South. Fla.
Sphaerophthalma (!) pennsylvanica(!) floridensis Schuster, 1944. Brooklyn Ent. Soc., Bul.
39: 142. 9.
Taxonomy: Mickel, 1974. Ent. Soc. Amer., Ann. 67: 466. d.
pensylvanica pensylvanica (Lepeletier). N. C., Ga., cent. Fla., La., Mo., Kans., Tex. Host:
Sceliphron caementarium (Drury), Trypargilum politum (Say), T. clavatum (Say);
anthidiine sp.
Mutilla pensylvanica Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 628. d.
Mutilla (Sphaeropthalma) pennsylvanica (!) Blake, 1871. Amer. Ent. Soc., Trans. 3: 233.

3.
Mutilla (Sphaeropthalma) balteola Blake, 1871. Amer. Ent. Soc., Trans. 3: 248. 2.

Taxonomy: Bradley, 1916. Amer. Ent. Soc., Trans. 42: 331.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 7 (host). —Rau, 1928. Acad. Sci. St. Louis,
Trans. 25: 437 (host).
pensylvanica scaeva (Blake). Mass. to N. C., Ohio, Ill., Mo., Kans., Tex. Host: Auplopus a.
architectus (Say), A. m. mellipes (Say); Trypargilum clavatum (Say), T. collinum

1296 Hymenoptera in America North of Mexico

rubrocinctum (Pack.), T. politwm (Say), T. lactitarse (Sauss.), Sceliphron caementarium
(Dru.). Parasite: Melittobia chalybu Ashm.
Mutilla (Sphaeropthalma) scaeva Blake, 1871. Amer. Ent. Soe., Trans. 3: 232. 6.

Taxonomy: Bradley, 1916. Amer. Ent. Soc., Trans. 42: 331.

Biology: Rau and Rau, 1918. Wasp studies afield, pp. 88-89 (host). —Rau, 1922. Acad. Sci. St.
Louis, Trans. 24, no. 7: 8 (host). —Krombein, 1951. U. S. Dept. Agr., Monog. 2: 752 (hosts).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 476-478, figs. 135-139 (life history).

Genus PHOTOMORPHUS Viereck

Taxonomy: Schuster, 1944. Brooklyn Ent. Soc., Bul. 39: 147-151 (east spp. only). —Krombein,
1954. Amer. Ent. Soc., Trans. 80: 6-7 (key to eastern females).

Genus PHOTOMORPHUS Subgenus PHOTOMORPHINA Schuster

Photomorphus subg. Photomorphina Schuster, 1952. Brooklyn Ent. Soc., Bul. 47: 53.
Type-species: Photomorphus (Photomorphina) aurifera Schuster. Orig. desig.
Photomorphus subg. Photomorphina Schuster, 1958. Ent. Amer. (n.s.) 37: 8 (in key), 39.
Preoce. by Photomorphus subg. Photomorphina Schuster, 1952, and a syn. of it.
Type-species: Photomorphus (Photomorphina) quadriangulata Schuster. Orig.
desig.
aurifer Schuster. Ariz.
Photomorphus (Photomorphina) aurifera Schuster, 1952. Brooklyn Ent. Soce., Bul. 47: 53.
é.
Photomorphus (Photomorphina) aurifera Schuster, 1958. Ent. Amer. (n. s.) 37: 44. 6.
Preoce. by and a syn. of Photomorphus (Photomorphina) aurifera Schuster, 1952.
auriventris Schuster. Tex.
Photomorphus (Photomorphina) auriventris Schuster, 1958. Ent. Amer. (n. s.) 37: 40. ¢.
bequaertii (Schuster), n. comb. (W. E. Ferguson). Tex.
Odontophotopsis (Odontophotopsis) bequaertii Schuster, 1958. Ent. Amer. (n. s.) 37: 52. d.
californicus Schuster. Calif.
Photomorphus (Photomorphina) californica Schuster, 1958. Ent. Amer. (n. s.) 37: 44. 6.
clandestinus (Viereck). Tex., N. Mex.
Odontophotopsis clandestinus Viereck, 1903. Acad. Nat. Sci. Phila., Proc. 54: 740. 3.
cobabi Schuster. Ariz.; Mexico (Sonora, Baja California).
Photomorphus (Photomorphina) cobabi Schuster, 1958. Ent. Amer. (n. s.) 37: 41. d.
cochisae Schuster. Ariz.; Mexico.
Photomorphus (Photomorphina) cochisae Schuster, 1958. Ent. Amer. (n. s.) 37: 48. 6.
coloradellus Schuster. Colo.
Photomorphus (Photomorphina) coloradella Schuster, 1958. Ent. Amer. (n. s.) 37: 44. d.
comobabi Schuster. Ariz.; Mexico.
Photomorphus (Photomorphina) comobabi Schuster, 1958. Ent. Amer. (n. s.) 37: 45. 6.
crepusculus (Viereck). Kans. (Morton Co.).
Mutilla (Photomorphus?) crepuscula Viereck, 1906. Amer. Ent. Soc., Trans. 32: 188. ¢.
dichrous Schuster. Ariz.
Photomorphus (Photomorphina) dichrous Schuster, 1958. Ent. Amer. (n. s.) 37: 41. 6.
hebes (Melander). Tex., N. Mex., Ariz.; Mexico (Sonora, Coahuila, Tamaulipas).
Mutilla hebes Melander, 1903. Amer. Ent. Soe., Trans. 29: 311. ¢.
Photomorphus (Photomorphina) taeniatus Schuster, 1958. Ent. Amer. (n. s.) 37: 42. 6.
Photomorphus (Photomorphina) taeniatus parksi Schuster, 1958. Ent. Amer. (n. s.) 37: 42.
3.
imperialoides Schuster. Ariz.
Photomorphus (Photomorphina) imperialoides Schuster, 1958. Ent. Amer. (n. s.) 37: 40. 6.
jason (Fox). Tex., Okla.
Mutilla jason Fox, 1899. Amer. Ent. Soc., Trans. 25: 250. ¢.

Superfamily SCOLIOIDEA 1297

juanita Schuster. Tex.; Mexico (Tamaulipas).

Photomorphus (Photomorphina) juanita Schuster, 1952. Brooklyn Ent. Soc., Bul. 47: 57. d.

Photomorphus (Photomorphina) juanita Schuster, 1958. Ent. Amer. (n. s.) 37: 43. 6.

Preoce. by and a syn. of Photomorphus (Photomorphina) juanita Schuster, 1952.

minimus Schuster. Ariz. (Wellton).

Photomorphus (Photomorphina) minima Schuster, 1958. Ent. Amer. (n. s.) STASMS
myrmicoides (Cockerell). Ala., Tex., Ill., lowa, Kans., S. Dak.

Mutilla parvula Blake, 1886. Amer. Ent. Soc., Trans. 13: 206. 2. Preoce.

Sphaerophthalma (!) myrmicoides Cockerell, 1895. Ent. News 6: 62. °.

Mutilla impar Melander, 1903. Amer. Ent. Soc., Trans. 29: 321. °.

Odontophotopsis subtenuis Viereck, 1904. Amer. Ent. Soc., Trans. 30: 85. d.

Taxonomy: Bradley, 1916. Amer. Ent. Soc., Trans. 42: 336. ¢. —Mickel, 1934. Ent. Soc.
Amer., Ann. 27: 611. 2, ¢. —Mickel, 1965. Ent. Soc. Wash., Proc. 67: 3-4 (synonymy).
nanullus Schuster. Ariz.
Photomorphus (Photomorphina) nanulla Schuster, 1958. Ent. Amer. (n. s.) 37: 44. 6.
obscurus Schuster. Calif., Ariz.
Photomorphus (Photomorphina) obscura Schuster, 1958. Ent. Amer. (n. s.) 37: 45. d.
quadriangulatus Schuster. Ariz.; Mexico (Sonora).
Photomorphus (Photomorphina) quadriangulata Schuster, 1958. Ent. Amer. (n. s.) 37: 42.
3.
sarpedon (Fox). Tex.; Mexico (Coahuila).
Mutilla sarpedon Fox, 1899. Amer. Ent. Soc., Trans. 25: 267. 6.
spinci (Bradley). N. C. to Fla., Ala.
Photopsis (Odontophotopsis) spinci Bradley, 1916. Amer. Ent. Soc., Trans. 42: 335. d.
Photopsis (Odontophotopsis) spinci floridensis Schuster, 1944. Brooklyn Ent. Soe., Bul. 39:
155. o.
spinciformis Schuster. Tex.
Photomorphus (Photomorphina) spinciformis Schuster, 1958. Ent. Amer. (n. s.) 37: 44. d.
thamyras (Fox). Tex.
Mutilla thamyras Fox, 1899. Amer. Ent. Soc., Trans. 25: 267. d.
trunculus (Viereck). Kans., Okla., Tex., Ariz.
Mutilla simpliciventris Melander, 1903 (Oct.). Amer. Ent. Soc., Trans. 29: 315. d. Preoce.
by Andre, July, 1903.
Odontophotopsis trunculus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 85. d.
Odontophotopsis melanderella Krombein, 1951. U. S. Dept. Agr., Monog. 2: 757. N. name.
wheeleri (Melander). Tex. (Austin).
Mutilla Wheeleri Melander, 1903. Amer. Ent. Soc., Trans. 29: 316. d.
vegas Schuster. N. Mex., Ariz., Utah.
Photomorphus (Photomorphina) vegas Schuster, 1958. Ent. Amer. (n. s.) 37: 42. 3d.

Genus PHOTOMORPHUS Subgenus PHOTOMORPHUS Viereck

Photomorphus Viereck, 1903. Ent. News 14: 249.
Type-species: Photomorphus Johnsoni Viereck. Orig. desig.

alogus Viereck. Ga., Fla., Miss.
Photomorphus alogus Viereck, 1903. Ent. News 14: 251. 6.
Taxonomy: Krombein, 1954. Amer. Ent. Soc., Trans. 80: 3. °.
banksi (Bradley). N. J. south to Fla., Ala.
Sphaerophthalma (!) (Photomorphus) banksi Bradley, 1916. Amer. Ent. Soe., Trans. 42:
Boleor
bradleyi (Schuster). S. C. (Clemson).
Photopsis bradleyi Schuster, 1944. Brooklyn Ent. Soc., Bul. 39: 152. °.
johnsoni johnsoni Viereck. N. Y. to Ga., Okla.
Photomorphus Johnsoni Viereck, 1903. Ent. News 14: 249. d.

1298 | Hymenoptera in America North of Mexico

Photomorphus johnsoni var. argentipilis Schuster, 1944. Brooklyn Ent. Soc., Bul. 39: 148.

johnsoni natchitoches Schuster. La.
Photomorphus (Photomorphus) johnsoni natchitoches Schuster, 1958. Ent. Amer. (n. s.) 37:
46. d.
paulus (Bradley). Ga., Fla.
Photopsis (Odontophotopsis) paula Bradley, 1916. Amer. Ent. Soc., Trans. 42: 338. 6.

Taxonomy: Schuster, 1944. Brooklyn Ent. Soe., Bul. 39: 154. ¢.
quintilis (Viereck). Iowa, Kans., La.
Mutilla (Photomorphus) quintilis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 187. ¢.
rubroscutellatus (Bradley). Va. (Falls Church).
Sphaerophthalma (!) (Photomorphus) rubroscutellatus Bradley, 1916. Amer. Ent. Soc.,
Mransy423ea2. 16 -

Genus ODONTOPHOTOPSIS Viereck
Revision: Viereck, 1904. Amer. Ent. Soc., Trans. 30: 81-92.
Genus ODONTOPHOTOPSIS Subgenus ODONTOPHOTOPSIS Viereck

Odontophotopsis Viereck, 1903. Acad. Nat. Sci. Phila., Proc. 54: 738.
Type-species: Odontophotopsis exogyrus Viereck. Orig. desig.
Tetraphotopsis Ashmead, 1903. Canad. Ent. 35: 305.
Type-species: Tetraphotopsis Hubbardi Ashmead. Orig. desig. (Not Mutilla
Hubbardii Fox as cited by Mickel, 1928. U. S. Natl. Mus., Bul. 143: 37).
acmaea Viereck. Ariz., south. Calif.; Mexico (Sonora).
Odontophotopsis acmaeus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 84. d.
adonis (Fox). N. Mex. (Las Cruces).
Mutilla adonis Fox, 1899. Amer. Ent. Soc., Trans. 25: 265. 6.
alemon (Fox). Tex., N. Mex.
Mutilla alemon Fox, 1899. Amer. Ent. Soc., Trans. 25: 266. 3.
Mutilla trita Melander, 1903. Amer. Ent. Soc., Trans. 29: 317. ¢.
Odontophotopsis crucis Viereck, 1904. Amer. Ent. Soc., Trans. 30: 86. d.
annulata Baker. Calif. (Claremont).
Odontophotopsis annulatus Baker, 1905. Invertebrata Pacifica 1: 94. 3.
anomala Schuster. Tex.
Odontophotopsis (Odontophotopsis) anomala Schuster, 1958. Ent. Amer. (n. s.) 37: 60. ¢.
appacheorum appacheorum Schuster. Tex.
Odontophotopsis (Odontophotopsis) appacheorum appacheorum Schuster, 1958. Ent.
Amer. (n. s.) 37: 55. do.
appacheorum rostrata Schuster. Ariz.
Odontophotopsis (Odontophotopsis) appacheorum rostratus Schuster, 1958. Ent. Amer. (n.
s.) 87: 55. 3.
argentipilis (Provancher). Published type locality Fla. is erroneous.
Sphaerophthalma (!) argentipilis Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym.,
py2oleice
armata Schuster. Nev.
Odontophotopsis (Odontophotopsis) armata Schuster, 1958. Ent. Amer. (n. s.) 37: 60. d.
augusta Viereck. N. Mex. (St. Augustine).
Odontophotopsis augustus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 90. d.
bicolor (Blake). Tex., Ariz.
Agama bicolor Blake, 1879. Amer. Ent. Soc., Trans. 7: 252. 6.
Mutilla Madejskii Dalla Torre, 1897. Cat. Hym., v. 8, p. 56. N. name for Agama bicolor
Blake, not Mutilla bicolor Pallas.

Taxonomy: Fox, 1899. Amer. Ent. Soc., Trans. 25: 259-260. 3d (in key to group imperialis).

Superfamily SCOLIOIDEA 1299

biramosa Schuster. Calif. (Holtville).
Odontophotopsis (Odontophotopsis) biramosa Schuster, 1952. Brooklyn Ent. Soc., Bul. 47:
48, fig. 1. d.
braccata Schuster. Tex., N. Mex.
Odontophotopsis (Odontophotopsis) braccatus Schuster, 1958. Ent. Amer. (n. s.) 37: 54. d.
clypeata Schuster. Ariz., Nev., Calif.
Odontophotopsis (Odontophotopsis) clypeatus Schuster, 1958. Ent. Amer. (n. s.) 37: 59. d.
cockerelli (Melander). Tex. (Cuero).
Mutilla Cockerelli Melander, 1903. Amer. Ent. Soc., Trans. 29: 307. ¢.
conifera Schuster. Ariz.
Odontophotopsis (Odontophotopsis) coniferus Schuster, 1958. Ent. Amer. (n. s.) 37: 48. d.
cookii Baker. Calif., Nev.
Odontophotopsis cookii Baker, 1905. Invertebrata Pacifica 1: 93. ¢.
delodonta Viereck. Ariz.
Odontophotopsis delodontus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 91. d.
edentata distans Schuster. Tex. (Devils River).
Odontophotopsis (Odontophotopsis) edentata distans Schuster, 1958. Ent. Amer. (n. s.) 37:
53. d.
edentata edentata Schuster. Ariz.
Odontophotopsis (Odontophotopsis) edentata edentata Schuster, 1958. Ent. Amer. (n. s.) 37:
53. d.
erebus (Melander). Kans., Okla., Tex., N. Mex., Ariz., Utah; Mexico (Tamaulipas).
Mutilla Erebus Melander, 1903. Amer. Ent. Soc., Trans. 29: 312. 6.
Odontophotopsis avellanus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 88. 6.
eubule (Cameron). N. Mex.; Mexico (Sonora). Host: Probably Evaniella neomexicana (Ashm.)
in ootheca of Arenivaga genitalis Caud., or. less likely, the ootheca itself.
Sphaerophthalma (!) ewbule Cameron, 1896. Biol. Cent.-Amer., Hym., v. 2, p. 383. d.
Mutilla hamata Melander, 1903. Amer. Ent. Soc., Trans. 29: 314. ¢.

Taxonomy: Mickel, 1965. Ent. Soc. Wash., Proc. 67: 1 (synonymy).

Biology: Mickel, 1974. Ent. Soc. Amer., Ann. 67: 468 (host).

exogyra Viereck. Calif. (La Jolla).

Odontophotopsis exogyrus Viereck., 1903. Acad. Nat. Sci. Phila., Proc. 54: 738. d.
fallax Viereck. Nev.

Odontophotopsis fallax Viereck, 1904. Amer. Ent. Soc., Trans. 30: 89. d.

Odontophotopsis ocellatus Baker, 1905. Invertebrata Pacifica 1: 98. ¢.
grandiceps Schuster. Ariz. (Globe).

Odontophotopsis (Odontophotopsis) grandiceps Schuster, 1958. Ent. Amer. (n. s.) 37: 53. ¢.
grata (Melander). N. Mex. (La Cueva).

Mutilla grata Melander, 1903. Amer. Ent. Soc., Trans. 29: 308. 6.
hexadonta Schuster. Calif.

Odontophotopsis (Odontophotopsis) hecadontus Schuster, 1958. Ent. Amer. (n. s.) 37: 51. 6.
inconspicua brunnea Schuster. Ariz.

Odontophotopsis (Odontophotopsis) inconspicua brunnea Schuster, 1958. Ent. Amer. (n. s.)

St b4 aon

inconspicua inconspicua (Blake). Calif., Nev.

Photopsis inconspicuus Blake, 1886. Amer. Ent. Soc., Trans. 13: 272. ¢.

Mutilla infelix Dalla Torre, 1897. Cat. Hym., v. 8, p. 50. N. name for Photopsis

inconspicuus Blake, not Mutilla inconspicua Smith.

lamellifera Schuster. Ariz.

Odontophotopsis (Odontophotopsis) lamellifera Schuster, 1958. Ent. Amer. (n. s.) 37: 56. 6.
melicausa melicausa (Blake). B. C., Mont., Calif., Ariz., Tex.

Agama melicausa Blake, 1871. Amer. Ent. Soc., Trans. 3: 261. d.

Mutilla brevicornis Fox, 1899. Amer. Ent. Soc., Trans. 25: 255. d.

Odontophotopsis mellicornis Baker, 1905. Invertebrata Pacifica 1: 96. d.

1300 Hymenoptera in America North of Mexico

Taxonomy: Schuster, 1958. Ent. Amer. (n. s.) 37: 58 (synonymy).
melicausa piceipes Schuster. Mont.
Odontophotopsis (Odontophotopsis) mellicausa (!) piceipes Schuster, 1958. Ent. Amer. (n.
s.) 37: 58. 6.
melicausa westcottii (Melander). Tex., N. Mex.
Mutilla Westcottii Melander, 1903. Amer. Ent. Soc., Trans. 29: 310. 6.
Odontophotopsis indotatus Viereck, 1904. Amer. Ent. Soe., Trans. 30: 89. 3.
microdonta Ferguson. Nev., Calif.
Odontophotopsis (Odontophotopsis) microdonta Ferguson, 1967. Brigham Young Univ. Sci.,
Bul., Ser. 8, no. 4: 22. 6.
obliqua Viereck. B. C., Mont., Idaho, Wash., Nev., Calif.; Mexico (Baja California).
Odontophotopsis obliquus Viereck, 1925. Canad. Ent. 56: 112. 3.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci., Bul., Ser. 8, no. 4: 23 (redescription).

parksiana Schuster. Tex.

Odontophotopsis (Odontophotopsis) parksiana Schuster, 1958. Ent. Amer. (n. s.) 37: 57. 6.
parva Schuster. Ariz. (Arlington).

Odontophotopsis (Odontophotopsis) parva Schuster, 1958. Ent. Amer (n. s.) 37: 55. 6.
pudica (Melander). Wash., Calif.

Mutilla pudica Melander, 1903. Amer. Ent. Soc., Trans. 29: 309. 3.

Odontophotopsis atripes Mickel, 1938. Pan-Pacifie Ent. 14: 182. 3.
quadridentata Schuster. Calif.

Odontophotopsis (Odontophotopsis) quadridentata Schuster, 1958. Ent. Amer. (n. s.) 37: 51.

Or

quadrispinosa Schuster. Nev., Calif.; Mexico (Baja California).
Odontophotopsis (Odontophotopsis) quadrispinosa Schuster, 1958. Ent. Amer. (n. s.) 37: 51.
3.

Taxonomy: Ferguson, 1967. Brigham Young Univ. Sci., Bul., Ser. 8, no. 4: 24 (redescription).

serca Viereck. Nev., Calif.; Mexico (Baja California). Host: Anthophora linsleyi Timberlake.
Odontophotopsis sercus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 87. 3.

Biology: Linsley and MacSwain, 1942. Amer. Midland Nat. 27: 409 (host).

setifera Schuster. South. Calif., Nev., Ariz.
Odontophotopsis (Odontophotopsis) setifera Schuster, 1952. Brooklyn Ent. Soc., Bul. 47: 47.
3.

succinea Viereck. Calif.

Odontophotopsis succineus Viereck, 1903. Acad. Nat. Sci. Phila., Proc. 54: 741. d.
tapajos aulus (Blake). Tex.

Agama Aulus Blake, 1872. Amer. Ent. Soc., Trans. 4: 75. 6.
tapajos tapajos (Blake). Tex., Ariz.

Agama Tapajos Blake, 1871. Amer. Ent. Soc., Trans. 3: 262. 6.

Agama Astyanax Blake, 1879. Amer. Ent. Soc., Trans. 7: 254. 6.
tenuiptera Schuster. Ariz.; Mexico.

Odontophotopsis (Odontophotopsis) tenuiptera Schuster, 1958. Ent. Amer. (n. s.) 37: 47. 3.
territa obscura Schuster. Ariz.

Odontophotopsis (Odontophotopsis) territa obscura Schuster, 1958. Ent. Amer. (n. s.) 37:

DOME

territa territa (Cockerell). N. Mex. (Las Cruces).

Photopsis territus Cockerell, 1894. Ent. News 5: 200. 6.

unicornis Schuster. Ariz., Calif.
Odontophotopsis (Odontophotopsis) wnicornis Schuster, 1958. Ent. Amer. (n. s.) 37: 52. 3.
venusta (Blake). Ariz.
Photopsis venustus Blake, 1886. Amer. Ent. Soc., Trans. 13: 270. ¢.
Mutilla pretiosissima Dalla Torre, 1897. Cat. Hym., v. 8, p. 74. N. name for Photopsis
venustus Blake, not Mutilla venusta Smith.
Tetraphotopsis Hubbardi Ashmead, 1903. Canad. Ent. 35: 305. 3.

Superfamily SCOLIOIDEA 1301

Taxonomy: Schuster, 1958. Ent. Amer. (n. s.) 387: 54 (synonymy).

viereckii Baker. Nev. (Ormsby Co.).
Odontophotopsis viereckivi Baker, 1905. Invertebrata Pacifica 1: 97. d.

Genus ODONTOPHOTOPSIS Subgenus PERIPHOTOPSIS Schuster

Odontophotopsis subg. Periphotopsis Schuster, 1958. Ent. Amer. (n. s.) 37: 8 (in key), 60.
Type-species: Odontophotopsis (Periphotopsis) mamatus Schuster. Orig. desig.
mamata Schuster. Ariz., Nev., Calif.
Odontophotopsis (Periphotopsis) mamatus Schuster, 1958. Ent. Amer. (n. s.) 37: 60. d.

Genus PROTOPHOTOPSIS Schuster

Genus PROTOPHOTOPSIS Subgenus PROTOPHOTOPSIS Schuster

Protophotopsis Schuster, 1947. Ent. Soc. Amer., Ann. 39: 693.
Type-species: Protophotopsis scudderi Schuster. Orig. desig.

Only the typical subgenus occurs in North America.

Taxonomy: Schuster, 1949. Ent. Amer. (n. s.) 29: 82-85.
scudderi Schuster. Kans., Tex., Colo.
Protophotopsis scudderi Schuster, 1947. Ent. Soc. Amer., Ann. 39: 694. 6.

venenaria (Melander). Kans., Tex., Calif. Possibly the opposite sex of scudderi Schuster.
Mutilla venenaria Melander, 1903. Amer. Ent. Soc., Trans. 29: 320. 2.

Taxonomy: Schuster, 1949. Ent. Amer. (n. s.) 29: 83.
TRIBE PSEUDOMETHOCINI

All species are diurnal. The authenticated host records suggest that members of this tribe are
parasitic on various kinds of ground-nesting bees, principally halictids belonging to the subfami-
lies Halictinae and Nomiinae. Fattig, 1943 (Emory Univ. Mus. Bul. 1: 4) records frigida (Smith)
as having been reared from mud cells of the eumenid wasp Ancistrocerus lutonidus Boh.; this
may be based on a misidentification.

Revision: Mickel, 1924. U.S. Natl. Mus., Proc. 64(15): 1-51, pls. 1-4. —Mickel, 1935. Amer.
Ent. Soc., Trans. 61: 383-398.

Genus PS9EUDOMETHOCA Ashmead

Pseudomethoca Ashmead, 1896. Amer. Ent. Soc., Trans. 23: 181.
Type-species: Photopsis Cressoni Fox. Orig. desig. (=Pseudomethoca frigida
(Smith)).
Nomiaephagus Ashmead, 1899. N. Y. Ent. Soc., Jour. 7: 56.
Type-species: Mutilla (Sphaeropthalma) Sanbornii Blake. Orig. desig.
albicoma Mickel. Tex. (Rio Grande in Brewster Co.).
Pseudomethoca albicoma Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 29. ¢.
anthracina (Fox). Oreg., Calif.
Sphaerophthalma (!) anthracina Fox, 1892. Ent. News 3: 172. d.
Mutilla anthracicolor Dalla Torre, 1897. Cat. Hym., v. 8, p. 9. N. name for
Sphaerophthalma (!) anthracina Fox, not Mutilla anthracina Gerstaecker.
Mutilla harpalyce Fox, 1899. Amer. Ent. Soc., Trans. 25: 227. °.

Taxonomy: Hurd, 1951. Pan-Pacifie Ent. 27: 156 (synonymy).

athamas (Fox). Oreg., Calif.
Mutilla athamas Fox, 1899. Amer. Ent. Soc., Trans. 25: 225. d.

aureovestita Bradley. N. Mex., Ariz.
Pseudomethoca aureovestita Bradley, 1924. In Mickel, U.S. Natl. Mus., Proc. 64 (15): 22.

Oeror

bequaerti Mickel. Tex., N. Mex., Ariz., Colo., Mont.

Pseudomethoca bequaerti Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 13. 2.

1302 | Hymenoptera in America North of Mexico

brazoria (Blake). Okla., Tex.
Mutilla (Sphaeropthalma) brazoria Blake, 1871. Amer. Ent. Soe., Trans. 3: 255. 2.

carbonaria Mickel. Tex.
Pseudomethoca carbonaria Mickel, 1924. U.S. Natl. Mus., Proc. 64 (15): 43. d.
cephalargia Mickel. Ariz. (Santa Catalina Mts.).
Pseudomethoca cephalargia Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 17. 9.

connectens (Cresson). Calif.

Mutilla connectens Cresson, 1865. Ent. Soe. Phila., Proc. 4: 387. °.
contumax (Cresson). Iowa, S. Dak., Nebr., Kans., Tex., Colo., Ariz.; Mexico.

Mutilla contumax Cresson, 1865. Ent. Soc. Phila., Proc. 4: 437. 9.

Mutilla microphthalma Gerstaecker, 1874. Arch. Naturgesch. 40: 64. 3.
contumeliosa Mickel. Utah, Idaho, Calif.

Pseudomethoca contumeliosa Mickel, 1935. Amer. Ent. Soc., Trans. 61: 393. 9.
damia (Cameron). Ariz.; Mexico (Guerrero).

Sphaerophthalma (!) damia Cameron, 1894. Biol. Cent.-Amer., Hym., v. 2, p. 383. 3.

Mutilla sphaerophthalmica Dalla Torre, 1897. Cat. Hym., v. 8, p. 87. N. name.

dentifrontalis Bradley. Oreg., Calif.
Pseudomethoca dentifrontalis Bradley, 1924. In Mickel, U. S. Natl. Mus., Proe. 64 (15): 11.

OF
dentigula Mickel. Tex. (Trans-Pecos).
Pseudomethoca dentigula Mickel, 1935. Amer. Ent. Soc., Trans. 61: 391. 9.
donaeanae (Cockerell). Tex., N. Mex., Ariz., Calif.
Sphaerophthalma (!) donae-anae Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci.
Phila., Proc. 49: 127. °.
flammigera Mickel. Ariz.
Pseudomethoca flammigera Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 21. °.
flavida (Blake). Tex.
Mutilla (Sphaeropthalma) flavida Blake, 1871. Amer. Ent. Soc., Trans. 3: 249. ¢.
frigida (Smith). Southern Canada and U.S. east of Rocky Mts. Host: Dialictus zephyrus (Sm.),
D. coeruleus (Robt.), D. rohweri (Ellis)?, D. laevissimus (Sm.)?, D. versatus (Robt.), D.
imitatus (Sm.)?, Evylaeus cinctipes (Prov.), Augochlorella striata (Prov.)?
Mutilla frigida Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 60. 2.
Mutilla (Sphaeropthalma) canadensis Blake, 1871. Amer. Ent. Soc., Trans. 3: 252. 2.
Photopsis Cressoni Fox, 1890. Ent. News 1: 138. ¢.
Sphaerophthalma (!) alveolata Provancher, 1895. Nat. Canad. 22: 110. 2.
Mutilla neojerseiensis Dalla Torre, 1897. Cat. Hym., v. 8, p. 65. N. name for Phytopsis (!)
Cressoni Fox, not Mutilla Cressoni Blake.

Taxonomy: Brothers, 1972. Kans. Univ. Sci. Bul. 50: 25-31, figs. 11-40 (egg, larvae, pupae).

Biology: Melander and Brues, 1903. Biol. Bul. 5: 4-7, fig. 4 (behavior in host nest).
—Krombein, 1938. Brooklyn Ent. Soc., Bul. 33:14-15 (behavior in host nest). —Fattig, 1943.
Emory Univ. Mus. Bul. 1: 4 (questionable host record). —Shappirio, 1947. Brooklyn Ent.
Soc., Bul. 42: 163 (mating behavior). —Michener and Wille, 1961. Kans. Univ. Sci. Bul. 42:
1129-1130 (behavior in host nests). —Lin, 1964. Ins. Sociaux 11: 187-192 (behavior in host
nests). —Batra, 1965. Kans. Ent. Soc., Jour. 38: 383-386, figs. 6 a-c, 7 a-b (life history,
behavior). —Michener, 1966. Kans. Ent. Soc., Jour. 39: 196-197 (host record). —Knerer and
Atwood, 1967. Ent. Soc. Ontario, Proce. 97: 106 (host record). —Brothers, 1972. Kans. Univ.
Sei. Bul. 50: 1-25, figs. 1-10 (life history, behavior).
gila (Blake). Tex. (Brownsville).

Mutilla (Sphaeropthalma) Gila Blake, 1871. Amer. Ent. Soc., Trans. 3: 250. 6.
ilione (Fox). Tex., Okla.

Mutilla ilione Fox, 1899. Amer. Ent. Soc., Trans. 25: 268. °.

Mutilla aprica Melander, 1903. Amer. Ent. Soc., Trans. 29: 322. 2.

Taxonomy: Mickel, 1965. Ent. Soe. Wash., Proc. 67: 2 (synonymy).

klotsi Mickel. Colo. (Hugo).
Pseudomethoca klotsi Mickel, 1935. Amer. Ent. Soc., Trans. 61: 389. °.

Superfamily SCOLIOIDEA 1303

manca Mickel. Utah.

Pseudomethoca manca Mickel, 1924. U.S. Natl. Mus., Proc. 64 (15): 45. ¢.
meritoria Mickel. Tex.

Pseudomethoca meritoria Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 27. 2.
mulaiki Mickel. Tex. (Cameron Co.).

Pseudomethoca mulaiki Mickel, 1938. Ent. Soe. Amer., Ann. 31: 150. 2.
nephele (Fox). Tex.

Mutilla nephele Fox, 1899. Amer. Ent. Soc., Trans. 25: 223. °.
nigricula Mickel. Ariz.

Pseudomethoca nigricula Mickel, 1924. U.S. Natl. Mus., Proc. 64 (15): 26. d.
nudula Mickel. N. Mex. (Pecos). Host: Possibly Calliopsis sp.

Pseudomethoca nudula Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 15. @.
oceola (Blake). N. J. south to Fla., west to S. Dak. and Ariz.

Mutilla (Sphaeropthalma) Oceola Blake, 1871. Amer. Ent. Soe., Trans. 3: 248. d.

Mutilla hippodamia Fox, 1899. Amer. Ent. Soc., Trans. 25: 227. °.
oculata (Banks). N. C., Ga., Fla.

Nomiaephagus oculatus Banks, 1921. Ent. Soc. Amer., Ann. 14: 24. °.
oculissima Mickel. N. Mex. (Pecos).

Pseudomethoca oculissima Mickel, 1924. U. S. Natl. Mus., Proce. 64 (15): 14. @.
paludata Mickel. Nebr., Kans., Wyo., Colo., N. Mex., Ariz.

Pseudomethoca paludata Mickel, 1924. U. S. Natl. Mus., Proce. 64 (15): 29. °.

Pseudomethoca scrupulosa Mickel, 1924. U. S. Natl. Mus., Proce. 64 (15): 31. 6.
pigmentata Mickel. Tex.

Pseudomethoca pigmentata Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 24. °.
praeclara (Blake). Ariz., Calif.

Sphaerophthalma (!) praeclara Blake, 1886. Amer. Ent. Soc., Trans. 13: 252. 9.

Mutilla aegaeon Fox, 1899. Amer. Ent. Soc., Trans. 25: 229. 6.

Nomiaephagus acuum Cockerell, 1915. Entomologist 48: 250. °.

Pseudomethoca (Nomiaephagus) aegeon (!) Bradley, 1916. Amer. Ent. Soe., Trans. 42: 321.

3.
propinqua (Cresson). Minn. south to Tex., west to Alta. and Ariz. Host: Nomia m. melanderi
Ckll.

Mutilla propinqua Cresson, 1865. Ent. Soe. Phila., Proc. 4: 433. d.

Mutilla montivaga Cresson, 1865. Ent. Soc. Phila., Proc. 4: 436. °.

Pseudomethoca sacatona Caldwell, 1939. Pan-Pacifie Ent. 15: 94. 9°.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 103 (host record).
quadrinotata Mickel. Ariz. (Tucson).
Pseudomethoca quadrinotata Mickel, 1938. Ent. Soe. Amer., Ann. 31: 149. °.
russeola Mickel. Tex. (San Diego).
Pseudomethoca russeola Mickel, 1924. U. S. Natl. Mus., Proce. 64 (15): 44. d.
sanbornii aeetis (Fox). ?Conn., N. C. south to Fla., west to Tex. in L. Austr. Zone.

Mutilla aeetis Fox, 1899. Amer. Ent. Soc., Trans. 25: 228. 9.
Pseudomethoca (Nomiaephagus) aetis (!) Bradley, 1916. Amer. Ent. Soe., Trans. 42: 312,

316. 2.
Taxonomy: Krombein, 1953. Wasmann Jour. Biol. 10: 308 (reduced to subsp. rank).
sanbornii sanbornii (Blake). Mass. south to Fla., west to Nebr. and Tex. in Transition and U.
Austr. Zones. Host: Nomia pattoni Cockerell.
Mutilla (Sphaeropthalma) Sanbornii Blake, 1871. Amer. Ent. Soe., Trans. 3: 248. ¢.
Biology: Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 38 (host).

scaevolella (Cockerell and Casad). Tex., N. Mex.
Sphaerophthalma (!) scaevolella Cockerell and Casad, 1895. Amer. Ent. Soe., Trans. 22:
298. 9°.
simillima (Smith). Mass. south to Fla., west to Nebr. and Tex.
Mutilla simillima Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 62. 2.

1304 Hymenoptera in America North of Mexico

Mutilla geryon Fox, 1899. Amer. Ent. Soc., Trans. 25: 255. d.

Mutilla Henshawi Melander, 1903. Amer. Ent. Soc., Trans. 29: 303. 6.

Mutilla daeckei Rohwer, 1910. Ent. Soe. Wash., Proc. 12: 49. ¢. :

Pseudomethoca simillima Mickel, 1924. U. S. Natl. Mus., Proc. 64 (15): 32. 2 (¢ misdet.).
Taxonomy: Krombein, 1948. Ent. News 59: 187-189.

Biology: Shappirio, 1947. Brooklyn Ent. Soc., Bul. 42: 163 (mating behavior).

torrida Krombein. Fla.
Pseudomethoca frigida torrida Krombein, 1954. Amer. Ent. Soc., Trans. 80: 7, fig. 7. 2, d.

toumeyi (Fox). Tex., Ariz.; Mexico (Sonora).
Sphaerophthalma (!) toumeyi Fox, 1894. Ent. News 5: 297. °.
Taxonomy: Mickel, 1974. Ent. Soc. Amer., Ann. 67: 468. ¢.
vanduzei Bradley. Va., S. C., Ga., Fla., Tex. Host: Nomia maneei Ckll.
Pseudomethoca (Nomiaephagus) vanduzei Bradley, 1916. Amer. Ent. Soc., Trans. 42: 321.
3.
Pseudomethoca fattigi Mickel, 1938. Ent. Soc. Amer., Ann. 31: 147. °.
Pseudomethoca vanduzeei (!) Krombein, 1951. U. S. Dept. Agr., Monog. 2: 761. Emend.
Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 6 (host record).

wickhami (Cockerell and Casad). Tex. (Fedor).
Sphaerophthalma (!) wickhami Cockerell and Casad, 1895. Amer. Ent. Soc., Trans. 22: 297.
OR

Genus MYRMILLOIDES Andre

Myrmilloides Andre, 1903. In Wytsman, Gen. Ins., fase. 11, p. 26.
Type-species: Mutilla (Sphaeropthalma) grandiceps Blake. Orig. desig.

grandiceps (Blake). Iowa, Nebr., Colo., Kans., Mo., Okla., Ark., Tex., La., Tenn. Host: Dialictus
zephyrus (Sm.)?, Augochlorella striata (Prov.)?, A. persimilis (Vier.)?
Mutilla (Sphaeropthalma) grandiceps Blake, 1872. Amer. Ent. Soc., Trans. 4: 74. 9, d.
Biology: Melander, 1903. Amer. Ent. Soc., Trans. 29: 293 (host record). —Ordway, 1964. Kans.
Ent. Soc., Jour. 37: 149 (host records).

TRIBE DASYMUTILLINI
Members of this tribe are diurnal except for a few species of Dasymutilla.

Genus DASYMUTILLA Ashmead

Dasymutilla Ashmead, 1899. N. Y. Ent. Soc., Jour. 7: 57.

Type-species: Mutilla (Sphaeropthalma) Gorgon Blake. Orig. desig.
Bruesia Ashmead, 1903. Canad. Ent. 35: 306.

Type-species: Mutilla harmonia Fox. Orig. desig.
Pycnomutilla Ashmead, 1904. Canad. Ent. 36: 8.

Type-species: Mutilla (Sphaeropthalma) Waco Blake. Orig. desig.

Revision: Mickel, 1928. U. S. Natl. Mus., Bul. 143: 1-351, 5 pls., 28 text figs. —Mickel, 1936.
Ent. Soe. Amer., Ann. 29: 29-60, 3 figs. (revised key to spp.).

Taxonomy: Hurd, 1951. Calif. Ins. Survey Bul. 1: 89-118, 1 pl. (Calif. spp.)
Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14, p. 43, pl. 14 A-F (male
genitalia).
albiceris Mickel. Calif. (San Bernardino Co.); Mexico (Baja California).

Dasymutilla albiceris Mickel, 1936. Pan-Pacific Ent. 12: 91. 9.
alesia Banks. Mass. south to Fla., west to II. and Okla.
Dasymutilla alesia Banks, 1921. Ent. Soc. Amer., Ann. 14: 24. 9.
angulata Krombein. Mich., IIl., Iowa, Mo., S. Dak., Nebr., Kans.
Mutilla anguliceps Fox, 1899. Amer. Ent. Soc., Trans. 25: 240. 2. Preoce. by Andre, 1897.
Dasymutilla angulata Krombein, 1951. U. S. Dept. Agr., Monog. 2: 762. N. name.

Superfamily SCOLIOIDEA 1305

apicalata (Blake). Tex., Ariz.; Mexico (Guerrero).
Mutilla (Sphaeropthalma) apicalata Blake, 1871. Amer. Ent. Soc., Trans. 3: 238. ¢.
Sphaerophthalma (!) perfidiosa Cameron, 1895. Biol. Cent.-Amer., Hym., v. 2, p. 368. ¢.
Preoce. in Mutilla.
Sphaerophthalma (!) guerreroensis Cameron, 1895. Biol. Cent.-Amer., Hym., v. 2, p. 369. ¢.
Mutilla perfida Dalla Torre, 1897. Cat. Hym., v. 8, p. 72. N. name.

Taxonomy: Mickel, 1965. Ent. Soe. Wash., Proc. 67: 2 (synonymy).
arcana Mickel. Nebr., Okla., Tex., N. Mex., Colo.
Dasymutilla arcana Mickel, 1928. U.S. Natl. Mus., Bul. 143: 217. 3.
arenivaga Mickel. Ariz., Calif. D. megalophthalma Mick. may be the male.
Dasymutilla arenivaga Mickel, 1928. U.S. Natl. Mus., Bul. 143: 278. 9.
arenivaga var. unicolor Mickel. Ariz. (Ft. Mojave).
Dasymutilla arenivaga var. unicolor Mickel, 1936. Ent. Soc. Amer., Ann. 29: 59. 9.
asopus asopus (Cresson). Man., Minn., N. Dak., Mont., S. Dak., Nebr., Kans., Colo., Tex. Host:
Dianthidium sp.?
Mutilla Asopus Cresson, 1865. Ent. Soe. Phila., Proc. 4: 435. 6.
Ephuta sparsiformis Cockerell and Rohwer, 1908. Psyche 15: 4. 9.

Biology: Hicks, 1926. Colo. Univ., Studies, p. 249. —Mickel, 1928. U. S. Natl. Mus., Bul. 143:
61.
asopus bexar (Blake). Mass. south to Fla., Miss., Tex., Mo., Ind. Host: Anthophora abrupta
Say, A. ursina Cr.; Dianthidium curvatum sayi Ckll., Paranthidiwm j. jugatorium Say.
Mutilla (Sphaeropthalma) Bexar Blake, 1871. Amer. Ent. Soc., Trans. 3: 238. 3.
Mutilla harmonia Fox, 1899. Amer. Ent. Soe., Trans. 25: 229. 9.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 4-5 (host records). —Krombein, 1958. U. S.
Dept. Agr., Monog. 2, Sup. 1: 103 (host record). —Michener, 1975. Kans. Ent. Soce., Jour. 48:
199 (host).
asopus cassandra Mickel. Fla.
Dasymutilla cassandra Mickel, 1928. U. S. Natl. Mus., Bul. 143: 64. ¢.
asteria Mickel. Ariz. (Pima Co.). Host: Microbembex nigrifrons (Prov.).
Dasymutilla asteria Mickel, 1936. Ent. Soc. Amer., Ann. 29: 57. 2.

Biology: Haddock, 1967. In Krombein, U.S. Dept. Agr., Monog. 2, Sup. 2: 339 (host record).

atricauda Mickel. Calif., Ariz.
Dasymutilla atricauda Mickel, 1936. Pan-Pacifie Ent. 12: 92. °.

atrifimbriata Mickel. Kans., Ark., Okla., Tex., Ariz.
Dasymutilla atrifimbriata Mickel, 1928. U. S. Natl. Mus., Bul. 148: 169. °.

atrifulva Mickel. Ariz.
Dasymutilla atrifulva Mickel, 1928. U. S. Natl. Mus., Bul. 143: 290. 3.

aureola (Cresson). Nev., Oreg., Calif. Host: Anthophora stanfordiana Ckll., Melissodes sp.
Mutilla aureola Cresson, 1865. Ent. Soc. Phila., Proc. 4: 386. 9.
Sphaerophthalma (!) parmosa Blake, 1886. Amer. Ent. Soc., Trans. 13: 210. 2.
Sphaerophthalma (!) mollissima Blake, 1886. Amer. Ent. Soc., Trans. 13: 215. 3.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 103 (host records).

aureola var. pacifica (Cresson). B. C., Calif.

Mutilla pacifica Cresson, 1875. Amer. Ent. Soc., Trans. 5: 120. 2.
biguttata (Cockerell). Miss., La., Tex., Ark., Kans., Ariz.

Sphaerophthalma (!) quadriguttata var. biguttata Cockerell, 1895. Ent. News 6: 63. 9.
bioculata (Cresson). Man. south to La., west to B. C. and N. Mex. Host: Bembix pruinosa Fox,

B. americana spinolae Lep.?, Microbembex monodonta (Say).

Mutilla bioculata Cresson, 1865. Ent. Soc. Phila., Proce. 4: 431. 6.

Mutilla Aegina Cresson, 1865. Ent. Soc. Phila., Proc. 4: 435. 9.

Mutilla chlamydata Melander, 1903. Amer. Ent. Soc., Trans. 29: 299. 9.

1306 Hymenoptera in America North of Mexico

Biology: Mickel, 1924. Ent. News 35: 236-242. —Krombein, 1958. U. S. Dept. Agr., Monog. 2,
Sup. 1: 103 (host record). —Cottrell, 1972. Jn Brothers, Kans. Univ. Sci. Bul. 50: 31-36, figs.
41-42 (life history).
birkmani (Melander). Nebr., Wyo., Kans., Okla., Tex.

Mutilla Birkmani Melander, 1903. Amer. Ent. Soc., Trans. 29: 313. 9.
bollii (Fox). Miss., La., Tex., Okla., Kans.

Mutilla Bollii Fox, 1899. Amer. Ent. Soc., Trans. 25: 242. 2.

Dasymutilla blawana Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 461. 9.
bonita Mickel. Ariz.

Dasymutilla bonita Mickel, 1928. U. S. Natl. Mus., Bul. 143: 208. 9°.

Dasymutilla poliothrix Mickel, 1928. U.S. Natl. Mus., Bul. 143: 285. 6.
californica (Radoszkowski). B. C., Oreg., Utah, Calif.

Mutilla Californica Radoszkowski, 1861. Soc. Ent. Rossica, Horae 1: 86. 2.

Dasymutilla abdita Mickel, 1928. U. S. Natl. Mus., Bul. 143: 293.
californica var. clio (Blake). B. C., Idaho, Nev., Calif.

Mutilla Clio Blake, 1879. Amer. Ent. Soc., Trans. 7: 251. 9.
calorata Mickel. Kans., Okla., Tex., N. Mex., Colo.

Dasymutilla calorata Mickel, 1928. U. S. Natl. Mus., Bul. 148: 227. °.
campanula Mickel. Kans., Colo.

Dasymutilla campanula Mickel, 1928. U. S. Natl. Mus., Bul. 143: 178. °.
candida Mickel. Ariz (Douglas).

Dasymutilla candida Mickel, 1928. U.S. Natl. Mus., Bul. 143: 296. 3.
canella (Blake). Mass. to Pa., west to Man. and Tex. Host: Dialictus pruinosus (Robt.)?

Mutilla (Sphaeropthalma) canella Blake, 1871. Amer. Ent. Soc., Trans. 3: 239. 6.

Mutilla rugulosa Fox, 1899. Amer. Ent. Soc., Trans. 25: 240. 2.

Mutilla infensa Melander and Brues, 1903. Biol. Bul. 5: 24. 2.

Biology: Viereck, 1902. Ent. News 13: 72. —Melander and Brues, 1903. Biol. Bul. 5: 25

(putative host).

caneo (Blake). Minn., N. Dak., south to Tex. and N. Mex., west to Alta. and B. C.
Mutilla caneo Blake, 1879. Amer. Ent. Soc., Trans. 7: 250. @.
Mutilla mixtura Blake, 1879. Amer. Ent. Soc., Trans. 7: 251. 9°.
Mutilla myrrha Fox, 1899. Amer. Ent. Soc., Trans. 25: 258. 9.

castor (Blake). S. C., Ga., Fla., Ala., Miss., La., Tex. Host: Polistes fuscatus (F.).
Mutilla (Sphaeropthalma) castor Blake, 1871. Amer. Ent. Soe., Trans. 3: 237. 6.
Dasymutilla mediatoria Mickel, 1938. Ent. Soc. Amer., Ann. 31: 152. °.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 6 (host record).
chattahoochei Bradley. N. C. south to Fla.
Dasymutilla (Dasymutilla) chattahoochei Bradley, 1916. Amer. Ent. Soc., Trans. 42: 324.

oe
Dasymutilla (Dasymutilla) arenerronea Bradley, 1916. Amer. Ent. Soc., Trans. 42: 324. 9.

Taxonomy: Krombein, 1954. Amer. Ent. Soc., Trans. 80: 9, fig. 5 (description male).

chiron (Blake). Tex.
Mutilla (Sphaeropthalma) Chiron Blake, 1872. Amer. Ent. Soc., Trans. 4: 72. 3.
Mutilla Ursula var. texana Cresson, 1875. Amer. Ent. Soc., Trans. 5: 120. 6.
chiron var. ursula (Cresson). B. C. south to Ariz. and Calif. and southeast to Tex.
Mutilla Ursula Cresson, 1875. Amer. Ent. Soc., Trans. 5: 120. 6.

chisos Mickel. Tex.
Dasymutilla chisos Mickel, 1928. U.S. Natl. Mus., Bul. 143: 284. ¢.
clotho (Blake). Okla., Tex.
Mutilla (Sphaeropthalma) Clotho Blake, 1872. Amer. Ent. Soc., Trans. 4: 72. 2.
clytemnestra (Fox). Oreg., Calif.
Mutilla clytemnestra Fox, 1899. Amer. Ent. Soc., Trans. 25: 246. °.
coccineohirta (Blake). Idaho, Nev., Calif., Oreg., Wash.
Mutilla (Sphaeropthalma) coccineohirta Blake, 1871. Amer. Ent. Soe., Trans. 3: 235. ¢.

Superfamily SCOLIOIDEA 1307

Mutilla ochracea Blake, 1879. Amer. Ent. Soc., Trans. 7: 247. 6.
Sphaerophthalma (!) venifica Blake, 1886. Amer. Ent. Soe., Trans. 13: 210. °.
Mutilla progne Fox, 1899. Amer. Ent. Soc., Trans. 25: 247. 2.
Dasymiutilla aletina Cockerell, 1915. Entomologist 48: 249. ¢.
columbiana Mickel. B. C. (Nicola).
Dasymutilla columbiana Mickel, 1928. U. S. Natl. Mus., Bul. 143: 119. d.
connectens (Cameron). Ariz.; Mexico (Chihuahua, Jalisco, Baja California).
Sphaerophthalma (!) connectens Cameron, 1895. Biol. Cent.-Amer., Hym., v. 2, p. 362. 9°.
Secondary homonym in Mutilla.
Mutilla eggeri Dalla Torre, 1897. Cat. Hym., v. 8, p. 33. N. name.
Dasymutilla helva Mickel, 1928. U.S. Natl. Mus., Bul. 148: 259. °.

Taxonomy: Mickel, 1965. Ent. Soc. Wash., Proc. 67: 2 (synonymy).

corcyra Mickel. Tex.

Dasymutilla corcyra Mickel, 1928. U.S. Natl. Mus., Bul. 143: 180. 9.
cotulla Mickel. Tex.

Dasymutilla cotulla Mickel, 1928. U. S. Natl. Mus., Bul. 143: 75. 6.
creon (Blake). Tex., Ark.

Mutilla (Sphaeropthalma) Creon Blake, 1872. Amer. Ent. Soc., Trans. 4: 73. d.
creusa (Cresson). Ark., Tex., Okla., Kans., Nebr., Colo., N. Mex.

Mutilla Creusa Cresson, 1865. Ent. Soc. Phila., Proc. 4: 431. 9.
creusa var. bellona (Cresson). Ark., Kans., Nebr., Wyo., Colo. Host: Bembix amoena Handl.

Possibly a synonym of bioculata (Cr.).
Mutilla Bellona Cresson, 1865. Ent. Soe. Phila., Proc. 4: 434. 2.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, p. 288 (host record).
curialis Mickel. N. Mex. (Jemez Springs).
Dasymutilla curialis Mickel, 1928. U.S. Natl. Mus., Bul. 143: 274. 6.
curticeps Mickel. Kans., Tex.
Dasymutilla curticeps Mickel, 1928. U.S. Natl. Mus., Bul. 143: 178. 2.
cypris (Blake). N. J., W. Va., Ga., Fla., Miss., Mo., Minn. Host: Bembix cinerea Handl.
Mutilla (Sphaeropthalma) Cypris Blake, 1871. Amer. Ent. Soc., Trans. 3: 246. °.
Dasymutilla mutata miamensis Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 462. °.
Dasymutilla hora Mickel, 1928. U.S. Natl. Mus., Bul. 148: 196. d.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 4, 7 (host record, mating behavior).
dammersi Mickel. Calif. (Palm Springs).
Dasymutilla dammersi Mickel, 1936. Pan-Pacific Ent. 12: 94. 2.
dawsoni Mickel. Nebr. (Halsey).
Dasymutilla dawsoni Mickel. 1936. Ent. Soc. Amer., Ann. 29: 40. 2.
digressa Mickel. N. Mex. (Pecos).
Dasymutilla digressa Mickel, 1928. U. S. Natl. Mus., Bul. 148: 273. d.
dilucida Mickel. Ariz.
Dasymutilla dilucida Mickel, 1928. U. S. Natl. Mus., Bul. 143: 267. °.
dionysia Mickel. Ariz.
Dasymutilla dionysia Mickel, 1928. U.S. Natl. Mus., Bul. 143: 104. °.
dorippa Mickel. Nev. (Ormsby Co.).
Dasymutilla dorippa Mickel, 1928. U.S. Natl. Mus., Bul. 143: 297. ¢.
dugesii (Cockerell and Casad). Kans., Colo., Tex., Ariz. Host: Anthophora occidentalis Cr.
Sphaerophthalma (!) dugesii Cockerell and Casad. 1894. Ent. News 5: 294. °.
Ephuta (Ephuta) Dugesi (!) Andre, 1903. In Wytsman, Gen. Ins., fase. 11, p. 59.

Biology: Krombein, 1967. U. S. Dept. Agr., Monog. 2, Sup. 2: 339 (host record).
electra (Blake). Tex., Okla.
Mutilla Electra Blake, 1872. Amer. Ent. Soc., Trans. 4: 75. 2.

eminentia Mickel. Ariz., Calif.; Mexico (Baja California).
Dasymutilla eminentia Mickel, 1928. U.S. Natl. Mus., Bul. 143: 79. 6.

1308 Hymenoptera in America North of Mexico

errabunda Mickel. Ariz., Calif.

Dasymutilla errabunda Mickel, 1928. U.S. Natl. Mus., Bul. 143: 269. d.
eurynome Mickel. Tex. (Valentine in Presidio Co.).

Dasymutilla eurynome Mickel, 1928. U.S. Natl. Mus., Bul. 143: 209. 9.
fasciventris Mickel. Ariz. (Globe).

Dasymutilla fasciventris Mickel, 1938. Ent. Soc. Amer., Ann. 31: 155. 6.
ferruginea (Smith). Ariz.; Mexico (Puebla, Oaxaca).

Mutilla ferruginea Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 226. 9.

Dasymutilla chrysocoma Mickel, 1928. U. S. Natl. Mus., Bul. 143: 266. 9.

Taxonomy: Mickel, 1965. Ent. Soc. Wash., Proc. 67: 3 (synonymy).

flammifera Mickel. Idaho, Oreg., Calif., Ariz.
Dasymutilla flammifera Mickel, 1928. U. S. Natl. Mus., Bul. 143: 240. 9.
foxi (Cockerell). N. Mex., Ariz. Host: Diadasia diminuta (Cr.).
Sphaerophthalma (!) fori Cockerell, 1894. Ent. News 5: 199. 6.
Sphaerophthalma (!) heterochroa Cockerell and Casad, 1894. Ent. News 5: 298. 2.
Sphaerophthalma (!) foxi var. arizonica Cockerell, 1900. Entomologist 33: 65. 2.

Biology: Cockerell, 1896. Nature 54: 461.
fulgida Mickel. Ariz. Possibly a synonym of erythrina (Say) from Mexico.
Dasymutilla fulgida Mickel, 1928. U.S. Natl. Mus., Bul. 143: 76. ¢.

Taxonomy: Mickel, 1974. Ent. Soc. Amer., Ann. 67: 470. 6.

furina Mickel. Ariz. (Douglas).
Dasymutilla furina Mickel, 1928. U. S. Natl. Mus., Bul. 143: 82. 9.
gentilis Mickel. Kans., Tex., Rocky Mts.
Dasymutilla gentilis Mickel, 1928. U.S. Natl. Mus., Bul. 148: 199. 6.
gibbosa (Say). N. H. south to Ga., west to Minn. and Mo.
Mutilla (Ephuta) gibbosa Say, 1836. Boston Jour. Nat. Hist. 1: 298. d.
Mutilla cariniceps Fox, 1899. Amer. Ent. Soc., Trans. 25: 241. @.
Dasymutilla scrobinata Rohwer, 1912. U.S. Natl. Mus., Proe. 41: 462. °.
Dasymutilla carniceps (!) Fattig, 1948. Emory Univ. Mus. Bul. 1: 9, 16. 9°.
gloriosa (Saussure). Tex., Utah, Nev., Ariz., Calif.; Mexico (Baja California). Host:
Microbembex nigrifrons (Prov.)
Mutilla gloriosa Saussure, 1868 (1867). Soc. Ent. France, Ann. (4) 7: 359. @.
Mutilla tecta Cresson, 1875. Amer. Ent. Soc., Trans. 5: 119. °.
Dasymutilla reperticia Mickel, 1928. U. S. Natl. Mus., Bul. 148: 287. ¢.

Taxonomy: Mickel, 1936. Ent. Soc. Amer., Ann. 29: 56 (synonymy).

Biology: Haddock, 1967. In Krombein, U.S. Dept. Agr., Monog. 2, Sup. 2: 339 (host record).

glycera Mickel. Ariz. (Florence).
Dasymutilla glycera Mickel, 1928. U.S. Natl. Mus., Bul. 148: 299. ¢.
gorgon (Blake). La., Okla., Tex., N. Mex.
Mutilla (Sphaeropthalma) Gorgon Blake, 1871. Amer. Ent. Soc., Trans. 3: 233. ?.
Mutilla Tisiphone Blake, 1879. Amer. Ent. Soc., Trans. 7: 249. 2.
Dasymutilla gorgons (!) Ashmead, 1899. N. Y. Ent. Soc., Jour. 7: 57, 60. 2, 3.
hector (Blake). Iowa, Nebr., Kans., Colo.
Mutilla (Sphaeropthalma) Hector Blake, 1871. Amer. Ent. Soc., Trans. 3: 237. ¢.
heliophila (Cockerell). Ariz., Calif.
Sphaerophthalma (!) heliophila Cockerell, 1900. Entomologist 33: 65. 2.
Dasymutilla welltonensis Bradley and Bequaert, 1923. Amer. Mus. Novitates 82: 1. 9.
hersilia Mickel. Tex. (Bexar Co.).
Dasymutilla hersilia Mickel, 1936. Ent. Soc. Amer., Ann. 29: 51. ¢.
hispidaria Mickel. Tex. (Hondo).
Dasymutilla hispidaria Mickel, 1936. Ent. Soc. Amer., Ann. 29: 55. @.

homole Mickel. N. Mex. (Organ Mts.).
Dasymutilla homole Mickel, 1928. U. S. Natl. Mus., Bul. 143: 72. 9.

Superfamily SCOLIOIDEA ~— 1309

intermixta Mickel. N. Mex., Ariz.

Dasymutilla intermixta Mickel, 1928. U.S. Natl. Mus., Bul. 148: 256. d.
interrupta Banks. Mass. south to Ga., west to N. Dak. and Colo.

Dasymutilla cypris var. interrupta Banks, 1921. Ent. Soc. Amer., Ann. 14: 24. @.

Taxonomy: Krombein, 1953. Wasmann Jour. Biol. 10: 310. 3.

klugii (Gray). Kans. south to Tex., west to Utah and Ariz. Host: Sphecius grandis (Say).
Mutilla Klugii Gray, 1872. In Griffith, Cuvier’s Animal Kingdom 15 (Insecta 2), p. 516. °.
Mutilla Orcus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 428. 2, d.

Biology: Bradley, 1920. Ent. News 31: 112-113 (host record).

klugioides Mickel. Okla., Tex.
Dasymutilla klugioides Mickel, 1936. Ent. Soc. Amer., Ann. 29: 54. 2.

leda (Blake). S. Dak. south to Tex., Colo., N. Mex.
Mutilla (Sphaeropthalma) Leda Blake, 1872. Amer. Ent. Soc., Trans. 4: 72. @.
Mutilla prognoides Viereck, 1906. Amer. Ent. Soc., Trans. 32: 186. 2.

lepeletierii (Fox). Mass. south to Fla., west to Iowa and Tex.
Mutilla fenestrata Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 627. ¢. Preoce.
Mutilla Lepeletierti Fox, 1899. Amer. Ent. Soc., Trans. 25: 244. N. name.
Mutilla vierecki Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 49. °.
Dasymutilla ferrugata var. balabetei Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 456. 9.
Dasymutilla georgiana Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 456. °.
Dasymutilla plesia Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 456. 2.
Dasymutilla (Dasymutilla) ferrugata var. ballabetei (!) Bradley, 1916. Amer. Ent. Soc.,
Trans. 42: 327. 9.

Biology: Shappirio, 1948. Brooklyn Ent. Soc., Bul. 42: 163 (mating).

macilenta (Blake). Ga., Fla., Tex.

Mutilla (Sphaeropthalma) macilenta Blake, 1871. Amer. Ent. Soc., Trans. 3: 239. d.
macra (Cresson). N. Dak. south to Ark., west to Alta. and N. Mex.

Mutilla macra Cresson, 1865. Ent. Soc. Phila., Proc. 4: 429. 3.

Sphaerophthalma (!) hispida Blake, 1886. Amer. Ent. Soc., Trans. 13: 226. d.
magna (Cresson). Ariz., Calif.; Mexico (Baja California).

Mutilla magna Cresson, 1865. Ent. Soc. Phila., Proc. 4: 385. 9.
magnifica Mickel. N. Mex., Ariz., Nev., Calif.; Mexico (Baja California).

Dasymutilla magnifica Mickel, 1928. U. S. Natl. Mus., Bul. 143: 234. 9, d.
medea (Cresson). Colo., N. Mex. Host: Microbembex monodonta (Say)?

Mutilla Medea Cresson, 1865. Ent. Soc. Phila., Proc. 4: 432. °.

Biology: Melander, 1903. Amer. Ent. Soc., Trans. 29: 297 (host ?).

megalophthalma Mickel. Ariz., Calif. D. arenivaga Mick. may be the female.
Dasymutilla megalophthalma Mickel, 1928. U.S. Natl. Mus., Bul. 143: 282. d.
melanippe Mickel. Tex.
Dasymutilla melanippe Mickel, 1928. U. S. Natl. Mus. Bul. 148: 152. 9.
Dasymutilla lauta Mickel, 1928. U.S. Natl. Mus., Bul. 143: 154. @.

Taxonomy: Mickel, 1936. Ent. Soc. Amer., Ann. 29: 46 (synonymy).

melanippe var. conformis Mickel. Tex.
Dasymutilla melanippe var. conformis Mickel, 1928. U. S. Natl. Mus., Bul. 148: 153. @.

meracula Mickel. Kans., Okla., Tex.
Dasymutilla meracula Mickel, 1928. U. S. Natl. Mus., Bul. 143: 201. 6.
monticola (Cresson). Minn. south to Kans., west to B. C. and Ariz.
Mutilla monticola Cresson, 1865. Ent. Soe. Phila., Proc. 4: 430. ¢.
Mutilla eximia Blake, 1886. Amer. Ent. Soc., Trans. 13: 200. d.
Ephuta boulderensis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 133. d.

montivagoides (Viereck). Kans. (Hamilton Co.).
Mutilla (Timulla) montivagoides Viereck, 1906. Amer. Ent. Soc., Trans. 32: 185. @.

1310 | Hymenoptera in America North of Mexico

mutata (Blake). N. Y. south to Fla., west to Mo. and Tex. Host: Myzinum obscurum (F.).
Mutilla (Sphaeropthalma) mutata Blake, 1871. Amer. Ent. Soc., Trans. 3: 247. °.
Dasymutilla allardi Rohwer, 1912. U. S. Natl. Mus., Proc. 41: 463. °.

Dasymutilla rubricosa Mickel, 1928. U. S. Natl. Mus., Bul. 143: 192. ¢.

Taxonomy: Mickel, 1938. Ent. Soe. Amer., Ann. 31: 152. 2.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 4, 7 (host record, mating).

myrice Mickel. Nebr., Kans., Okla., Tex., Colo.
Dasymutilla myrice Mickel, 1928. U.S. Natl. Mus., Bul. 148: 121. 6.
nigricauda (Viereck). Kans., Okla., Tex.
Mutilla (Timulla) nigricauda Viereck, 1906. Amer. Ent. Soc., Trans. 32: 187. °.
nigripes (Fabricius). Mass. to Fla., west to Alta. and Ariz. Host: Philanthus gibbosus (F.)?,
Cerceris f. flavofasciata Sm.
Mutilla nigripes Fabricius, 1787. Mantissa Insectorum, v. 1, p. 313. 2.
Mutilla sparsa Fox, 1899. Amer. Ent. Soc., Trans. 25: 240. °.
Dasymutilla blawa Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 457. °.
Dasymutilla segregata Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 459. 2.
Dasymutilla segregata finni Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 459. 9.
Dasymutilla bruneri Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 460. 2.
Dasymutilla champlaini Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 461. °.
Dasymutilla hirticula Mickel, 1923. Minn. State Ent. Rpt. 19: 107. d.

Taxonomy: Mickel, 1936. Ent. Soc. Amer., Ann. 29: 43-44 (synonymy).

Biology: Shappirio, 1948. Brooklyn Ent. Soc., Bul. 42: 163 (mating behavior). —Shappirio,
1949. Brooklyn Ent. Soe., Bul. 43: 158 (host record). —Krombein, 1959. Ent. Soc. Wash.,
Proce. 61: 198 (host record). —Evans and Miller, 1969. Mich. Ent. 2: 74 (adult female
overwintering). —Evans, Miller and Bartlett, 1973. Kans. Ent. Soc., Jour. 46: 343-346 (host
searching range).

nitidula Mickel. Kans., Okla., Tex.
Dasymutilla nitidula Mickel, 1928. U.S. Natl. Mus., Bul. 143: 174. 9.
nitidula var. nigridia Mickel. Tex.
Dasymutilla nitidula var. nigridia Mickel, 1936. Ent. Soc. Amer., Ann. 29: 47. °.
nocturna Mickel. Calif., Nev.
Dasymutilla nocturna Mickel, 1928. U.S. Natl. Mus., Bul. 143: 279. °.
nogalensis Mickel. Ariz.
Dasymutilla nogalensis Mickel, 1928. U.S. Natl. Mus., Bul. 143: 258. 9.
nupera Mickel. Tex., N. Mex.
Dasymutilla nupera Mickel, 1928. U. S. Natl. Mus., Bul. 143: 103. @.
occidentalis comanche (Blake). Iowa and S. Dak., south to Tex., Colo., Ariz., west to Idaho
and Oreg.
Mutilla (Sphaeropthalma) Comanche Blake, 1871. Amer. Ent. Soc., Trans. 3: 234. 2, d.
occidentalis occidentalis (Linnaeus). Conn. south to Fla., west to Mo. and Tex. Host: Bombus
fraternus Smith.
Mutilla occidentalis Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 582. 2.
Mutilla bifasciata Swederus, 1787. Svensk. Vetensk. Akad. Hand. 8: 285. 9.
Mutilla coccinea Fabricius, 1793. Ent. System. 2: 366. °.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 5 (host).

ocydrome Mickel. Ariz.

Dasymutilla ocydrome Mickel, 1928. U.S. Natl. Mus., Bul. 143: 292. ¢.
paenulata Mickel. Ariz., Calif.

Dasymutilla paenulata Mickel, 1928. U. S. Natl. Mus., Bul. 143: 206. °.
paranocturna Barr and Hurd. Calif.

Dasymutilla paranocturna Barr and Hurd, 1947. Pan-Pacific Ent. 23: 88. °.

parksi Mickel. Tex. (Bexar Co.).
Dasymutilla parksi Mickel, 1936. Ent. Soc. Amer., Ann. 29: 50. °.

Superfamily SCOLIOIDEA 1311

perilla Mickel. Tex.

Dasymutilla perilla Mickel, 1928. U.S. Natl. Mus., Bul. 1438: 300. 3.
perilla var. gentilicia Mickel. Kans., Colo.

Dasymutilla perilla var. gentilicia Mickel, 1928. U.S. Natl. Mus., Bul. 143: 301. d.
phaon (Fox). Tex., N. Mex., Ariz., Utah.

Mutilla phaon Fox, 1899. Amer. Ent. Soc., Trans. 25: 243. 6.
phaon var. fimbrialis Mickel. Utah, Ariz., Calif.

Dasymutilla phaon var. fimbrialis Mickel, 1928. U.S. Natl. Mus., Bul. 148: 302. 6.
phoenix (Fox). Ariz., Calif. Host: Ptilothrix sumichrasti (Cr.).

Mutilla phoenix Fox, 1899. Amer. Ent. Soc., Trans. 25: 247. 9.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 104 (host record).

polia Mickel. Ariz.

Dasymutilla polia Mickel, 1928. U.S. Natl. Mus., Bul. 143: 215. 6.
praegrandis Mickel. Tex.

Dasymutilla praegrandis Mickel, 1928. U.S. Natl. Mus., Bul. 143: 148. 9, 6.
praegrandis var. russata Mickel. Tex.

Dasymutilla praegrandis var. russata Mickel, 1928. U.S. Natl. Mus., Bul. 148: 152. 6.
pseudopappus (Cockerell). Colo., N. Mex., Ariz., Calif.

Sphaerophthalma (!) gloriosa var. pseudopappus Cockerell, 1895. Psyche 7(sup.): 6. 2.
pyrrhus (Fox). Ga., Fla. Host: Sp. of Bembicini.

Mutilla pyrrhus Fox, 1899. Amer. Ent. Soe., Trans. 25: 243. d.

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 2-4, fig. 1 (host record).
quadriguttata (Say). Ill., Minn., and S. Dak. south to Miss. and Tex. Host: Myzinwm
quinquecinctum (F.).
Mutilla quadriguttata Say, 1823. West. Quart. Rptr. 2: 74. °.
Dasymutilla permista Mickel, 1923. Minn. State Ent. Rpt. 19: 108. d.

Taxonomy: Remington, 1944. Ent. Soe. Amer., Ann. 37: 198-200 (synonymy).

Biology: Swezey, 1915. Hawaii Ent. Soc., Proc. 3: 72 (host record; misdet. as castor Blake.)

—Hayes, 1924. Brooklyn Ent. Soc., Bul. 19: 153 (host).

reclusa Mickel. S. Dak. south to Tex., Mont., Wyo.
Dasymutilla reclusa Mickel, 1928. U.S. Natl. Mus., Bul. 1438: 219. ¢.

rubicunda Bradley. Ga., Fla.
Dasymutilla (Dasymutilla) rubicunda Bradley, 1916. Amer. Ent. Soc., Trans. 42: 325. 9.
Dasymutilla beameri Mickel, 1936. Ent. Soc. Amer., Ann. 29: 48. °.

sackenii (Cresson). Oreg., Calif., Nev.; Mexico (Baja California). Host: Bembix occidentalis

beutenmulleri Fox.

Mutilla Sackenti Cresson, 1865. Ent. Soe. Phila., Proc. 4: 385. 2.
Mutilla erudita Cresson, 1875. Amer. Ent. Soc., Trans. 5: 120. @.

Biology: Bohart and MacSwain, 1939. South. Calif. Acad. Sci., Bul. 38: 89, pl. 16, figs. 11-14
(life history).
saetigera Mickel. Ariz. (Baboquivari Mts.).
Dasymutilla saetigera Mickel, 1928. U. S. Natl. Mus., Bul. 148: 211. 9.
satanas Mickel. Ariz., Nev., Calif.; Mexico (Baja California).
Dasymutilla satanas Mickel, 1928. U. S. Natl. Mus., Bul. 143: 239. 9.
Dasymutilla mimula Mickel, 1928. U.S. Natl. Mus., Bul. 148: 255. ¢.

Taxonomy: Barr and Hurd, 1947. Pan-Pacific Ent. 23: 86. ?, d.
scaevola (Blake). Mass. south to Ga., west to Mont. and Ariz. Host: Cerceris clypeata Dahlb.?,
C. finitima Cr.?

Mutilla (Sphaeropthalma) obscura Blake, 1871. Amer. Ent. Soc., Trans. 3: 239. 6. Preoce.
Mutilla (Sphaeropthalma) scaevola Blake, 1871. Amer. Ent. Soc., Trans. 3: 247. @.
Mutilla (Sphaeropthalma) admetus Blake, 1872. Amer. Ent. Soc., Trans. 4: 74. d.
Sphaerophthalma (!) macer (!) Blake, 1886. Amer. Ent. Soc., Trans. 13: 227. 6.
Sphaerophthalma (!) macerata Blake, 1886. Amer. Ent. Soc., Trans. 13: 286. Emend.

1312 Hymenoptera in America North of Mexico

Mutilla macera (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 56.
Mutilla (Dasymutilla) apachea Viereck, 1908. Amer. Ent. Soc., Trans. 33: 386. 6.

Biology: Shappirio, 1948. Brooklyn Ent. Soc., Bul. 42: 162-163 (host record). —Shappirio,
1949. Brooklyn Ent. Soc., Bul. 43: 157-158 (host records).

scitula Mickel. Utah, Nev., Calif.
Dasymutilla scitula Mickel, 1928. U.S. Natl. Mus., Bul. 143: 244. 2, ¢.
serenitas Mickel. Tex. (Cotulla, Eastland Co.).
Dasymutilla serenitas Mickel, 1928. U. S. Natl. Mus. , Bul. 143: 289. 3.
sicheliana (Saussure). Ariz.; Mexico (Durango).
Mutilla Sicheliana Saussure, 1868 (1867). Soe. Ent. France, Ann. (4) 7: 360. @.
Sphaerophthalma (!) prunotincta Cockerell, 1895. Ent. News 6: 60. 2.
Sphaerophthalma (!) thera Cameron, 1895. Biol. Cent.-Amer., Hym., v. 2, p. 358. 2. Preoce.
in Mutilla. ,
Mutilla gynaecologica Dalla Torre, 1897. Cat. Hym., v. 8, p. 45. N. name.

Taxonomy: Mickel, 1965. Ent. Soc. Wash., Proc. 67: 3 (synonymy).
snoworum (Cockerell). Nebr. south to Tex., west to Mont. and Ariz.
Sphaerophthalma (!) snoworum Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci.
Phila., Proc. 49: 135. 6.
Mutilla poecilonota Melander, 1903. Amer. Ent. Soc., Trans. 29: 301. @.

sophrona Mickel. Ariz.
Dasymutilla sophrona Mickel, 1928. U.S. Natl. Mus., Bul. 143: 271. ¢.
stevensi Mickel. Iowa, N. Dak., south to Tex., Colo., Utah, N. Mex., Ariz.
Dasymutilla stevensi Mickel, 1928. U. S. Natl. Mus., Bul. 143: 99. 2.
Dasymutilla medora Mickel, 1928. U. S. Natl. Mus., Bul. 143: 101. 6.

Taxonomy: Mickel, 1936. Ent. Soc. Amer., Ann. 29: 44.(synonymy).

subhyalina Mickel. Calif. (Andrade).

Dasymutilla subhyalina Mickel, 1928. U.S. Natl. Mus., Bul. 143: 281. 6.
sulcatulla Mickel. Tex. (Rio Grande in Brewster Co.).

Dasymutilla sulcatulla Mickel, 1928. U. S. Natl. Mus., Bul. 143: 155. 9.

testaceiventris (Fox). Oreg., Calif.
Mutilla testaceiventris Fox, 1899. Amer. Ent. Soc., Trans. 25: 242. ¢.
texanella Mickel. Kans., Tex., Colo.
Mutilla texana Blake, 1879. Amer. Ent. Soc., Trans. 7: 250. 2. Preoce.
Dasymutilla texanella Mickel, 1928. U. S. Natl. Mus., Bul. 143: 96. 2. N. name.
thetis (Blake). Ariz.
Sphaerophthalma (!) thetis Blake, 1886. Amer. Ent. Soc., Trans. 13: 214. °.
vandala Mickel. Tex.
Dasymutilla vandala Mickel, 1928. U.S. Natl. Mus., Bul. 143: 74. ¢.
vesta errans Rohwer. S. Dak. south to Tex., Colo.
Dasymiutilla errans Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 457. 9.
Dasymutilla bosquensis Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 457. @.
Dasymutilla texensis Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 460. @.

vesta sappho (Fox). N. C. south to Fla., Miss. Host: Bembix cinerea Handl.
Mutilla sappho Fox, 1899. Amer. Ent. Soc., Trans. 25: 239. 2.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 104 (host).
vesta vesta (Cresson). East of Rocky Mts. in Canada and U.S. Host: Trypargilum politwm

(Say); Nomia m. melanderi Ckll.

Mutilla Vesta Cresson, 1865. Ent. Soc. Phila., Proc. 4: 436. 2.

Scolia wnicincta Provancher, 1882. Nat. Canad. 13: 6. 3 (2 misdet.).

Mutilla monozona Dalla Torre, 1897. Cat. Hym., v. 8, p. 64. N. name for Scolia wnicincta
Provancher, not Mutilla wnicincta Lueas.

Mutilla agenor Fox, 1899. Amer. Ent. Soe., Trans. 25: 245. 6.

Mutilla zella Rohwer, 1910. Ent. Soc. Wash., Proe. 12: 50. 2.

Pycnomutilla harmoniiformis Rohwer, 1912. U. S. Natl. Mus., Proe. 41: 455. ¢.

Superfamily SCOLIOIDEA 1313

Dasymutilla ferrugatella Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 458. 9.

Dasymutilla coloradella Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 458. @.

Dasymutilla coloradella virginica Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 459. °.
Dasymutilla coloradella kamloopsensis Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 459. 2.
Dasymutilla mesillae Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 461. @.

Dasymutilla carolina Rohwer, 1912. U. S. Natl. Mus., Proc. 41 462. 9.

Biology: Fattig, 1943. Emory Univ. Mus. Bul. 1: 4 (host record). —Krombein, 1958. U.S.
Dept. Agr., Monog. 2, Sup. 1: 104 (host record).
vestita (Lepeletier). N. Dak. south to Tex., west to Alta. and Calif., south in Mexico to Oaxaca.

Host: Anthophora occidentalis (Cr.), Diadasia enavata (Cr.); Megachile perihirta Ckll.

Mutilla vestita Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 634. 2, 6.

Mutilla Montezumae Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 634. 9.

Mutilla fulvohirta Cresson, 1865. Ent. Soe. Phila., Proc. 4: 433. 3.

Sphaerophthalma (!) townsendi Cockerell, 1894. Ent. News 5: 199. d.

Sphaerophthalma (!) aspasia Cameron, 1895. Biol. Cent.-Amer., Hym., v. 2, p. 370. d.
Preoce. in Mutilla.

Mutilla aspasioides Dalla Torre, 1897. Cat. Hym., v. 8, p. 12. N. name.

Ephuta californica var. euchroa Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 513. °.

Taxonomy: Mann, 1915. Psyche 22: 178, fig. 1 (gynandromorph).
Biology: Mickel, 1928. U.S. Natl. Mus., Bul. 143: 72. —Krombein, 1958. U.S. Dept. Agr.,
Monog. 2, Sup. 1: 104 (host records).

waco (Blake). Tex.
Mutilla (Sphaeropthalma) Waco Blake, 1871. Amer. Ent. Soc., Trans. 3: 238. 9.

wileyae Mickel. Tex.
Dasymutilla wileyae Mickel, 1928. U.S. Natl. Mus., Bul. 143: 177. 2.

zelaya (Blake). Okla., Tex., N. Mex., Ariz.
Mutilla (Sphaeropthalma) Zelaya Blake, 1871. Amer. Ent. Soc., Trans. 3: 234. 6.

Genus LOMACHAETA Mickel

Lomachaeta Mickel, 1936. Ent. Soc. Amer., Ann. 29: 289.
Type-species: Lomachaeta hicksi Mickel. Orig. desig.
Revision: Mickel, 1940. Pan-Pacific Ent. 16: 127-131.
coloradensis Mickel. Colo. (Boulder Co.).
Lomachaeta coloradensis Mickel, 1936. Ent. Soc. Amer., Ann. 29: 292. 6.
formosula Mickel. Ga. (Clarke Co.); Mexico (Coahuila).
Lomachaeta formosula Mickel, 1940. Pan-Pacifie Ent. 16: 130. d.
hicksi Mickel. Calif.
Lomachaeta hicksi Mickel, 1936. Ent. Soc. Amer., Ann. 29: 289. 2, d.
minutula Mickel. Okla., Tex. (Type from Tenn., Mo. or Ark.).
Lomachaeta minutula Mickel, 1936. Ent. Soc. Amer., Ann. 29: 294. 9.
punctinota Mickel. Ill. (Carbondale).
Lomachaeta punctinota Mickel, 1936. Ent. Soc. Amer., Ann. 29: 293. d.
variegata Mickel. Wash., Calif., Nev., Ariz., N» Mex. Host: Solierella plenoculoides similis
(Brid.), S. blaisdelli (Brid.), nesting in stems of Eriogonum elatum.
Lomachaeta variegata Mickel, 1940. Pan-Pacifie Ent. 16: 128. 2, d.

Biology: Parker, 1962. Pan-Pacific Ent. 38: 116 (host records). —Parker and Bohart, 1966.
Pan-Pacific Ent. 42: 95 (host records).

Genus SMICROMUTILLA Mickel

Smicromutilla Mickel, 1964. Pan-Pacifiec Ent. 40: 108.
Type-species: Smicromutilla powelli Mickel. Orig. desig.
powelli Mickel. Calif. (San Luis Obispo Co.). Host: Diodontus occidentalis Fox?
Smicromutilla powelli Mickel, 1964. Pan-Pacifie Ent. 40: 108, 1 fig. d, 9.

1314. Hymenoptera in America North of Mexico

UNPLACED TAXA OF MUTILLIDAE

Mutilla contracta Say, 1836. Boston Jour. Nat. Hist. 1: 295. ¢. Type destroyed.
Mutilla exulans Fabricius, 1775. Systema Ent., p. 397. 2. Described from Drury coll.; type

probably lost.
Mutilla versicolor Fabricius, 1775. Systema Ent., p. 398. 2. Described from Drury coll.; type

probably lost.
Mutilla vigilans Say, 1836. Boston Jour. Nat. Hist. 1: 296. ¢. Type destroyed.

Family SCOLITDAE

The scoliid wasps are external parasites of larvae of Scarabaeidae in the soil or in debris of
wood-rat nests. Members of some exotic genera parasitize scarabaeid larvae in decaying wood.

Morphology: Betrem, 1972 (1971). Nederland. Ent. Ver., Monog. 6: 13-20, 6 figs. (adults).
SUBFAMILY CAMPSOMERINAE

Revision: Betrem, 1972 (1971). Nederland. Ent. Ver., Monog. 6: 1-326, 6pls. (African species;
erects tribal classification).

TRIBE TRIELIDINI

Taxonomy: Bradley, 1950. Eos, Tomo extraord., pp. 427-438 (primitive character of group).
—Betrem, 1972 (1971). Nederland. Ent. Ver., Monog. 6: 26.

Genus TRIELIS Saussure

Revision: Bradley, 1928. Amer. Ent. Soc., Trans. 54: 195-214, 4 text figs., pl. 22.
Taxonomy: Betrem, 1962. Ent. News 73: 146.

Genus TRIELIS Subgenus TRIELIS Saussure

Trielis Saussure, 1863. Soc. Ent. France, Ann. (4) 3: 18.
Type-species: Elis xantiana Saussure. Monotypic.
_Campsoscolia Betrem, 1933. Stettin. Ent. Ztg. 94: 240.
Type-species: Scolia sexmaculata Fabricius. Orig. desig.

Only the typical subgenus occurs in North America. Our species belong to Species Group In-
terrupta which is found also in the Palaearctic Region.

Taxonomy: Betrem, 1962. Ent. News 73: 146 (type species).
octomaculata hermione (Banks). Austrorip., N. J. to Fla., Ala., Ill.
Trielis hermione Banks, 1912. Canad. Ent. 44: 200. ¢.
octomaculata octomaculata (Say). U. Austr. Zone west of Allegheny Mts. and east of 100th
Meridian.
Scolia vitimaculata Say, 1823. West. Quart. Rptr. 2: 74. @.
Scolia octo-maculata Say, 1825. American Entomology, pl. 29. Emend.
octomaculata texensis (Saussure). U. and L. Sonoran Zones.
Elis texensis Saussure, 1858. Soe. Ent. France, Ann. (8) 6: 224. ¢.
Scolia regina Cresson, 1865. Ent. Soc. Phila., Proc. 4: 447. 2.
Scolia consors Cresson, 1865. Ent. Soc. Phila., Proc. 4: 449. ¢. Preoce.
Scolia flavosignata Cresson, 1865. Ent. Soc. Phila., Proc. 4: 449. 6.
Elis (Trielis) zonaria Cresson, 1868. Amer. Ent. Soc., Trans. 1: 378. N. name.
Elis luwpina Cresson, 1872. Amer. Ent. Soc., Trans. 4: 202. 2.
pollenifera (Viereck). Kans., N. Mex., Ariz.
Elis (Trielis) pollenifera Viereck, 1906. Amer. Ent. Soc., Trans. 32: 190. @.
Elis (Trielis) pollenifera var. a Viereck, 1906. Amer. Ent. Soc., Trans. 32: 191. 2, 6.

Genus CRIOSCOLIA Bradley

Crioscolia Bradley, 1950. Eos, Tomo extraord., p. 431.
Type-species: Campsomeris (Trielis) flammicoma Bradley. Orig. desig.

Superfamily SCOLIOIDEA 1315

Only the typical subgenus occurs in North America.

alcione (Banks). U. Sonoran Zone.
Trielis alcione Banks, 1917. Mus. Compar. Zool., Bul. 61: 112. d.

flammicoma (Bradley). L. Sonoran Zone.
Campsomeris (Trielis) flammicoma Bradley, 1928. Amer. Ent. Soc., Trans. 54: 209. 2, d.

TRIBE CAMPSOMERINI
Taxonomy: Betrem, 1972 (1971). Nederland. Ent. Ver., Monog. 6: 76.

Genus CAMPSOMERIS Guerin

Taxonomy: Bradley, 1957. Amer. Ent. Soc., Trans. 83: 65-77 (subgeneric reclassification
Amer. spp.). —Bradley, 1964. Ent. News 75: 101-108 (added notes on subgenera and spp.).

Genus CAMPSOMERIS Subgenus CAMPSOMERIS Guerin

Campsomeris Guerin, 1838. In Duperrey, Voy. Coquille, Zool., v. 2, p. 247.
Type-species: Scolia atrata Fabricius. Desig. by Bequaert, 1926.
Colpa Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 534.
Type-species: Colpa peregrina Lepeletier. Desig. by Betrem, 1928.

The typical subgenus does not occur in North America.

Genus CAMPSOMERIS Subgenus DIELIS Saussure and Sichel

Elis subg. Dielis Saussure and Sichel, 1864. Cat. Spec. Gen. Scol., p. 14.
Type-species: Scolia radula Fabricius. Desig. by Betrem, 1928.

Revision: Bradley, 1928. Acad. Nat. Sci. Phila., Proc. 80: 313-337, 2 text figs., pl. 26.
Taxonomy: Betrem, 1962. Ent. News 73: 207 (type of Dielis).
SPECIES GROUP PLUMIPES

Taxonomy: Bradley, 1957. Amer. Ent. Soc., Trans. 83: 69.
plumipes confluenta (Say). U. Austr. between Allegheny and Rocky Mts. Ecology: On flowers
of Rubus, Arabis, Melilotus, Solidago, and Cephalanthus. Host: Cotalpa lanigera (L.).
Scolia confluwenta Say, 1823. West. Quart. Rptr. 2: 74. °.

Biology: Kurezewski, 1963. Ent. News 74: 21-24 (host record). —Kurezewski, 1966. Kans.
Ent. Soc., Jour. 39: 156 (host record). —Kurezewski, 1967. Kans. Ent. Soc., Jour. 40:
208-209 (host record).

plumipes fossulana (Fabricius). Austrorip. from Md. to Fla., west to Tex.
Scolia fossulana Fabricius, 1805. Systema Piezatorum, p. 242. °.
Campsomeris szetschwanensis forma D Betrem, 1932. Ent. Ber. 9: 414. ¢.

Taxonomy: Betrem, 1964. Jn Bradley, Spolia. Zool. Mus. Hauniensis 21: 35 (synonymy).

plumipes plumipes (Drury). Carol. Zone from Mass. to Ga., Ky.
Sphea plumipes Drury, 1770. Illus. Nat. Hist., v. 1, p. 104, pl. 44, fig. 5. 9.
Scolia radula Fabricius, 1775. Systema Ent., p. 355. N. name for plumipes. Preoce. in
Campsomeris by Tiphia radula Fabricius, 1775, p. 354.
Scolia quadricincta Klug, 1805. In Weber and Mohr, Beitr. z. Naturk. 1: 37. 2.

Taxonomy: Bradley, 1964. Spolia Zool. Mus. Hauniensis 21: 23 (nomenclature radula).
—Betrem, 1964. Jn Bradley, Spolia Zool. Mus. Hauniensis 21: 34-35 (radula specimens in
Fabricius’ collection). —Bradley, 1967. Jn Bradley and Betrem, Deut. Ent. Ztschr. 15: 331
(lectotype quadricincta).

tolteca (Saussure). Tex., Ariz., Calif., south into Mexico, Haiti.
Elis tolteca Saussure, 1857. Rev. Mag. Zool. (2) 9: 282. 9.
Elis dives Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 410. 2.
Elis 4-cincta Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 411. 6.

1316 Hymenoptera in America North of Mexico

trifasciata nassauensis Bradley. Fla. (Key Vaca); Bahamas.
Campsomeris (Campsomeris) trifasciata nassauensis Bradley, 1928. Acad. Nat. Sci. Phila.,
Proc. 80: 322. 2, 6.
trifasciata trifasciata (Fabricius). Fla. (Miami) and Greater Antilles.
Tiphia trifasciata Fabricius, 1793. Ent. System., v. 2, p. 226. °.
Colpa Alexandri Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 548. d.

Taxonomy: Bradley, 1964. Spolia Zool. Mus. Hauniensis 21: 26 (holotype trifasciata).
SPECIES GROUP PILIPES

Bradley (1964. Ent. News 75: 102) notes that a new subgenus will be described for this species
group.

Taxonomy: Bradley, 1957. Amer. Ent. Soc., Trans. 83: 69-70.
pilipes (Saussure). U. and L. Sonor. Zones.

Elis pilipes Saussure, 1858. Soc. Ent. France, Ann. (8) 6: 246. 9.
Biology: Linsley, 1946. Jour Econ. Ent. 39: 27-28 (pollinating alfalfa; misdet. as plwmipes).

Genus CAMPSOMERIS Subgenus XANTHOCAMPSOMERIS Bradley

Campsomeris subg. Xanthocampsomeris Bradley, 1957. Amer. Ent. Soc., Trans. 83: 70.
Type-species: Tiphia tricincta Fabricius. Orig. desig.

Revision: Rohwer, 1927. Wash. Acad. Sci., Jour. 17: 150-154.
completa completa Rohwer. Ariz. (Chiricahua and Santa Rita Mts.), Tex. (Hidalgo Co.);
Mexico to El Salvador. Another subsp. occurs in Mexico.
Campsomeris (Campsomeris) completa Rohwer, 1927. Wash. Acad. Sci., Jour. 17: 151. @.
fulvohirta (Cresson). Fla. (Miami); Cuba.
Scolia (Elis) fulvohirta Cresson, 1865. Ent. Soe. Phila., Proc. 4: 119. 2, 6.
Campsomeris (Campsomeris) fulvhirta (!) Rohwer, 1927. Wash. Acad. Sci., Jour. 17: 154.
ORO:
limosa (Burmeister). Ariz.; Mexico.
Scolia limosa Burmeister, 1853. Naturf. Gesell. Halle, Abhandl. 1 (4): 28. 2, ¢.
Dielis fulvopilosa Banks, 1912. Canad. Ent. 44: 200. °.

Taxonomy: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 776 (synonymy). —Bradley, 1966. In
Bradley and Betrem, 1966. Beitr. z. Ent. 16: 76 (lectotype limosa).

Genus CAMPSOMERIS Subgenus PYGODASIS Bradley

Campsomeris subg. Pygodasis Bradley, 1957. Amer. Ent. Soc., Trans. 83: 72.
Type-species: Scolia quadrimaculata Fabricius. Orig. desig.

SPECIES GROUP QUADRIMACULATA

quadrimaculata (Fabricius). Largely L. Austr. from Mass. south to Fla., west to Okla. and
Tex.
Vespa maculata Drury, 1773. Illustr. Nat. Hist. 2: 74, pl. 39, fig. 2. 2. Preoce.
Scolia quadrimaculata Fabricius, 1775. Syst. Ent., p. 355. °.
Scolia quatuormaculata Christ, 1791. Naturgesch. Class. Nomencl. Ins., p. 267. N. name.
Scolia quadrinotata Fabricius, 1805. Systema Piezatorum, p. 240. °.
Scolia Abotii Klug, 1810. In Weber and Mohr, Beitr. z. Naturk. 2: 213. ¢.
Colpa Pensylvanica Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 546. d.
Elis quadriguttata (!) Viereck, 1906. Amer. Ent. Soc., Trans. 32: 222.
Campsomeris druryi Cockerell, 1907. Entomologist 40: 50. N. name.

Taxonomy: Bradley, 1964. Spolia Zool. Mus. Hauniensis 21: 21-22 (quadrinotata
nomenclature; notes that it is a variety of quadrimaculata). —Bradley, 1967. In Bradley
and Betrem, Deut. Ent. Ztschr. 15: 324 (holotype abotii).

Superfamily SCOLIOIDEA 1317

SPECIES GROUP EPHIPPIUM

ephippium ephippium (Say). Tex. (Victoria, Seguin); south to northern South America.
Another subsp. occurs in South America.
Scolia ephippium Say, 1837. Boston Jour. Nat. Hist. 1: 333. d.
Scolia Petitii Guerin, 1838. In Duperrey, Voy. Coquille, Zool., v. 2, p. 249.

Taxonomy: Bradley, 1945. Ent. Venez., Bul. 4: 18 (notes on type series pettitii (!)).
—Krombein, 1949. U. S. Natl. Mus., Proe. 100: 56-57, figs. 4-12 (gynandromorph).

Genus MICROMERIELLA Betrem

Micromeris Betrem, 1967. In Bradley and Betrem, Brit. Mus. (Nat. Hist.), Bul., Ent. 20:
294. Preocc.
Type-species: Scolia marginella Klug. Orig. desig.
Micromeriella Betrem, 1972 (1971). Nederland. Ent. Ver., Monog. 6: 116. N. name.

This genus is not a member of the Nearctic fauna.

marginella modesta (Smith). Liberated in N. J., Conn., but not established; Philippines.
Introduced from Hawaii where it had been successfully established from the
Philippines. Host: Anomala orientalis Waterh.
Scolia modesta Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 91. 2, (¢ misdet.).
Scolia manilae Ashmead, 1904. N. Y. Ent. Soc., Jour 12: 8. 9.

Genus CAMPSOMERIELLA Betrem

Campsomeris subg. Campsomeriella Betrem, 1941. Notes d’ent. Chin. 8, fase. 4: 86.
Type-species: Scolia thoracica Fabricius. Orig. desig.

This genus is not a member of the Nearctic fauna.

Genus CAMPSOMERIELLA Subgenus ANNULIMERIS Betrem

Campsomeriella subg. Annulimeris Betrem, 1967. Ent. Ber. 27: 28.
Type-species: Tiphia annulata Fabricius. Orig. desig.
annulata (Fabricius). Liberated in N. J. but not established. Introduced from China and Japan.

Host: Popilla japonica Newm.

Tiphia annulata Fabricius, 1793. Ent. System., v. 2, p. 225. 2.

Campsomeris Servillei Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 501. 2. Preoce.
(secondary homonym).

Biology: Clausen, King and Teranishi, 1927. U. S. Dept. Agr., Bul. 1429: 31-33 (life history).
—Clausen, Jaynes and Gardner, 1933. U. S. Dept. Agr., Tech. Bul. 366: 28-30 (life history).

SUBFAMILY SCOLIINAE
Genus SCOLIA Fabricius

Revision: Bartlett, 1912. Ent. Soc. Amer., Ann. 5: 293-340, 2 pls.
Taxonomy: Hurd, 1952. Calif. Ins. Survey, Bul. 1: 141-152, 2 pls. (Calif. spp.).

Genus SCOLIA Subgenus SCOLIA Fabricius

Scolia Fabricius, 1775. Systema Ent., p. 355.

Type-species: Scolia quadripunctata Fabricius. Desig. by Latreille, 1810.
Lacosi Guerin, 1838. In Duperrey, Voy. Coquille, Zool., v. 2, p. 247.

Type-species: Scolia quadripunctata Fabricius. Desig. by Bequaert, 1926.
Lisoca Costa, 1858. Fauna Napoli Scoliid., p. 8.

Type-species: Scolia quadripunctata Fabricius. Desig. by Krombein, 1951.

The typical subgenus does not occur in North America.

1318 Hymenoptera in America North of Mexico

Genus SCOLIA Subgenus DISCOLIA Saussure

Scolia subg. Discolia Saussure, 1863. Soc. Ent. France, Ann. (4) 3: 18.
Type-species: Scolia nobilitata Fabricius. Desig. by Betrem and Bradley, 1964.
bicincta Fabricius. U. and L. Austr. Zones, east of 100th Meridian.
Scolia bicincta Fabricius, 1775. Systema Ent., p. 356.
Scolia obscura Klug, 1805. In Weber and Mohr., Beitr. z. Naturk. 1: 33.
Scolia undata Klug, 1810. In Weber and Mohr, Beitr. z. Naturk. 2: 212. 9.

Taxonomy: Brimley, 1920. Ent. News 31: 261-262. —Bradley, 1964. Spolia Zool. Mus.

Hauniensis 21: 10 (type series bicincta).

consors Saussure. Colo.; Mexico (Baja California).
Scolia consors Saussure, 1863. Soc. Ent. France, Ann. (4) 3: 18. 6.
Scolia amoena Cresson, 1865. Ent. Soe. Phila., Proce. 4: 447. 6.

dubia dubia Say. U. and L. Austr. Zones, Mass. to Fla., west to Colo. and Ariz. Host: Cotinis

nitida (L.)?

Scolia dubia Say, 1837. Boston Jour. Nat. Hist. 1: 364.
Scolia aulica Burmeister, 1853. Naturf. Gesell. Halle, Abhandl. 1 (4): 33. 9, 6.
Discolia thalia Banks, 1912. Canad. Ent. 44: 199. 6.

Taxonomy: Bradley, 1966. Jn Bradley and Betrem, Beitr. z. Ent. 16: 74 (lectotype awlica).

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 129-133. —Rau, 1932. Brooklyn Ent.
Soc., Bul. 27: 59-62.
dubia haematodes Burmeister. Tex., N. Mex., Ariz., Calif.
Scolia haematodes Burmeister, 1853. Naturf. Gesell. Halle, Abhandl. 1 (4): 33. 2, 6.
Elis Americana Saussure, 1857. Rev. Mag. Zool. (2) 9: 282. ¢.

Taxonomy: Bradley, 1966. Jn Bradley and Betrem, Beitr. z. Ent. 16: 76 (lectotype
haematodes).
guttata Burmeister. Tex.; Mexico.
Scolia guttata Burmeister, 1853. Naturf. Gesell. Halle, Abhandl. 1 (4): 36. 9.
Discolia Hecate Kirby, 1889. Ent. Soc. London, Trans., p. 449. d.

Taxonomy: Bradley, 1966. In Bradley and Betrem, Beitr. z. Ent. 16: 76 (lectotype guttata).
mexicana Saussure. West. Tex., N. Mex., Ariz.; Mexico. Host: Cotinis texana Casey.
Scolia (Lacosi) mexicana Saussure, 1858. Soc. Ent. France, Ann. (8) 6: 219. 2.
Scolia monticola Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 223. 2, ¢.
Scolia nigrescens Bartlett, 1912. Ent. Soc. Amer., Ann. 5: 331.

Taxonomy: Bradley, 1964. Jn Betrem and Bradley, Zool. Meded. 40: 96 (synonymy).

nobilitata nobilitata Fabricius. U. and L. Austr. Zones, Mass. to Fla., west to Colo. and Tex.
Scolia nobilitata Fabricius, 1805. Systema Piezatorum, p. 244.
Scolia tricolor Klug, 1805. In Weber and Mohr, Beitr. z. Naturk. 1: 35.
Scolia maculata Guerin, 1838. In Duperrey, Voy. Coquille, Zool., v. 2, p. 255. 2. Preoce.
Scolia ornata Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 96. N. name.

nobilitata otomita Saussure. Calif., Nev., Ariz.; Mexico. Host: Scarabaeid larvae in debris of

wood-rat nest.

Scolia otomita Saussure, 1858. Soe. Ent. Fance, Ann. (3) 6: 223. 6.
Scolia fulviventris Bartlett, 1912. Ent. Soc. Amer., Ann. 5: 323. 9.

Taxonomy: Bradley, 1964. Jn Betrem and Bradley, Zool. Meded. 40: 96 (subspecifie status).

Biology: Ryckman, 1956. Pan-Pacifie Ent. 32: 180 (host record).

nobilitata tricincta Say. West. Kans. and Tex., Colo., N. Mex., Ariz.
Scolia tricincta Say, 1823. West. Quart. Rptr. 2: 74.? 6.
Scolia Ridingsti Cresson, 1865. Ent. Soc. Phila., Proc. 4: 445. 2.
Scolia inconstans Cresson, 1865. Ent. Soe. Phila., Proc. 4: 446. ¢.
Scolia (Discolia) Lecontei Cresson, 1868. Amer. Ent. Soc., Trans. 1: 376. °.
Scolia (Discolia) flavocostalis Cresson, 1868. Amer. Ent. Soc., Trans. 1: 377. 6.
Scolia Lewisvi (!) Cresson, 1868. Amer. Ent. Soc., Trans. 1: 377. Lapsus for lecontez.
Scolia flavicostalis (!) Banks, 1912. Canad. Ent. 44: 200.

Superfamily SCOLIOIDEA 1319

UNPLACED TAXON OF SCOLIA SUBGENUS DISCOLIA SAUSSURE

bifasciata (Swederus). N. Y.
Sphex (Scolia) bifasciata Swederus, 1787. Svensk. Vetensk.-Akad., Handl. 8: 281. This is

possibly the same as typical nobilitata (F.).
Genus TRISCOLIA Saussure

Scolia subg. Triscolia Saussure, 1863. Soe. Ent. France, Ann. (4) 3: 17.
Type-species: Scolia (Triscolia) badia Saussure. Monotypic.

Taxonomy: Betrem and Bradley, 1964. Zool. Meded. 39: 436-437 (description).
ardens (Smith). Tex., N. Mex., Ariz., Calif., ?0reg., south into Mexico.
Scolia fervida Burmeister, 1853. Naturf. Gesell. Halle, Abhandl. 1 (4): 20. 2, 3. Preoce.
Scolia ardens Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 112. N. name.
Scolia Montezumae Saussure, 1857. Rev. Mag. Zool. (2) 9: 281. 9.

Taxonomy: Bradley, 1966. In Bradley and Betrem, 1966, Beitr. z. Ent. 16: 75-76 (lectotype
fervida).

Family SAPYGIDAE

All of the known records confirm that members of this family are parasitic in the nests of
other aculeate Hymenoptera. The sapygid egg is apparently inserted through the cell closure in
the host nest and usually hatches before the host egg. The newly hatched larvae of Sapyga cen-
trata Say, S. lowisi Krombein and S. pumila Cresson destroy the host egg and then develop on
the pollen stored by the host bee; the larvae of S. confluenta Cresson and of Eusapyga rubripes
proxima (Cresson) feed on the host larva after the latter has spun its cocoon. Development to
the adult stage appears to be concurrent with that of the host. Sapygids parasitic on vernal bees
transform to adults late in the summer, as do the host bees, and overwinter in the cocoons,
emerging in the spring.

Revision: Pate, 1947. Acta Zool. Lilloana (Tucuman) 4: 396-402.

Taxonomy: Pate, 1946. Ent. News 57: 219-221 (list of type-species). —Tobias, 1965. Zool.
Zhur. (Moscow) 44: 706-715, 4 figs. (family classification).

SUBFAMILY FEDTSCHENKIINAE

This subfamily includes only the primitive fossorial genus Fedtschenkia Sauss. It has a
disjunct distribution in xeric Holarctic areas, the five described species being known from Turk-
menia, Uzbekistan, Tadzhikistan, Iran, Lebanon, Israel and western United States. Modern Eu-
ropean workers have considered that Fedtschenkia constitutes a separate family based on the
erroneous assumption that it is a free-living fossorial form, probably parasitic on soil-dwelling
larvae as in the Scoliidae. The single North American species of Fedtschenkia is a parasite of
the ground-nesting eumenid wasp, Pterochilus trichogaster Bohart.

Genus FEDTSCHENKIA Saussure

Fedtschenkia Saussure, 1880. In Fedtschenko, Reise in Turkestan, II, Zool. Th., 13
Hymenoptera: Scoliidae, p. 13.
Type-species: Fedtschenkia grossa Saussure. Monotypic.
Cosilella Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 237.
Type-species: Cosila plutonis Banks. Orig. desig.

Taxonomy: Bradley, 1955. Ent. News 66: 230-233.
anthracina (Ashmead). Wash., Calif., Colo., N. Mex. Ecology: Visits flowers of Chaenactis,
Chorizanthe, Cryptantha, Eriogonum, Eriophyllum, Melilotus and Wislizenia. Host:
Pterocheilus trichogaster Bohart in soil.
Telephoromyia anthracina Ashmead, 1898. Psyche 8: 251. 6.
Plesia (Myzine) nigropilosella Cameron, 1908. Amer. Ent. Soc., Trans. 34: 237. °.
Cosila plutonis Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 237. 9.

Biology: Bohart and Schuster, 1972. Pan-Pacific Ent. 48: 149 (host record).

1320 Hymenoptera in America North of Mexico
SUBFAMILY SAPYGINAE

North American species have been reared only from megachilid bees. However, extralimital
species have been reared from Odynerus sens. lat., Anthophora and Xylocopa as well as from

Megachilidae.
Genus SAPYGA Latreille

Sapyga Latreille, 1796. Precis Caract. Ins., p. 134.
Type-species: Scolia quinquepunctata Fabricius. Desig. by Latreille, 1802.
Hellus Fabricius, 1805. Systema Piezatorum, p. xiii.
Type-species: Sapyga sexpunctata (Fabricius). Desig. by Shuckard, 1837.
aculeata Cresson. Minn., Wyo., Colo., Alta, Oreg., Calif. in Transit. Zone. Host: Hoplitis
producta (Cr.), H. producta gracilis (Mich.), H. fulgida platyura (Ckll.), H. hypocrita
(Ckll.), H. sambuci Titus. Parasite: Sphaeropthalma amphion (Fox).
Sapyga aculeata Cresson, 1865. Ent. Soc. Phila., Proc. 4: 450. 2.
Eusapyga aciculata (!) Hicks, 1934. Univ. Colo. Studies 21: 268.

Biology: Davidson, 1896. Ent. News 7: 218. —Hicks, 1934. Univ. Colo., Studies 21: 268.
—Linsley and Michener, 1942. Pan-Pacifie Ent. 18: 28. —Linsley, 1944. Brooklyn Ent. Soc.,
Bul. 39: 54. —Parker and Bohart, 1966. Pan-Pacific Ent. 42: 96-97. —Parker and Bohart,
1968. Pan-Pacifie Ent. 44: 5.

angustata Cresson. Wyo., Colo., Utah, Ariz., Nev., Wash., Oreg., Calif. in Transit. and U. and

L. Sonor. Zones. Host: Osmia atrocyanea CklIl., O. pikei Ckll., O. lignaria Say.,
Megachile angelarum Ckll.

Sapyga angustata Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. d.

Sapyga moesta Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. d.

Sapyga obscura Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. 9.

Sapyga fulvicornis Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. ¢.

Sapyga maesta (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 192.

Sapyga russellensis Roberts, 1929. Psyche 36: 360. °.

Biology: Linsley and Michener, 1942. Pan-Pacifie Ent. 18: 28. —Linsley, 1944. Brooklyn Ent.
Soc., Bul. 39: 54. —Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 97.
centrata Say. Ont. south to S. C., W. Va., IIll., Tex. Host: Osmia bucephala Cr., O. pumila Cr.,
Hoplitis truncata (Cr.)?
Sapyga centrata Say, 1836. Boston Jour. Nat. Hist. 1: 301.
Sapyga americana Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. 2.
Sapyga pelopaei Ashmead, 1896. Amer. Ent. Soc., Trans. 23: 179. d.

Biology: Krombein, 1952. Ent. Soe. Wash., Proc. 54: 175. —Krombein, 1967. Trap-nesting
wasps and bees, pp. 479-481 (life History). —Medler, 1967. Ent. Soc. Amer., Ann. 60: 342
(host record).

confluenta Cresson. N. H., Colo. Host: Osmia hesperella Ckll., O. lignaria propinqua Cr., O.
cordata Robt.
Sapyga confluenta Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. ¢.
Sapyga emarginata Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. 9.

Biology: Hicks, 1934. Univ. Colo., Studies 21: 267-268.
elegans Cresson. Colo., Nev., Idaho, Calif., Oreg., U. Sonor. and Transit. Zones. Ecology: Visits
flowers of Arctostaphylos nevadensis.
Sapyga elegans Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. d, 2.
Sapyga coloradensis Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. d.
Sapyga truncata Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. d.

Biology: Linsley and Michener, 1942. Pan-Pacifie Ent. 18: 28.

interrupta Roberts. Colo.
Sapyga interrupta Roberts, 1929. Psyche 36: 359. @.

louisi Krombein. N. Y., N. J., Fla., Mich., Tex. Host: Heriades carinata Cr.
Sapyga lowisi Krombein, 1938. Ent. Soc. Amer., Ann. 31: 467. 6, 9.

Biology: Matthews, 1965. Amer. Ent. Inst., Contrib. 1, no. 3: 24-26 (life history).

Superfamily SCOLIOIDEA 1321

maculata Provancher. Que.
Sapyga maculata Provancher, 1882. Nat. Canad. 18: 9. 2.
martinii Smith. Canada, N. H., Colo., Utah, Wyo., Wash.
Sapyga martinii Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 117. d, °.
nevadica Cresson. Tex., Utah, Nev., Idaho, Wash., Oreg., Calif. Host: Dianthidium dubium
dilectum Timb.
Sapyga nevadica Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xxi. d.
Sapyga montana Cresson, 1880. amer. Ent. Soc., Tans. 8: Proc., p. xxi. 9.

Biology: Hurd and Linsley, 1950. N. Y. Ent. Soc., Jour. 58: 247.

pumila Cresson. Nebr., Colo., N. Mex., Utah, Nev., Alta., Calif. Host: Ashmeadiella aridula
CkIl., A. meliloti Ckll., Anthocopa copelandica (Ckll.), Dianthidium consimile (Ashm.),
Megachile rotundata (F.), Osmia sp., Heriades variolosa (Cr.).
Sapyga pumila Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. 2.
Sapyga minor Roberts, 1933. Kans. Ent. Soc., Jour. 6: 96. ¢.

Biology: Hicks, 1934. Univ. Colo., Studies 21: 268. —Linsley, 1944. Brooklyn Ent. Soc., Bul.
39: 54. —Parker and Bohart, 1966. Pan-Pacific Ent. 42: 96-97. —Parker and Bohart, 1968.
Pan-Pacific Ent. 44: 3-4. —Torchio, 1972. Melanderia 10: 1-30, 60 figs. (life history, control).
—Parker, 1975. Pan-Pacifie Ent. 51: 119 (host).

Morphology: Torchio, 1972. Melanderia 10: 8-10, figs. 26-55 (larval instars, pupa).

Genus EUSAPYGA Cresson

Eusapyga Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx.
Type-species: Sapyga rubripes Cresson. Desig. by Ashmead, 1903.

Host records include only the megachilid genus Dianthidiwm.

californica (Cresson). Calif.

Sapyga californica Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. d.
intermedia Roberts. Calif. Host: Dianthidium sp.

Eusapyga intermedia Roberts, 1929. Psyche 36: 361. 2.
nigripes (Cresson). Nev.

Sapyga nigripes Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. ¢.
rubripes carolina Banks. N. C.

Eusapyga carolina Banks, 1912. Canad. Ent. 44: 208. 6.
rubripes proxima (Cresson). Colo., Wyo., Mont. Host: Dianthidium pudicum (Cr.), probably

subsp. decorum Timb.
Sapyga proxima Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. 6.

Biology: Hicks, 1927. Psyche 34: 193.

rubripes rubripes (Cresson). Tex., Colo. Host: Dianthidium pudicum (Cr.), probably subsp.
decorum Timb.
Sapyga rubripes Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. d, 9.

Biology: Hicks, 1934. Univ. Colo., Studies 21: 268.

verticalis (Cresson). Nev., Calif. in Transit. Zone. Host: Dianthidium consimile (Ashm.), D. wu.
ulkei (Cr.).
Sapyga verticalis Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xx. 9.

Biology: Hicks, 1934. Univ. Colo., Studies 21: 268. —Parker and Bohart, 1966. Pan-Pacific
Ent. 42: 96.

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1323

Superfamily FORMICOIDEA

By Davi R. SMITH

Family FORMICIDAE

Ants are social insects that live in colonies in various ecological situations, most commonly in
the soil, rotting wood, and plant cavities. They are practically ubiquitous being extremely prolific
in numbers of individuals even though relatively low in numbers of species. Although most ants
are free-living, some are parasites on other species of ants or live as inquilines in the nests of
other ants. Most ants are omnivorous, but many have a more highly specialized food require-
ment. A number of forms are of concern to man because of their feeding, nesting, or stinging
habits. The species that nest in or near houses or other buildings are sometimes considerably an-
noying to man. Because of the adaptive ability of many species and their habits of nesting in
plants and soil, many forms have been spread throughout the world by commerce. A number of
these tramp species are found in North America.

A colony of ants is usually composed of one or more reproductive females (queens) and wor-
kers. At certain times of the year a nest also contains males and virgin females. There are three
distinct castes of ants, the female, males, and workers. The female is winged but loses her wings
soon after finding a suitable place to begin a new colony; her main purpose is reproduction. The
male is winged and is short-lived; he dies soon after mating. The workers are not winged, and
their main functions are to build the nest, feed the colony, care for the young, and defend the
nest. In some cases, workers are also reproductive forms. Many ants are monomorphic, that is,
the workers are all the same size. Other ants are polymorphic, in which case the workers vary
considerably in size. In polymorphic species, the largest workers are commonly termed soldiers
or maxims and the smaller workers termed minors or minims. The size of a colony of ants, de-
pending on the species and age of the colony, varies and may contain only a few dozen to many
thousands of individuals.

The literature on ants in voluminous. It is impossible to cite every reference on the subject.
The following list contains some of the more significant works on a variety of subjects. Further
references may be found in the literature cited sections of these citations. There have been
many local or state faunal lists and treatments of ants; many of these are given by Smith (1947)
who listed 84 publications by states. They are not repeated here.

I appreciate the cooperation of the following myrmecologists who have reviewed all or parts
of this section: A. C. Cole, University of Tennessee, Knoxville; A. Francoeur, Universite du
Quebec a Chicoutimi; M. R. Smith, Arlington, Virginia; R. R. Snelling, Los Angeles County
Museum, California; and J. F. Watkins II, Baylor University, Waco, Texas. This section is based
in large part on the Formicidae sections of the first Hymenoptera Catalog (1951) and its supple-
ments (1958, 1967) by M. R. Smith.

Revision: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 1-585, 57 pls. (keys to
genera and species of N. Amer-.).

1324 Hymenoptera in America North of Mexico

Taxonomy: Dalla Torre, 1893. Cat. Hym., v. 7, 289 pp. (world cat.). —Emery, 1910. In
Wytsman, Gen. Ins., fase. 102, 34 pp. (Dorylinae, world cat.). —Emery, 1911. In Wytsman,
Gen. Ins., fase. 118, 124 pp. (Ponerinae, world cat.). —Wheeler, 1911. N. Y. Acad. Sci., Ann.
21: 157-175 (type-species). —Emery, 1912. In Wytsman, Gen. Ins., fase. 137, 50 pp.
(Dolichoderinae, world cat.). —Wheeler, 1920. Psyche 27: 46-55 (subfamilies). —Emery,
1921. In Wytsman, Gen. Ins., fase. 174, 379 pp. (Myrmicinae, world cat.). —Wheeler, 1922.
Amer. Mus. Nat. Hist., Bul. 45: 631-710 (keys to world genera and subgenera). —Emery,
1925. In Wytsman, Gen. Ins., fase. 183, 302 pp. (Formicinae, world cat.). —Carpenter, 1930.
Harvard Univ., Mus. Comp. Zool., Bul. 70: 1-66 (fossil ants of N. Amer.). —Donisthorpe,
1943. Ann. and Mag. Nat. Hist. (11) 10: 617-648, 649-688, 721-737 (type-species). —Smith,
1943. Amer. Midland Nat. 30: 273-321 (key to N. Amer. genera based on males). —Smith,
1947. Amer. Midland Nat. 37: 521-647 (key to N. Amer. genera based on workers; list of
publications on ants by states). —Van Pelt, 1948. Fla. Ent. 30: 57-67 (spp. of Alachua Co.,
Fla.). —Brown, 1954. Insectes Sociaux 1: 21-31 (phylogeny and subfamily classification).
—Van Pelt, 1956. Amer. Midland Nat. 56: 358-387 (spp. of Welaka Reserve, Fla.).
—Kannowski, 1956. Amer. Midland Nat. 56: 168-185 (spp. of Ramsey Co., N. Dak.). —Van
Pelt, 1958. Amer. Midland Nat. 59: 1-57 (list of spp. of Welaka Reserve, Fla.). —Wenner,
1959. Amer. Midland Nat. 62: 174-182 (spp. of Bidwell Park, Chico, Calif.). —Wheeler and
Wheeler, 1960. Psyche 67: 87-94 (techniques for study of larvae). —Carter, 1962. Elisha
Mitchell Sci. Soc., Jour. 78: 150-204 (N. C.). —Carter, 1962. Elisha Mitchell Sci. Soc., Jour.
78: 1-18 (N. C. Piedmont). —Wheeler and Wheeler, 1963. Ants of N. Dak., Univ. N. Dak.
Press, Grand Forks, 326 pp. —Gregg, 1963. Ants of Colo., Univ. Colo. Press, Boulder, 792
pp. —Wilson, 1964. Breviora 210: 1-14 (ants of Fla. Keys). —Young and Howell, 1964. Okla.
Agr. Expt. Sta. MP 71 (spp. of Okla.). —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol.
Ser. 7 (3): 1-27 (spp. of Nev. Test Site). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:
1-109 (spp. of Polynesia). —Wilson, Carpenter, and Brown, 1967. Psyche 74: 1-19 (first
Mesozoic ant; a new subfamily). —Warren and Rouse, 1969. Ark. Univ., Agr. Expt. Sta.,
Div. Agr. Bul. 742, 67 pp. (spp. of Ark.). —Ross, Rotramel, and LaBerge, 1971. Ill. Nat.
Hist. Survey, Biol. Notes No. 71, 22 pp. (common and economic ants of IIl.). —Wheeler and
Wheeler, 1972. Ent. Soc. Wash., Proc. 74: 35-45 (subfamilies). —Kempf, 1972. Studia Ent.
15: 3-344 (cat. of Neotropical ants). —Wheeler and Wheeler, 1973. Ants of Deep Canyon,
Philip L. Boyd Deep Canyon Research Center, Univ. Calif., Riverside, 159 pp. —Brown,
1973. In Meggers, et al., Tropical forest ecosystems in Africa and S. Amer., Smithsn. Inst.
Press, Wash., D. C., pp. 161-185 (list of world generic and subgenerie names; suggested
synonymy). —Wheeler and Wheeler, 1976. Ent. Soc. Wash., Mem. 7, 108 pp. (ant larvae:
review and synthesis).

Biology: McCook, 1882. The honey ants of the Garden of the Gods, and the occident ants of
the American plains, Philadelphia, 188 pp. —Buckingham, 1911. Amer. Acad. Arts and Sci.,
Proc. 46: 425-507 (division of labor). —Wheeler, 1910. Ants, their structure, development,
and behavior, Columbia Univ. Press, N. Y., 663 pp. (reprinted 1926, 1960). —Wheeler, 1928.
The social insects, their origin and evolution, Harcourt, Brace and Co., N. Y., 378 pp.
—Forel, 1928. The social world of the ants, G. P. Putnam’s Sons, Ltd., London and N. Y.,
v. I, 551 pp., v. II, 445 pp. —Jones, 1929. Colo. Agr. Col., Expt. Sta. Bul. 341, 96 pp.
(relation to aphids). —Wheeler, 1936. Amer. Acad. Arts and Sci., Proc. 71: 159-243
(ecological relations to termites). —Haskins and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37:
97-162 (sociological an physiological features). —Haskins, 1939. Of ants and men,
Prentice-Hall Inc., N. Y., 244 pp. —Lafleur, 1941. N. Y. Ent. Soc., Jour. 49: 227-231 (civil
disturbances in ant communities). —Lafleur, 1941. N. Y. Ent. Soc., Jour. 49: 199-204
(communal disaffection). —Lafleur, 1942. Sci. Monthly 65: 467-471 (behavior in the face of
obstacles). —Smith and Weiss, 1942. U. S. Dept. Agr., Tech. Bul. 798, 44 pp. (relation to
azalea flower spot). —Wheeler, 1942. Harvard Univ., Mus. Comp. Zool., Bul. 40: 1-252
(Neotropical ant-plants and their ants). —MacGregor, 1948. Behaviour 1: 267-296 (odor as
a basis for orientated movement). —Brackbill, 1948. Auk 65: 66-77 (anting by birds).
—Nixon, 1951. The association of ants with aphids and coccids, Commonwealth Inst. Ent.
(London). 35 pp. — Valentini, 1951. Ann. Sci. Nat. (Zool.) 11: 249-276 (adaptation of larvae).
—F landers, 1951. Canad. Ent. 83: 93-98 (role of ants in biological control of homopterous
insects). —Flanders, 1952. Jour. Econ. Ent. 45: 38-39 (ovisorption as the mechanism

Superfamily FORMICOIDEA 1325

causing worker development). —Flanders, 1953. Sci. Monthly 76: 142-148 (caste
determination). —Talbot, 1953. Mich. Univ., Contrib. Lab. Vertebrate Biol. 63, 13 pp.
(fauna and populations). —Wilson, 1953. Psyche 60: 15-20 (caste determination). —Simeone,
1954. State Univ. N. Y., Syracuse, Col. Forestry Bul. 34, 19 pp. (carpenter ants and their
control). —Chapman, 1954. Pan-Pacifie Ent. 30: 93-102 (swarming of ants on mountain
summits). —Gosswald, 1955. Rev. Suisse de Zool. 62: 372-386 (caste determination).
—O’Rourke, 1956. Insectes Sociaux 3: 107-118 (medical and veterinary importance).
—Wilson and Eisner, 1957. Insectes Sociaux 4: 157-166 (liquid transmission of food).
—Talbot, 1957. Insectes Sociaux 4: 375-384 (populations). —Brian, 1957. Ann.Rev. Ent. 2:
107-120 (caste determination). —Weber, 1958 (1956). Tenth Internat. Cong. Ent., Proc. 2:
459-473 (evolution of ants and feeding habits). —Eisner and Wilson, 1958 (1956). Tenth
Internat. Cong. Ent., Proc. 2: 509-513 (food transmission). —Kannowski, 1959. Insectes
Sociaux 6: 115-162 (flight activities and colony founding of bog ants in Mich.). —Wilson,
1959. Anat. Rec. 134: 653 (pheromones in organization of ant societies). —Bartlett, 1961.
Ent. Soc. Amer., Ann. 54: 543-551 (influence of ants on parasites, predators, and scale
insects). —Downey, 1962. Ent. News 73: 57-66 (association with larvae of a lycaenid
butterfly). —Ayre, 1962. N. Y. Ent. Soc., Jour. 70: 159-167 (use of Lincoln Index for
estimating size of colonies). —Reid, 1962. Ga. Agr. Expt. Sta. and Univ. Ga. Handbook, 71
pp. (as intermediate hosts of chicken and turkey tapeworms). —Van Pelt, 1963. Amer.
Midland Nat. 69: 205-223 (distribution in south. Blue Ridge Mtns.). —Way, 1963. Ann. Rev.
Ent. 8: 307-344 (mutualism between ants and honeydew producing Homoptera). — Kincaid,
1963. Amer. Micros. Soc., Trans. 82: 101-105 (pollination of plants). —Wilson, 1963. Ann.
Rev. Ent. 8: 345-368 (social biology). —Wilson, 1963. Evolution 17: 249-253 (social
modification). —Ayre, 1963. Canad. Ent. 95: 712-715 (feeding habits). —Wilson and
Bossert, 1963. Recent Progress 19: 673-716 (chemical communication). —Orlog, 1964 (1963).
Ent. Expt. and Appl. 6: 95-106 (plant virus transmission). —Talbot, 1965. Insectes Sociaux
12: 19-47 (populations in a low field). —Brian, 1965. Social insect populations, Academic
Press, London and N. Y., 135 pp. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326, 105 pp.
(house-infesting ants of east. U. S.). —Janzen, 1966. Evolution 20: 249-275 (coevolution of
mutualism between ants and acacias). —Beck, Allred, and Despain, 1967. Great Basin Nat.
27: 67-78 (predaceous-scavenger ants in Utah). —Sudd, 1967. An introduction to the
behavior of ants, St. Martin’s Press, N. Y., 200 pp. —Markin, 1968. Jour. Econ. Ent. 61:
1744-1745 (handling techniques for large quantities of ants). —Wilson, 1971. The insect
societies, Belknap Press of Harvard Univ., 548 pp. —Lettendre, et al., 1971. Nat. Canad.
98: 591-606 (spp. from St. Hippolyte, Que.). —Payne and Mason, 1971. Ent. Soc. Wash.,
Proc. 73: 135-136, 138 (ants associated with pig carrion). —Jennings, 1971. Ent. Soc. Amer.,
Ann. 64: 384 (ants preying on jack-pine budworm larvae). —Finnegan, 1971. Canad. Ent.
103: 1489-1493 (indigenous ants as limiting agents of forest pests in Que.). —Bhatkar and
Whitcomb, 1970. Fla. Ent. 53: 229-232 (artificial diet for rearing various spp.). —Nielsson,
et al., 1971. Fla. Ent. 54: 245-248 (ants associated with aphids in Fla.). —Whitcomb, e¢ al.,
1972. Fla. Ent. 55: 129-142 (ants of Fla. soybean fields). —Gregg, 1972. Canad. Ent. 104:
1073-1091 (northward distribution of ants in N. Amer.). —Hickman, 1974. Science 184:
1290-1292 (pollination by ants: a low-energy system). —Lettendre and Pilon, 1973. Nat.
Canad. 100: 195-235 (ant fauna of Laurentide, Que.). —Finnegan, 1974. Entomophaga 7:
53-59 (ants as predators of forest pests). —Gurney, 1975. Insect World Digest 2 (5): 19-25
(stinging ants).

Morphology: Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 367-446 (thorax of wingless and
short-winged Hymenoptera). —Brown and Nutting, 1950. Amer. Ent. Soc., Trans. 75:
113-132 (wing venation). —Wilson, 1953. Quart. rev. Biol. 28: 136-156 (polymorphism).
—Wilson, 1954. Insectes Sociaux 1: 75-80 (polymorphism). —Eisner, 1957. Harvard Univ.,
Mus. Comp. Zool., Bul. 116: 439-490 (studies of the proventriculus). —Glockner, 1957.
Insectes Sociaux 4: 83-90 (effect of hormones on metamorphosis). —Eisner and Brown,
1958 (1956). Tenth Internat. Cong. Ent., Proc. 2: 503-508 (evolution and social significance
of the proventriculus). —Nachtweg, 1961. Insectes Sociaux 8: 369-381. —Tulloch, Shapiro,
and Hershenov, 1962. Brooklyn Ent. Soc., Bul. 77: 91-101 (ultrastructure of metasternal
glands). —Roth and Eisner, 1962. Ann. Rev. Ent. 7: 107-136 (chemical defenses). — Wilson,
1963. Sci. Amer. 208: 100-106 (pheromones). —Nachtweg, 1963. Insectes Sociaux 10: 43-57

1326 Hymenoptera in America North of Mexico

(sound organs). —Nachtweg, 1964 (1963) Insectes Sociaux 10: 359-378 (acoustics).
—Etterschank and Brown, 1964. Ent. Monthly Mag. 100: 5-7 (Malphigian tubules as
meristic characters). —Williams and Williams, 1964. Soc. Expt. Biol. Med. 116: 161-163
(toxicity studies of ant venom). —Cavil and Robertson, 1965. Science 149: 1337-1345 (ant
venoms, attractants and repellants). —Law, Wilson, and McCloskey, 1965. Science 149:
544-546 (biochemical polymorphism). —Brown, 1968. Amer. Nat. 102: 188-191 (function of
metapleural glands). —Gotwald, 1969. Cornell Univ., Agr. Expt. Sta. Mem. 408, 150 pp.
(mouthparts). —Hermann, 1969. Ga. Ent. Soc., Jour. 4: 123-141 (poison apparatus). —Blum,
1969. Ann. Rev. Ent. 14: 57-80 (alarm pheromones).

SUBFAMILY DORYLINAE

Members of this subfamily are known as army ants and are sometimes referred to as legiona-
ry ants in the New World and driver ants in the Old World. They are predaceous and are known
for their foraging expeditions the size of which are sometimes exaggerated. Army ants exhibit a
number of morphological and biological pecularities not common to most ants such as wasplike
males, wingless termitelike females, blind workers, and their raiding and emigrating behavior.
Rettenmeyer (1963) outlined the following traits in which they differ from other ants: (1) They
feed almost exclusively on animal prey which is collected by large groups of raiding workers; (2)
their raiding columns usually connect to the nest by at least one continuous column; (3) the en-
tire colony periodically and frequently emigrates to new nest sites; (4) emigrations are largely
dependent on the size, caste, age, and range of ages of the brood (or broods); and (5) the colonies
are founded by division of an entire colony into two (or possibly several) daughter colonies.
Other ants may possess some of these traits, but not all of them.

Much of the biological work on army ants has been done in Central America on the terrestrial
species of Eciton which bivouac in large clusters above the ground and whose colonies may
number up to a million individuals. Most of the army ants, however, are subterranean in habit,
though the raiding columns of some may appear above ground. Raiding may be in columns only
several ants wide or in swarms of a fan-shaped pattern. Most of the prey is other Arthropods,
only occasionally vertebrates. All species have nomadic and statary activity cycles where the en-
tire colony moves from one area to another, a unique behavior studied by Schneirla and Retten-
meyer in the papers listed below.

Borgmeier’s revision of 1955 is the most definitive taxonomic work on this subfamily for the
New World. He recognized 137 species in 5 genera and 2 tribes. Only the tribe Ecitonini is found
in the United States; the other tribe, Cheliomyrmicini, includes a single genus of several species
found from Mexico to Brazil. In the United States, most species are found in the Southwest,
though several reach the Atlantic coast and range north to Iowa, Ohio, and Virginia. Most of the
taxonomy is based on males and workers, and for some species only one caste is known. Further
study and association of castes may result in some synonymy, especially in Netvamyrmex.

Revision: Smith, 1942. Amer. Midland Nat. 27: 537-590 (U. S.). —Borgmeier, 1955. Studia
Ent. 3: 1-716 (New World). —Watkins, 1976. The identification and distribution of New
World army ants, Baylor Univ. Press 102 pp. (keys to genera and spp. of workers and
males; distribution maps).

Taxonomy: Wheeler, 1943. Ent. Soc. Amer., Ann. 36: 319-332 (larvae). —Borgmeier, 1953.
Studia Ent. 2: 1-51. —Borgmeier, 1958. Studia Ent. 1(n. s.): 197-208 (Ecitonini;
identification of females to genus). —Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc.
66: 129-137 (larvae). —Watkins 1972. Kans. Ent. Soc., Jour. 45: 347-372 (U.S.
Neivamyrmesx).

Biology: Wheeler, 1900. Amer. Nat. 34: 563-574. —Wheeler, 1910. Ants, pp. 246-266.
—Schneirla, 1933. Jour. Comp. Psychology 15: 267-299 (in Panama). —Schneirla, 1934. Nat.
Acad. Sci., Proc. 20: 316-321 (raiding and other phenomena). —Schneirla, 1938. Jour. Comp.
Psychology 25: 51-90 (theory of army ant behavior). —Schneirla, 1940. Jour. Comp.
Psychology 29: 401-460 (mass organization in the swarm-raiders). —Schneirla, 1944. N. Y.
Ent. Soc., Jour. 52: 153-192 (reproductive functions of queen as pace-makers of group
behavior). —Schneirla 1944. Amer. Phil. Soc. Proc. 87: 438-457 (nomadism in Eciton
burchelli (Westwood)). —Schneirla, 1945. Biol. Bul. 88: 166-193 (nomad-statary relations in
swarmers; migration). —Schneirla, 1947. Amer. Mus. Novitates 1336: 1-20 (life and

Superfamily FORMICOIDEA 1327

behavior under dry season conditions with reference to reproductive functions).
—Schneirla, 1948. Zoologica (N. Y.) 33: 89-112 (life and behavior under dry season
conditions, appearance and date of males). —Schneirla, 1948. Sci. Amer. 178: 16-23
(general). —Schneirla, 1949. Amer. Mus. Nat. Hist., Bul. 94: 1-82 (life and behavior under
dry season conditions, course of reproduction and colony behavior). —Schneirla, 1950.
(Mex.) Inst. de Biol., An. 20: 371-384 (environmental adaptations). —Schneirla and Brown,
1950. Amer. Mus. Nat. Hist., Bul. 95: 269-233 (life and behavior under dry season
conditions, cyclic processes in behavioral and reproductive functions). —Schneirla and
Brown, 1952. Zoologica (N. Y. ) 37: 5-32 (sexual broods and production of young queens).
—Schneirla, 1953. Insectes Sociaux 1: 29-41 (army ant queen). —Schneirla, Brown and
Brown, 1954. Ecol. Monog. 24: 269-296. (bivouac or temporary nest as adaptive factor in
terrestrial species). —Tafuri, 1955. N. Y. Ent. Soc., Jour. 63: 21-41 (growth and
polymorphism of larva of Eciton hamatum (F.)). —Schneirla, 1956. Insectes Sociaux 3:
49-69 (colony division and related processes). —Schneirla, 1956. Smithsn. Inst. Ann. Rpt.,
1955 pp. 379-406 (army ants, general). —Schneirla, 1957. Insectes Sociaux 4: 259-298
(comparison of functional patterns in army ants). —Schneirla, 1957. Amer. Phil. Soc., Proc.
101: 106-133 (theoretical consideration of cyclic processes). —Schneirla, 1958. Insectes
Sociaux 5: 215-255 (behavior and biology of Neivamyrmesx nigrescens (Cresson) and N.
opacithorax (Emery)). —Brown, 1960. Psyche 66: 25-27 (alarm and attack behavior).
—Schneirla, 1960. Smithsn. Treasury of Science. Simon and Schuster, N. Y., pp. 664-696
(army ants, general). —Schneirla, 1961. Ztschr. f. Tierpsychol. 18: 1-32 (sexual broods and
colony division of Neivamyrmex nigrescens (Cresson)). —Rettenmeyer, 1961. Kans. Univ.,
Sci. Bul. 42: 993-1066 (biology and taxonomy of flies over swarms of army ants).
—Rettenmeyer, 1963. Univ. Kans., Sci. Bul. 44: 287-465 (behavioral studies of army ants in
Canal Zone and Kansas). —Schneirla, 1963. Animal Behavior 11: 583-595 (springtime
resurgence of cyclic function, Arizona species). —Rettenmeyer, 1960. XI. Internat. Kong.
fur Ent., Wien. 1960, 1: 610-612 (mites associated with army ants). —Rettenmeyer, 1962.
Kans. Ent. Soc., Jour. 35: 377-384 (millipedes associated with army ants). —Rettenmeyer,
1963. Ent. Soc. Amer., Ann. 56: 170-174 (Thysanura associated with army ants). —Watkins,
1964. Kans. Ent. Soc., Jour. 37: 22-28 (trail following). —Watkins and Cole, 1966. Tex. Jour.
Sei. 18: 254-265 (attraction of workers to secretion of queens). —Akre and Rettenmeyer,
1966. Kans. Ent. Soc., Jour. 39: 745-782 (Staphylinidae associated with army ants).
—Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail following and
trail preference). —Akre and Rettenmeyer, 1968. Kans. Ent. Soc., Jour. 41: 165-174 (trail
following by guests of army ants). —Akre, 1968. Pan-Pacific Ent. 44: 87-101 (Histeridae
associated with army ants). —Rettenmeyer and Akre, 1968. Ent. Soc. Amer., Ann. 61:
1317-1326 (ectosymbiosis between phorid flies and army ants.). —Kannowski, 1969.
Internat. Union for study of social insects, VI Cong., Proc. 6: 77-83 (daily and seasonal
periodicities in nuptial flights). —Torgerson and Akre, 1970. Melanderia 5: 1-28
(persistence of army ant chemical trails and their significance in Ecitonine-Ecitophile
association). —Torgerson and Akre, 1970. Kans. Ent. Soc., Jour. 43: 395-404 (interspecific
responses to trail and alarm pheromones). —Schneirla, 1971. Army Ants: A Study in Social
Organization. W. H. Freeman, San Francisco. 349 pp. —Topoff, 1971. Amer. Nat. 105:
529-548 (polymorphism related to division of labor and colony cyclic behavior).

Morphology: Hagen, 1954. Amer. Mus. Novitates 1663: 1-12 (anatomy of queen of Eciton).
—Hagen, 1954. Amer. Mus. Novitates 1664: 1-17 (Reproductive system of queen of
Eciton). —Hagen, 1954. Amer. Mus. Novitates 1665: 1-20 (oocyte cycle). —Lappano, 1958.
Insectes Sociaux 5: 31-66 (morphological study of larval development in all worker broods
of Eciton burchelli (Westwood). —Whelden, 1963. N. Y. Ent. Soc., Jour. 71: 158-178
(reproductive system of worker and female of Eciton hamatum(F.) and Eciton burchelli
(Westwood)). —Whelden, 1963. N. Y. Ent. Soc., Jour. 71: 246-261 (antennae and legs of E.
hamatum and E. burchelli).

TRIBE ECITONINI
Genus LABIDUS Jurine

Labidus Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 282.
Type-species: Labidus latreillii Jurine. Monotypic.

1328 Hymenoptera in America North of Mexico

Nycteresia Roger, 1861. Berlin. Ent. Ztschr. 5: 21.
Type-species: Formica coeca Latreille. Monotypic.
Pseudodicthadia Andre, 1885. Spec. Hym. Eur. Alg. 2: 838.
Type-species: Pseudodicthadia incerta Andre. Monotypic.

A neotropical genus of 8 species, one of which reaches the United States.
Revision: Borgmeier, 1955. Studia Ent. 3: 80-134.
Taxonomy: Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc. 66: 134 (larvae).

Biology: Rettenmeyer, 1963. Kans. Univ., Sci. Bul. 44: 403-424 (behavioral studies in Canal
Zone).
coecus (Latreille). S. Ark., La., Okla., Tex. s. to Argentina. Ecology: The large colonies, usually

with many thousands of individuals, are found in more or less temporary nests in
decayed logs and stumps or in ground beneath objects. They are subterranean and
nocturnal. Food consists of other arthropods, small mammals, birds, and nuts. Workers
are highly predaceous and are known to feed on injurious insects such as the immature
stages of the serewworm (Cochliomyia hominivorax (Coquerel)) and the secondary
serewworm (C. macellaria (F.)).

Formica omnivora Olivier, 1791. Encycl. Meth. Hist. Nat. 6: 496. ¥. Preoce. by Linnaeus,
1758.

Formica coeca Latreille, 1802. Hist. Nat. Fourmis, v. 9, p. 270. 3.

Labidus latreillii Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 283. ¢.

Labidus jurini Shuckard, 1840. Ann. Nat. Hist. 5: 198. 3.

Labidus servillei Westwood, 1842. Arcana Ent., v. 1, p. 75. d.

Mutilla (Labidus) fulvescens Blanchard, 1849. In Cuvier, Regne Animale, ed. 3, v. 2, pl.
118, fig. 2.

Labidus saji (!) Haldeman, 1852. In Stanbury, Exped. Great Salt Lake, p. 367. do.

Labidus atriceps Smith, 1859. Cat. Hym. Brit. Mus., v. 7, p. 5. 3.

Labidus pilosus Smith, 1859. Cat. Hym. Brit. Mus., v. 7, p. 7. o.

Labidus panzeri Smith, 1859. Cat. Hym. Brit. Mus., v. 7, p. 72. 3.

Eciton vastator Smith, 1860. Jour. Ent., London 1: 71. 3.

Eciton erratica Smith, 1860. Jour. Ent., London 1: 71. %.

Myrmica rubra Buckley, 1867. Ent. Soc. Phila., Proc. 6: 335. %. Preoce. in Myrmica by
Linnaeus, 1758.

Pseudodicthadia incerta Andre, 1885. Spec. Hym. Eur. Alg. 2: 838. ?.

Eciton smithii Dalla Torre, 1892. Cat. Hym., v. 7, p. 6. N. name for L. pilosus Smith.

Eciton coecum var. biloba Emery, 1901. Soc. Ent. Belg., Ann. 45: 51. 3.

Eciton nigrita Emery, 1901. Soc. Ent. Belg., Ann. 45: 52. d.

Eciton coecum var. kulowi Forel, 1901. Mitt. Nat. Mus. Hamburg 18: 47. 3.

Eciton selysi Forel, 1904. Soc. Ent. Belg., Ann. 48: 169. 3.

Eciton grassator Forel, 1911. Deut. Ent. Ztschr., p. 288. ¥.

Eciton (Labidus) coecum servillei var. hostilis Santschi, 1920. Soc. Ent. France, Ann. 88:
368. 3.

Eciton (Labidus) coecum var. opacifrons Wheeler, 1921. Amer. Acad. Arts and Sci., Proc.
56: 310. &.

Eciton (Labidus) coecum var. elsbethae Forel, 1922. Rev. Suisse Zool. 30: 91. d.

Eciton (Labidus) serpentis Weber, 1938. Ent. Soc. Amer., Ann. 31: 209. %.

Taxonomy: Weber, 1941. Amer. Midland Nat. 26: 238 (queen). —Wheeler, 1943. Ent. Soc.
Amer., Ann. 36: 332 (larva). —Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc. 66: 135
(larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 408-409. —Lindquist, 1942. Jour.
Econ. Ent. 35: 850 (as predators of serewworms). —Hess, 1958. Field and Lab. 26: 35-37.
—Kempf, 1961. Studia Ent. 4(n.s.): 551-552 (as a cave ant). —Rettenmeyer, 1963. Kans.
Univ., Sci. Bul. 44: 418-424. —Smith, 1965. U. S. Dept. Agr. Tech. Bul. 1326, pp. 17-18.
—Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-265 (attraction of workers to secretion of
queens). —Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail
following and trail preference).

Superfamily FORMICOIDEA 1329

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 390 (maxillary and labial palps).
—Gotwald, 1969. N. Y. Agr. Expt. Sta. (Cornell Univ.) Mem. 408, p. 127 (mouthparts).

Genus NOMAMYRMEX Borgmeier

Eciton subg. Nomamyrmex Borgmeier, 1936. Inst. Biol. Veg. Arq. 3: 55.
Type-species: Eciton crassicorne Smith. Orig. desig.

A genus of two species, a subspecies of only one reaching the United States. They are subter-
ranean though carrying on terrestrial raids. Raiding is in columns of several ants wide and a few
meters long and is sometimes done during the daylight hours.

Revision: Borgmeier, 1955. Studia Ent. 3: 135-161.

Taxonomy: Borgmeier, 1953. Studia Ent. 2: 4. —Borgmeier, 1958. Studia Ent. 1 (n. s.):
201-203.

Biology: Rettenmeyer, 1963. Kans. Univ. Sci. Bul. 44: 424-432 (behavioral studies in Canal
Zone; esenbeckii crassicornis (Smith)).

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 390 (maxillary and labial palps).
esenbeckii wilsoni (Santschi). Extreme s. Tex. to Costa Rica. N. esenbeckii esenbeckwi
(Westwood) is South American. Literature references for esenbeckii (Westwood) and
crassicornis (Smith) from Texas pertain to wilsoni.
Eciton (Labidus) Esenbecki Wilsoni Santschi, 1919. Soe. Ent. France, Ann. 88: 366. 3.
Eciton (Holopone) crassicornis mordax Santschi, 1928. Deut. Ent. Ztschr., p. 415. &.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 409 (male).

Genus NEIVAMYRMEX Borgmeier

Eciton subg. Acamatus Emery, 1894. Soc. Ent. Ital., Bol. 26: 181. Preoee. by Schoenherr,
1833.
Type-species: Eciton (Acamatus) schmitti Emery. Desig. by Wheeler, 1911.
Eciton subg. Neivamyrmex Borgmeier, 1940. Rev. de Ent. 11: 606. N. name for Acamatus
Emery.
Woitkowskia Enzmann, 1952. Iowa Acad. Sci., Proc. 59: 443.
Type-species: Woitkowskia connectens Enzmann. Orig. desig.

About 115 species are known for this New World genus, but only 23 are found north of Mex-
ico. Most species are found in the southwestern states with a few ranging east to the Atlantic
coast and as far north as Virginia, Ohio, Illinois, and Iowa. The species are hypogaeic and are
found in the soil under objects, though some have been reported from rotten logs and stumps.
Some may carry on their foraging and emigrating activities during daylight, but most are ap-
parently nocturnal in their activities. Most colonies have one functional queen, and new colonies
are produced by fission. The cyclic pattern of nomadic and statary phases is similar to the tropi-
cal species, but in most Nearctic species the activity stops in the autumn and resumes again in
the spring. Because many of the species listed below were described from a single caste, future
study and caste association may reveal that fewer species actually exist.

Revision: Smith, 1942. Amer. Midland Nat. 27: 537-590 (U. S. species). —Borgmeier, 1955.
Studia Ent. 3: 277-651 (New World species).

Taxonomy: Borgmeier, 1950. Rev. de Ent. 21: 624. —Watkins, 1971. Kans. Ent. Soc., Jour. 44:
93-103 (key to major workers and queens of U. S.). —Watkins, 1972. Kans. Ent. Soc., Jour.
45: 347-372 (keys for each caste and distribution of U. S. species).

Biology: Schneirla, 1958. Insectes Sociaux 5: 215-255 (last part of functional season, s.e.
Ariz.). —Schneirla, 1961. Ztschr. f. Tierpsychologie 18: 1-32 (sexual broods and colony
division). —Schneirla, 1963. Animal Behaviour 11: 583-595 (spring resurgence of cyclic
function, s.e. Ariz.). —Rettenmeyer, 1963. Kans. Univ., Sci. Bul. 44: 433-452 (behavioral
studies). —Watkins, 1964. Kans. Ent. Soc., Jour. 37: 22-28 (trail following). —Watkins and
Cole, 1966. Tex. Jour. Sci. 18: 254-265 (attraction of workers to secretions of queens).
—Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail following and

1330 Hymenoptera in America North of Mexico

trail preference). —Plsek, Kroll, and Watkins, 1969. Kans. Ent. Soc., Jour. 42: 452-456
(association with carabids).

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 392-393 (maxillary and labial palps).

agilis Borgmeier. S. Ariz.; Mexico. Only the worker is known.

Neivamyrmesx agilis Borgmeier, 1953. Studia Ent. 2: 45. 8.
andrei (Emery). N. Mex., Ariz.; Mexico. Only the male is known.

Eciton andrei Emery, 1901. Soc. Ent. Ital., Bol. 33: 53. ¢.

Eciton (Acamatus) oslari Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 415. d.
baylori Watkins. Tex. (Waco).

Neivamyrmex baylori Watkins, 1973. Kans. Ent. Soc., Jour. 46: 430-433. 3.
californicus (Mayr). Utah, Nev., Calif. Only the worker and queen are known.

Eciton californicum Mayr, 1870. Zool.-Bot. Gesell. Wien., Verh. 20: 969. %.

Eciton (Acamatus) californicum var. obscura Forel, 1914. Soc. Vaud. Sci. Nat., Bul. 50:

265. ¥.

Taxonomy: Watkins, 1972. Kans. Ent. Soc., Jour. 45: 363-366 (queen, distribution).

Biology: Mallis, 1938. Sci. Monthly 47: 220-226. —Mallis, 1941. South. Calif. Acad. Sci., Bul.
40: 62.
carolinensis (Emery). Va., N. C., S. C., Ga., Fla., Ohio, Tenn., Ala., Miss., La., Nebr., Kans., N.
Mex., Ariz. All castes are known.
Eciton (Acamatus) carolinense Emery, 1894. Soc. Ent. Ital., Bol. 26: 184. 8.

Biology: Smith, 1928. Ent. News 39: 245. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 278, 304.
—Watkins, 1964. Kans. Ent. Soc., Jour. 37: 22-28 (trail following). —Watkins and
Rettenmeyer, 1967. Psyche 74: 228-233 (effect of queen on longevity of workers).

fallax Borgmeier. La., Kans., Tex., N. Mex., Ariz.; Mexico; Guatemala. Only the worker is
known. Literature references for commutatum (Emery) for N. Amer. north of Mexico
should be referred to fallax.
Neivamyrmex fallax Borgmeier, 1953. Studia Ent. 2: 48. 8.
fuscipennis (Wheeler). Kans., e. Tex. Only the male is known.
Acamatus fuscipennis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 417. d. Misdet. as
spoliator Forel, but fuscipennis validated by pl. 26, fig. 12.

Taxonomy: Watkins, 1975. Southwest. Nat. 20: 85-90 (relationship to N. macropterus
Borgmeier; lectotype).
harrisii (Haldeman). Okla., Tex., N. Mex., Ariz.; Mexico. All castes are known. Records citing
this species from Utah are probably incorrect.
Labidus harrisii Haldeman, 1852. In Stanbury, Exped. Great Salt Lake, p. 367. 3.
Eciton wheeleri Emery, 1901. Soc. Ent. Ital., Bol. 33: 65. &.
Eciton (Acamatus) wheeleri dubia Creighton, 1932. Psyche 39: 75. %, 2.

Taxonomy: Watkins, 1968. Amer. Midland Nat. 80: 273-275 (association of castes).
—Borgmeier, 1958. Studia Ent. 1: 206-207.

Biology: Watkins and Cole, 1966. Tex. Jour. Sci. 12: 254-265 (attraction of workers to
secretion of queens). —Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40:
146-151 (trail following and trail preference). —Plsek, Kroll, and Watkins, 1969. Kans. Ent.
Soc., Jour. 42: 452-456 (carabids in raiding columns).

Morphology: Forbes and Do-Van-Quy, 1965. N. Y. Ent. Soc., Jour. 73: 95-111 (male
reproductive system).

leonardi (Wheeler). Okla., Tex., Calif.; Mexico. Only the worker is known.
Eciton (Acamatus) leonardi Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 392. 8.
Eciton (Acamatus) peninsulare Mann, 1926. Psyche 33: 98. 8.

Taxonomy: Borgmeier, 1953. Studia Ent. 2: 10. —Watkins, 1971. Kans. Ent. Soc., Jour. 44:
101-103 (worker; biological notes).

Superfamily FORMICOIDEA 1331

Biology: Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-265 (=pausxillus of authors, not *
Wheeler; attraction of workers to secretion of queens). —Watkins, Cole, and Baldridge,
1967. Kans. Ent. Soc., Jour. 40: 146-151 (~pausxillus of authors, not Wheeler; trail
following and trail preference).

macropterus Borgmeier. W. Tex., N. Mex., Ariz.; Mexico. only the male is known.
Neivamyrmex macropterus Borgmeier, 1953. Studia Ent. 2: 40. d.

Taxonomy: Watkins, 1975. Southwest. Nat. 20: 85-89 (relationship to fuscipennis (Wheeler)).
melanocephalus (Emery). S. Ariz. to Honduras. Only the worker is known.
Eciton (Acamatus) melanocephalum Emery, 1895. Zool. Jahrb. Abt. f. System. 8: 260. %.
Eciton (Acamatus) melanocephalum xipe Wheeler, 1914. N. Y. Ent. Soc., Jour. 22: 41. &.
melsheimeri (Haldeman). La., Okla., Tex.; s. to Costa Rica. Only the male is known. Records
citing this species from Utah are probably incorrect.
Labidus melshaemeri(!) Haldeman, 1852. In Stanbury, Exped. Great Salt Lake, p. 368. d.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 418.
microps Borgmeier. Ariz. (Phoenix). Only the male is known.
Neivamyrmex microps Borgmeier, 1955. Studia Ent. 3: 635. d.
minor (Cresson). Kans., Okla., Tex., N. Mex., Ariz., Nev., Calif.; Mexico. Only the male is
known. One of the smallest of our legionary ants.
Labidus minor Cresson, 1872. Amer. Ent. Soc., Trans. 4: 195. 3.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 418 (male).

Biology: Wheeler and Long, 1901. Amer. Nat. 35: 165. —Cole, 1966. Brigham Young Univ.,
Sci. Bul. 7: 26.
mojave (Smith). Calif. (Mojave Desert). Only the male is known.
Eciton (Neivamyrmex) mojave Smith, 1943. Lloydia 6: 196. 3.
moseri Watkins. La., Tex. Ecology: Collected from a nest of Atta texana (Buckley) in La. Only
the worker and queen are known.
Neivamyrmex moseri Watkins, 1968. Kans. Ent. Soc., Jour. 41: 528-531. &.

Taxonomy: Watkins, 1971. Kans. Ent. Soc., Jour. 44: 95-99 (worker, queen; also biological
notes).
nigrescens (Cresson). W. Va., Ky., Tenn., Ga., Ala., Miss., Ill., Iowa, Mo., Ark., La., Nebr., Kans.,

Okla., Tex., Colo., N. Mex., Ariz., Calif.; Mexico. Ecology: Temporary nesting sites are in
decayed logs or stumps or in the ground beneath stones and other objects. Many
foraging activities take place in daylight, and they are highly predaceous on other
insects. Large colonies contain 150,000 to 250,000 workers. Each colony has one
functional queen, and new colonies are formed by splitting. All castes are known.

Labidus nigrescens Cresson, 1872. Amer. Ent. Soc., Trans. 4: 194. d.

Eciton (Acamatus) Schmitti Emery, 1894. Soc. Ent. Ital, Bol. 26: 183. 8.

Taxonomy: Wheeler, 1943. Ent. Soc. Amer., Ann. 36: 331 (larva). —Watkins, 1972. Kans. Ent.
Soe., Jour. 45: 358-363 (worker, distribution).

Biology: Wheeler, 1900. Amer. Nat. 34: 565-574 (sumichrasti, not Norton). —Smith, 1927.
Ent. Soc. Amer., Ann. 20: 401-404. —Cole, 1953. Tenn. Acad. Sci., Jour. 28: 84. —Schneirla,
1958. Insectes Sociaux 5: 215-255. —Rettenmeyer, 1963. Univ. Kans., Sci. Bul. 44: 446-452.
—Gregg, 1963. Ants of Colo., pp. 286-288. —Watkins, 1964. Kans. Ent. Soc., Jour. 37: 22-28
(trail following). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326, pp. 18-19. —Watkins and
Cole, 1966. Tex. Jour. Sci. 18: 254-265 (attraction of workers to secretion of queens).
—Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail following and
trail preference). —Watkins, Gehlbach, and Baldridge, 1967. Southwest. Nat. 12: 455-462
(blind snake follows pheromone trails of nigrescens). —Watkins, Gehlbach, and Kroll,
1969. Ecology 50: 1098-1102 (attractant, repellant secretions). —Plsek, Kroll, and Watkins,
1969. Kans. Ent. Soc., Jour. 42: 452-456 (carabids in raiding columns). —Topoff, 1969. N. Y.
Ent. Soc., Jour. 77: 273-274 (communication). —Topoff, 1969. Psyche 76: 375-381 (predatory
association between carabids and nigrescens). —Topoff, 1970. N. Y. Ent. Soc., Jour. 78:
239-240 (cyclic behavior). —Watkins, Gehlbach, and Plsek, 1972. Tex. Jour. Sci. 23: 34.
(behavior of blind snakes in response to raiding columns).

1332 Hymenoptera in America North of Mexico

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 393 (maxillary and labial palpi).
opacithorax (Emery). Va., N. C., S. C., Ga., Fla. Tenn., Ala., Mo., Ark., Kans., Okla., Tex., N.
Mex., Ariz., Calif.; Mexico to Costa Rica. Ecology: Habits are similar to those of
nigrescens (Cresson). All castes are known.
Eciton (Acamatus) californicum opacithorax Emery, 1894. Soc. Ent. Ital., Bol. 26: 184. %.
Eciton (Acamatus) opacithorax castaneum Borgmeier, 1939. Rev. de Ent. 10: 416. 8.

Biology: Wheeler and Long, 1901. Amer. Nat. 35: 163, 172. —Smith, 1924. Ent. News 35: 84.
—Cole, 1953. Tenn. Acad. Sci., Jour. 28: 84. —Schneirla, 1958. Insectes Sociaux 5: 215-255.
—Rettenmeyer, 1963. Univ. Kans., Sci. Bul. 44: 437-446. —Watkins, 1964. Kans. Ent. Soc.,
Jour. 37: 22-28 (trail following). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326, pp. 19-20.
—Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-265 (attraction of workers to secretion of
queens). —Watkins, Cole, and Baldridge, 1967. Kans. Ent. Soc., Jour. 40: 146-151 (trail
following and trail preference). —Watkins, Gehlbach, and Baldridge, 1967. Southwest. Nat.
12: 455-462 (a blind snake follows pheromone trails of opicithorazx).

Morphology: Borgmeier, 1957. Rev. Brasil. Biol. 17: 393 (maxillary and labial palpi).
pauxillus (Wheeler). Tex.; Mexico. Only the worker and queen are known.
Eciton (Acamatus) pauxillum Wheeler, 1903. Psyche 10: 93. &.

Taxonomy: Watkins, 1971. Kans. Ent. Soc., Jour. 44: 99-101 (worker, queen; biological notes).

Biology: Smith, 1938. Ent. Soe. Wash., Proc. 40: 158.
pilosus mandibularis (Smith). N. Mex., Ariz. Only the male is known. N. pilosus pilosus
(Smith) is found in S. Amer.
Eciton (Neivamyrmex) pilosum mandibulare Smith, 1942. Amer. Midland Nat. 27: 543,
548. o.
pilosus mexicanus (Smith). Miss., Ark., La., Okla., Tex., Calif., s. to Colombia. All castes are
known.
Labidus Mexicanus Smith, 1859. Cat. Hym. Brit. Mus., v.7, p. 7. d.
Eciton clavicornis Norton, 1868. Amer. Ent. Soc., Trans. 2: 46. 9.
Eciton (Labidus) subsulcatum Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 440. 3.
Eciton (Acamatus) aztecum Forel, 1901. Mitt. Nat. Mus. Hamburg 18: 49. 9.
Eciton (Labidus) spininode militarium Santschi, 1929. Wien Ent. Ztg. 46: 85. 9.

Taxonomy: Borgmeier, 1936. Inst. Biol. Veg. Arg. 3: 60. —Reichensperger, 1939. Zool. Jahrb.
Abt. f. System. 73: 297-300.

Biology: Smith, 1924. Ent. News 35: 85. —Watkins and Cole, 1966. Tex. Jour. Sci. 18: 254-265
(attraction of workers to secretion of queens). —Watkins, Cole, and Baldridge, 1967. Kans.
Ent. Soc., Jour. 40: 146-151 (trail following and trail preference).

rugulosus Borgmeier. S. Ariz.; Mexico. Only the worker is known.
Neivamyrmex rugulosus Borgmeier, 1953. Studia Ent. 2: 49. 3.
swainsonii (Shuckard). La., Tex., N. Mex., Ariz., Calif.; s. to Argentina. Only the male is
known.
Labidus swainsonii Shuckard, 1840. Ann. Nat. Hist. 5: 201. d.
Eciton (Acamatus) arizonense Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 414. do.
texanus Watkins. Va., N. C., S. C., Ga., Fla., Tex., Colo., N. Mex., Ariz.; Mexico. All castes are
known.
Neivamyrmesx texanus Watkins, 1972. Kans. Ent. Soc., Jour. 45: 353-358. 9, 2, d.

Taxonomy: Wheeler and Long, 1901. Amer. Nat. 35: 161 (male described as schmitti).
—Borgmeier, 1955. Studia Ent. 3: 496 (male described as nigrescens).
UNPLACED TAXON OF DORYLINAE

Myrmica (Monomarium(!)) coeca Buckley, 1867. Ent. Soc. Phila., Proc. 6: 339. 3. Tex. (San
Saba Co.).

Superfamily FORMICOIDEA 1333

SUBFAMILY CERAPACHYINAE

This small subfamily, found in the tropical regions of the world, exhibits a blending of doryline
and ponerine traits both morphologically and biologically. Only three species are known from
north of Mexico. Little is known concerning the behavior of the New World forms, but they are
predaceous and carnivorous and the colonies are small. Wilson (1958) studied the behavior of
several species from Melanesia and Australia and found them all to be myrmecophagous, feeding
on the broods and sometimes adults of other species of ants. He suggested that these ants carry
on an alternating, group foraging and raiding behavior pattern by which the colony efficiently
exploits the surrounding territory.

After this section was completed, Brown (1975) published on the Cerapachyinae and regarded
the Cerapachyini and Acanthostichini as tribes of the Ponerinae, consequently not recognizing
the Cerapachyinae as a subfamily.

Revision: Brown, 1975. Search, Agr., Ent. (Ithaca) 15, 5 (1): 14-36 (Cerapachyini and
Acanthostichini as tribes of Ponerinae; biol. notes; keys to world genera and species).

Taxonomy: Wheeler, 1920. Psyche 27: 50-51. —Donisthorpe, 1921. London Ent. Soc., Proc.,
pp. xlv-xlvii. —Wheeler, 1922. Amer. Mus. Nat. Hist., Bul. 45: 51-52. —Morley, 1939. Soc.
Ent. France, Bul. 44: 114-118. —Reid, 1941. Roy. Ent. Soc. London, Trans. 91: 421-422.
—Creighton, 1950. Harvard Univ., Mus. Compar. Zool., Bul. 104: 56-59. —Wheeler, 1950.
Psyche 57: 102-113 (larvae). —Kuznezov, 1952. Dusenia 3: 115. —Brown, 1954. Insectes
Sociaux 1: 26-27. —Wheeler and Wheeler, 1964. Ent. Soc. Wash., Proc. 66: 65-71 (larvae).
—Wheeler and Wheeler, 1973. Psyche 80: 204-211 (larvae).

Biology: Wilson, 1958. Insectes Sociaux 5: 129-140 (behavior of certain Melanesian and
Australian species).

TRIBE CERAPACHYINI

Genus CERAPACHYS Smith

Cerapachys Smith, 1858. Linn. Soc. London, Jour. 2: 74.
Type-species: Cerapachys antennatus Smith. Monotypic.
Syscia Roger, 1861. Berlin. Ent. Ztschr. 5: 19.
Type-species: Syscia typhla Roger. Monotypic.
Parasyscia Emery, 1882. In Andre, Spec. Hym. Eur. Alg. 2: 235.
Type-species: Parasyscia piochardi Emery. Monotypic.
A pantropical genus with about 8 New World species, two of which are found in southwestern
United States. For additional generic synonymy, see Brown, 1975 Search, Agr., Ent. (Ithaca) 15,
5 (1): 18-19.

Taxonomy: Kempf, 1972. Studia Ent. 15: 76 (generic synonymy).
augustae Wheeler. Tex., Ariz.; Mexico. The type colony from Austin, Tex., was found 6 inches
below the surface of soil containing limestone chips; 10 workers and a female were
found. Other specimens were found in the stomach of an armadillo.
Cerapachys (Parasyscia) augustae Wheeler, 1902. Biol. Bul. 3: 182. 3, °.

Taxonomy: Smith, 1942. Ent. Soc. Wash., Proc. 44: 63 (male). —Wheeler, 1950. Psyche 57:
106-107 (larva).
Biology: Wheeler, 1903. Psyche 10: 205-209.
davisi Smith. Tex. (Ft. Davis, 5,000 ft.).
Cerapachys (Parasyscia) davisi Smith, 1942. Ent. Soc. Wash., Proc. 44: 64. d.
TRIBE ACANTHOSTICHINI
Genus CTENOPYGA Ashmead

Ctenopyga Ashmead, 1905. Canad. Ent. 37: 382. Nom. nud.
Ctenopyga Ashmead, 1906. Ent. Soc. Wash., Proc. 8: 29.
Type-species: Ctenopyga townsendi Ashmead. Monotypic.

1334

Hymenoptera in America North of Mexico

texanus (Forel). Tex.; Mexico.
Acanthostichus texanus Forel, 1904. Soc. Ent. Belg., Ann. 48: 168. 2.

Ctenopyga townsendi Ashmead, 1906. Ent. Soc. Wash., Proc. 8: 29-30. 2, d.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 400 (female). —Smith, 1955.
Brooklyn Ent. Soc., Bul. 50: 48-50 (synonymy).

SUBFAMILY PONERINAE

One of the smaller subfamilies of ants, with most species found in the tropical regions of the
world. Most of the North American forms are either northern extensions of Neotropical taxa or
have been introduced by commerce from other parts of the world. These are primitive ants

whic

h nest in small colonies of a few hundred individuals or less, mostly in soil or rotting wood.

They are predaceous and carnivorous.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 111-144 (larvae). —Wheeler
and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 443-462 (larvae). —Wheeler and Wheeler,

1971. Ent. Soc. Amer., Ann. 64: 1197-1217 (larvae). —Wheeler and Wheeler, 1974. Ent. Soc.

Wash., Proc. 76: 278-281 (larvae). —Wheeler and Wheeler, 1976. Amer. Ent. Soc., Trans.

102: 41-64 (supplementary studies on larvae).

Biology: Wheeler, 1910. Ants, pp. 225-245. —Wilson, 1958. Evolution 12: 24-31 (beginnings of
nomadic and group-predatory behavior).

TRIBE AMBLYOPONINI

Revision: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bul. 122: 145-230.

Genus AMBLYOPONE Erichson

Amblyopone Erichson, 1842. Arch. f. Naturgesch. 8: 260.
Type-species: Amblyopone australis Erichson. Monotypic.
Stigmatomma Roger, 1859. Berlin Ent. Ztschr. 3: 250.
Type-species: Stigmatomma denticulatum Roger. Desig. by Bingham, 1903.
Arotropus Provancher, 1881. Nat. Canad. 12: 205.
Type-species: Arotropus binodosus Provancher. Monotypic.
Amblyopopona Schulz, 1906. Spolia Hym., p. 154. Emend.
Stigmatomma subg. Xymmer Santschi, 1914. Bol. Lab. Zool. Gen. e Agr. Portici 8: 311.
Type-species: Stigmatomma (Xymmer) muticum Santschi. Monotypic.
Stigmatomma subg. Fulakora Mann, 1919. Harvard Univ., Mus. Comp. Zool., Bul. 63: 279.
Type-species: Stigmatomma (Fulakora) celata Mann. Orig. desig.
Amblyopone subg. Neoamblyopone Clark, 1927. In Wheeler, Amer. Acad. Arts and Sci.,
Proc. 62: 1.
Type-species: Amlyopone (Neoamblyopone) clarki Wheeler. Monotypic.
Amblyopone subg. Protamblyopone Clark, 1927. In Wheeler, Amer. Acad. Arts and Sci.,
Proc. 62: 1.
Type-species: Amblyopone (Protamblyopone) aberrans Wheeler. Monotypic.
Lithomyrmez Clark, 1928. Roy. Soc. W. Australia, Jour. 14: 30.
Type-species: Lithomyrmesx glauerti Clark. Orig. desig.
Ericapelta Kusnezov, 1955. Zool. Anz. 154: 273.
Type-species: E'ricapelta egregia Kusnezov, Monotypic.

This genus is represented in the tropical and temperate regions of the world though it is more
highly developed in the Australian Region than elsewhere. The Nearctic forms commonly occur
in wooded areas, especially those that are well shaded. They are subterranean, and the workers

are

timid and slow of movement. The female of pallipes forages for food during the period of

nest founding, an archaic habit in ants.
Revision: Creighton, 1940. Amer. Mus. Novitates 1079: 1-8. —Brown, 1960. Harvard Univ.,

Mus. Comp. Zool., Bul. 122: 155-169.

Taxonomy: Brown, 1949. Psyche 56: 81-88. —Wheeler and Wheeler, 1964. Ent. Soc. Amer.,

Ann. 57: 444-446 (larvae).

Superfamily FORMICOIDEA 1335

Biology: Haskins and Haskins, 1951. Amer. Midland Nat. 45: 482-445 (colony founding of A,
australis Erichson).
Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116: 476 (proventriculus).
oregonense (Wheeler). B. C., Wash., Oreg., Calif. Ecology: Usually at low elevations in the

coastal mountains.
Stigmatomma pallipes oregonense Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 389. 8,

Gr
Taxonomy: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bul. 122: 169, 183.
pallipes (Haldeman). Ont., Que. s. to Ga. w. to Wis., Iowa, Okla., Colo., Tex., Ariz. Ecology: The
small colonies are most common in areas of heavy cover and considerable precipitation.

Chilopods appear to be the main food source.
Typhlopone pallipes Haldeman, 1844. Acad. Nat. Sci. Phila., Proc. 2: 54. &.
Stigmatomma serratum Roger, 1859. Berlin. Ent. Ztschr. 3: 251. &.
Arotropus binodosus Provancher, 1881. Nat. Canad. 12: 206. °.
Stigmatomma pallidipes(!) var. Wheeleri Santschi, 1913. Soc. Ent. Belg., Ann. 57: 429. %,
OrOr
Stigmatomma pallipes arizonense Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 389. 9.
Stigmatomma pallipes montigena Creighton, 1940. Amer. Mus. Novitates 1079: 2, 7. 9, °.
Stigmatomma pallipes subterranea Creighton, 1940. Amer. Mus. Novitates 1079: 3, 8. 8.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 261-262 (each caste). —Wheeler and
Wheeler, 1952. Amer. Midland Nat. 48: 114 (larva). —Brown, 1960. Harvard Univ., Mus.
Comp. Zool., Bul. 122: 169, 183-185. —Francoeur and Beique, 1966. Canad. Ent. 98: 141
(Provancher types).

Biology: Wheeler, 1900. Biol. Bul. 2: 56-64. —Haskins, 1928. N. Y. Ent. Soc., Jour. 36: 179-184.
—Buren, 1944. Iowa State Col., Jour. Sci. 18: 279. —Cole, 1953. Tenn. Acad. Sci., Jour. 28:
84. —Gregg, 1963. Ants of Colo., pp. 279-282.

Morphology: Whelden, 1957. N. Y. Ent. Soc., Jour. 65: 1-21 (anatomy). —Gotwald, 1969. N. Y.
Agr. Expt. Sta. (Cornell Univ.), Mem. 408: 25-42 (mouthparts).

trigonignatha Brown. N. C. (Concord). Ecology: Sifted from leaf mold in Berlese funnel.
Amblyopone (Stigmatomma) trigonignatha Brown, 1949. Psyche 56: 81-84. &.

Taxonomy: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bul. 122: 169, 185.
Genus PRIONOPELTA Mayr

Prionopelta Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. K1. Sitzber. 53: 503.
Type-species: Prionopelta punctulata Mayr. Monotypic.

Renea Donisthorpe, 1947. Ann. and Mag. Nat. Hist. (11) 14: 183. Preoce. by Nevill, 1880.
Type-species: Renea testacea Donisthorpe. Orig. desig.

Examblyopone Donisthorpe, 1949. Ann. and Mag. Nat. Hist. (12) 2: 401.
Type-species: Examblyopone churchilli Donisthorpe. Orig. desig.

A tropicopolitan genus of small, soil-inhabiting ants.
Revision: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bul. 122: 173-178.

Taxonomy: Brown, 1951. Brooklyn Ent. Soc., Bul. 46: 102. —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 120 (arvae). —Brown, 1953. Breviora 11: 11. —Wheeler and
Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 447 (larvae; Prinopelta(!)).

antillana Forel. Fla. (Juniper Springs, Marion Co.); W. Indies, Central America to Brazil,
Bolivia (?). Ecology: Specimens have been found in soil.
Prionopelta punctulata antillana Forel, 1909. Deut. Ent. Ztschr., p. 239. %.

Taxonomy: Brown, 1960. Harvard Univ., Mus. Comp. Zool., Bul. 122: 178.
Biology: Kempf, 1961. Studia Ent. 4: 489-490 (in soil samples in Surinam).
TRIBE PLATYTHYREINI
Revision: Brown, 1975. Search, Agr., Ent. (Ithaca) 15, 5 (1): 4-11 (world genera and species).

1336 Hymenoptera in America North of Mexico

Taxonomy: Brown, 1952. Breviora 6: 1-6.
Genus PLATYTHYREA Roger

Platythyrea Roger, 1863. Berlin Ent. Ztschr. 7: 172.
Type-species: Pachycondyla punctata Smith. Desig. by Bingham, 1903.
A tropicopolitan genus with 8 Neotropical species, only one of which reaches the United
States.

Taxonomy: Brown, 1952. Breviora 6: 4 (in key). —Wheeler and Wheeler, 1964. Ent. Soc.
Amer., Ann. 57: 446 (larvae). —Wheeler and Wheeler, 1976. Amer. Ent. Soc., Trans. 102: 41
(revised characterization of larvae).

punctata (Smith). S. Fla. and s. Tex., s. to W. Indies, Central Amer., and Brazil. Ecology:
Nests in small colonies up to a few hundred individuals each, usually in rotten logs and
stumps or under the bark of trees in shady situations. Workers are active, forage singly,
and are carnivorous and predatory.
Pachycondyla punctata Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 108. 9, ¢.
Platythyrea pruinosa Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 962. %.

Taxonomy: Forel, 1893. Ent. Soc. London, Trans., p. 358 (worker, female). —Mann, 1916.
Harvard Univ., Mus. Comp. Zool., Bul. 60: 403. —Wheeler and Wheeler, 1952. Amer.
Midland Nat. 48: 119 (larva).

Biology: Forel, 1899. Rev. Suisse Zool. 9: 335 (in Barbados). —Wheeler, 1905. Amer. Mus.
Nat. Hist., Bul. 21: 80 (in Bahamas). —Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 824
(in Puerto Rico). —Brown, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116: 229 (in
Mexico).

TRIBE ECTATOMMINI
Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 175-362.
Genus ECTATOMMA Smith

Ectatomma Smith, 1859. Cat. Hym. Brit. Mus., v. 6, p. 102.
Type-species: Formica tuberculata Olivier. Desig. by Bingham, 1903.

A Neotropical genus of about 14 species, none of which are native to the United States. One
species was introduced into southern Texas for biological control purposes, but the attempt
failed.

Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 206-211, 295-299.
Taxonomy: Wheeler and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 449 (larvae).

Biology: Weber, 1946. Ent. Soc. Wash., Proc. 48: 1-16 (E. tuberculatum (Olivier) and E.
ruidum (Roger).
tuberculatum (Olivier). S. Tex. (?); Mexico to n. Argentina. Ecology: Nests are in the soil and

are composed of several hundred individuals. Workers are predatory and carnivorous
and also attend membracids and aphids on plants. Introduced into Victoria Co., Texas
from Guatemala in 1904-1905 to combat the cotton boll weevil. The attempted
introduction led to a fiery confrontation between O. F. Cook, instigator of the
experiment, and W. M. Wheeler. Known as the kelep.

Formica tuberculata Olivier, 1791. Eneycel. Meth., Dict. Ins., v. 6, p. 498. &.

Formica tridentata Fabricius, 1804. Systema Piezatorum, p. 42. 3.

Ectatomma ferrugineus Norton, 1868. Comm. Essex Institute 6: 5. 3, 6.

Ectatomma tuberculatum var. punctigerum Emery, 1890. Soc. Ent. France, Ann. 10: 56.
OF

Ectatomma tuberculatum var. acrista Forel, 1909. Deut. Ent. Ztschr., p. 254. 8, 2.

Ectatomma tuberculatum var. irregularis Santschi, 1921. Soc. Vaud. Nat. des Sci., Bul. 54:
83. 8.

Superfamily FORMICOIDEA 1337

Taxonomy: Lever, 1930. Ent. Monthly Mag. 66: 214 (female). —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 129-133 (larva). —Brown, 1958. Harvard Univ., Mus. Comp. Zool.,
Bul. 118: 209, 211, 298-299. —Kempf, 1962. Studia Ent. 5: 2-3.

Biology: Cook, 1904. U. S. Dept. Agr., Div. Ent., Bul. 49: 1-15. —Cook, 1904. Science 19:
862-864. —Cook, 1904. Science 20: 310-312 (pupation). —Wheeler, 1904. Science 20: 437-440
(pupation; feasibility of introduction). —Cook, 1904. Science 20: 611-612. —Wheeler, 1904.
Science 20: 766-768. —Cook, 1905. U. S. Dept. Agr., Bur. Ent., Tech. Ser. 10: 1-55.
—Wheeler, 1905. Science 21: 706-710 (criticism of Cook’s work). —Cook, 1906. Science 23:
187-189. —Wheeler, 1906. Science 23: 348-350. —Weber, 1946. Ent. Soc. Wash., Proc. 48:
1-16 (biology and economic significance).

Morphology: Haskins and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37: 100-162 (physiology).
—Gotwald, 1969. N. Y. Agr. Expt. Sta. (Cornell Univ.), Mem. 408: 25-42 (mouthparts).

Genus GNAMPTOGENYS Roger

Gnamptogenys Roger, 1863. Berlin Ent. Ztschr. 7: 174.
Type-species: Ponera tornata Roger. Desig. by Emery, 1911.
Ectatomma subg. Stictoponera Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37: 539.
Type-species: Ectatomma coxale Roger. Desig. by Bingham, 1903.
Ectatomma subg. Holcoponera Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37: 540.
Type-species: Gnamptogenys striatula Mayr. Desig. by Emery, 1911.
Alfaria Emery, 1896. Soc. Ent. Ital., Bul. 28: 41.
Type-species: Alfaria simulans Emery. Monotypic.
Ectatomma subg. Poneracantha Emery, 1897. Mus. Civ. Stor. Nat. Genova, Ann. 38: 547.
Type-species: Ectatomma (Holcoponera?) bispinosum Emery. Monotypic.
Rhopalopone Emery, 1897. Mus. Civ. Stor. Nat. Genova, Ann. 38: 549.
Type-species: Rhopalopone epinotalis Emery. Monotypic.
Emeryella Forel, 1901. Soc. Ent. Belg., Ann. 45: 334.
Type-species: E’meryella schmitti Forel. Monotypic.
Ectatomma subg. Mictoponera Forel, 1901. Soc. Ent. Belg., Ann. 45: 372.
Type-species: E'ctatomma (Mictoponera) diehli Forel. Monotypic.
Ectatomma subg. Parectatomma Emery, 1911. In Wytsman, Gen. Ins., fase. 118, p. 44.
Type-species: Ectatomma (Gnamptogenys) triangulare Mayr. Orig. desig.
Spaniopone Wheeler and Mann, 1914. Amer. Mus. Nat. Hist., Bul. 33: 11.
Type-species: Spaniopone haytiana Wheeler and Mann. Monotypic.
Wheeleripone Mann, 1919. Harvard Univ., Mus. Comp. Zool., Bul. 63: 282.
Type-species: Wheeleripone albiclava Mann. Orig. desig.
Opisthoscyphus Mann, 1922. U.S. Natl. Mus., Proc. 61: 4.
Type-species: Opisthoscyphus scabrosus Mann. Monotypic.
Ectatomma subg. Commateta Santschi, 1929. Zool. Anz. 82: 476.
Type-species: Ectatomma (Parectatomma) bruchi Santschi. Orig. desig.
Ectatomma subg. Tammoteca Santschi, 1929. Zool. Anz. 82: 476.
Type-species: E'ctatomma concinna Smith. Orig. desig.
Emeryella subg. Barbourella Wheeler, 1930. New England Zool. Club, Proc. 12: 10.
Type-species: Emeryella (Barbourella) banksi Wheeler. Orig. desig.

Brown (1958) treated 81 species of which about 64 are found in the New World tropics and di-
vided the genus into four species groups. The question has been raised as to whether our only
species is native or adventive.

Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 211-241, 299-330.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 133-134 (larvae). —Wheeler
and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 540 (larvae). —Wheeler and Wheeler, 1976.
Amer. Ent. Soc., Trans. 102: 43 (revised characterization of larvae).

hartmani (Wheeler). La. (Lucky), Tex. (Huntsville); Honduras. Ecology: Specimens have been
taken from soil under banana trees in Honduras and from nests of Trachymyrmex
septentrionalis (McCook) in La.
Ectatomma (Parectatomma) hartmani Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 390.
%.

1338 Hymenoptera in America North of Mexico

Taxonomy: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 228, 230, 234, 302.
—Brown, 1961. Psyche 68: 69 (from Honduras; single worker from Tex. described by
Wheeler possibly a locality error or adventive specimen).

Biology: Echols, 1964. Ent. Soc. Amer., Ann. 57: 137 (La.; first reproductive colonies taken in
U.S).

Genus PROCERATIUM Roger

Proceratium Roger, 1863. Berlin. Ent. Ztschr. 7: 171.
Type-species: Proceratium silaceum Roger. Monotypic.
Sysphingta Roger, 1863. Berlin. Ent. Ztschr. 7: 175.
Type-species: Sysphingta micrommata Roger. Monotypic.
Sysphincta Mayr, 1865. Reise d. Novara, Zool. 1 (1) Formicidae, p. 12. Emend.

About 24 world species are known, most of which are found in the warmer parts of the
northern temperate region of the world. They are hypogaeic, and the small colonies of two or
three dozen individuals are found in well-rotted, moist wood such as that of logs and stumps. A
constant, high moisture content is essential. Toward the cooler areas they may be under the

deepest rocks. Workers are sluggish and carnivorous, apparently feeding almost exclusively on
the eggs of other Arthropods.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 264-266. —Brown, 1958. Harvard
Univ., Mus. Comp. Zool., Bul. 118: 241-248.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 135-137. (larvae). —Wheeler
and Wheeler, 1964. Ent. Soc. Amer., Ann. 57: 451 (larvae). —Snelling, 1967. Los Angeles
Co. Mus., Contrib. Sci. 124: 1-10 (key to New World species). —Wheeler and Wheeler,
1971. Ent. Soc. Amer., Ann. 64: 1202 (larvae).

Biology: Brown, 1958. Psyche 65: 115 (predation of arthropod eggs).
californicum Cook. Calif. Found in several scattered localities from Sutter Co. to Los Angeles
Co. The worker is unknown.
Procratium(!) californicum Cook, 1953. The Ants of Calif., pp. 45-46. d.

Taxonomy: Snelling, 1967. Los Angeles Co. Mus., Contrib. Sci. 124: 1-10 (female, male).

croceum (Roger). Va. to Fla. w. to IIl., Tex.
Ponera crocea Roger, 1860. Berlin Ent. Ztschr. 4: 288. @.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 264 (worker, female). —Smith, 1930.
Ent. Soc. Amer., Ann. 23: 390-392 (male). —Wheeler and Wheeler, 1952. Amer. Midland
Nat. 48: 135 (larva).

Biology: Haskins, 1930. N. Y. Ent. Soc., Jour. 38: 121-126. —Brown, 1958. Harvard Univ.,
Mus. Comp. Zool., Bul. 118: 246-247.
melinum (Roger). “Carolina”; Europe. A European species that doubtfully occurs in North
America and has not been collected here since it was originally described; types may
have been mislabeled.
Ponera melina Roger, 1860. Berlin. Ent. Ztschr. 4: 291. %, 2, do.
Sysphingta Europaea Forel, 1886. Soc. Ent. Belg., Ann. (C. R.) 30: CLXIII. &.
Sysphincta europaea rossica Arnoldi, 1930. Zool. Anz. 91: 144. ¥, 9.
Sysphincta fialai Kratochvil, 1944. In “Mohelno.” Arch. Svaz. ochr. prir. dom. Morave,
Svazek 6: 54, 86.

Taxonomy: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 243, 246-248, 334
(doubtful in N. Amer.; also biological notes).
pergandei (Emery). Mass. to Fla. w. to Iowa, Ark., La.
Sysphincta pergandei Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 264. %.

Taxonomy: Smith, 1928. Ent. News 39: 242-243 (male). —Cole, 1940. Amer. Midland Nat. 24:
36. —Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 243, 246, 247-248, 336 (also
biological notes).

Superfamily FORMICOIDEA 1339

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 276, 304. —Wesson and Wesson, 1940. Amer.

Midland Nat. 24: 90-91 (ate only inside of gaster of dead workers of other ants.)
silaceum Roger. Mass., s. Ont. (Pelee Is. and vicinity) s. to n. Fla., w. to Ill, Ark., Okla.
Proceratium silaceum Roger, 1863. Berlin. Ent. Ztschr. 7: 172. %.
Proceratium crassicorne Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 265. §.
Proceratium crassicorne var. vestitum Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 266. %.
Proceratium silaceum rugulosum Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 390. &,
2

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 137 (larva). —Brown, 1958.

Harvard Univ., Mus. Comp. Zool., Bul. 118: 241, 245-248, 336 (also biology).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 373, 375. —Wheeler, 1916. Ind. Acad.
Sci., Proc. 26: 460. —Smith, 1928. Ent. News 29: 244. —Dennis, 1938. Ent. Soc. Amer., Ann.
31: 272, 273, 276, 304. —Wesson and Wesson, 1940. Amer. Midland Nat. 24: 91.

Morphology: Kennedy and Talbot, 1939. Ind. Acad. Sci., Proc. 48: 206-210 (each caste; also
biological notes).

Genus DISCOTHYREA Roger

Discothyrea Roger, 1863. Berlin. Ent. Ztschr. 7: 176.
Type-species: Discothyrea testacea Roger. Monotypic.
Pseudosysphincta Arnold, 1916. South. Afr. Mus., Ann. 14: 161.
Type-species: Pseudosysphincta poweri Arnold. Orig. desig.
Prodiscothyrea Wheeler, 1916. Roy. Soc. South. Aust., Trans. 60: 33.
Type-species: Prodiscothyrea velutina Wheeler. Monotypic.
Pseudosphincta Wheeler, 1922. Amer. Mus. Nat. Hist., Bul. 45: 645, 762. Variant spelling of
Pseudosysphincta.

Most of the 26 species of this genus are found in the tropical and southern temperate regions
of the world. Because of their small size and cryptobiotic habits, they are not commonly collected
and their biology is poorly known. The colonies are small and a number of forms have been col-
lected from leaf litter, humus, and rotten logs.

Revision: Brown, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 118: 248-253.

Taxonomy: Borgmeier, 1949. Rev. Brasil. Biol. 9: 205. —Borgmeier, 1957. Acad. Brasil. de
Cien., An. 29: 124-125 (male). —Wheeler and Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1202
(larvae).

Biology: Brown, 1958. Psyche 64: 115 (as predators of eggs of other arthropods).

testacea Roger. N. C. to Fla.; Okla. Ecology: Some have been found in pine forest litter and in
soil.
Discothyrea testacea Roger, 1863. Berlin. Ent. Ztschr. 7: 177. 3, °.

Taxonomy: Weber, 1939. Ent. Soc. Amer., Ann. 32: 99 (worker, female). —Smith and Wing,
1954. N. Y. Ent. Soc., Jour. 62: 110-112 (worker, female). —Smith, 1955. Brooklyn Ent.
Soc., Bul. 50: 98 (probable type locality; collector). —Brown, 1958. Harvard Univ., Mus.
Comp. Zool., Bul. 118: 253, 341-342.

TRIBE PONERINI

Taxonomy: Brown, 1963. Breviora 190: 1-10.
Genus PACHYCONDYLA Smith

Pachycondyla Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 105.
Type-species: Formica crassinoda Latreille. Desig. by Emery, 1901.
Neoponera Emery, 1901. Soc. Ent. Belg., Ann. 45: 40, 43.
Type-species: Formica villosa Fabricius. Orig. desig.
Euponera subg. Trachymesopus Emery, 1911. In Wytsman, Gen. Ins., fase. 118, p. 84.
Type-species: Formica stigma Fabricius. Orig. desig.

1340 Hymenoptera in America North of Mexico

Brown (1973) has synonymized Neoponera, Trachymesopus, and several other genera with
Pachycondyla. This is a large genus in the tropical regions of the world, but only three species
reach the United States.

Revision: Emery, 1890. Soc. Ent. France, Ann. 10: 71-74 (Pachycondyla).

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 119: 352 (Trachymesopus
not a subgenus of Euponera). —Kempf, 1960. Studia Ent. 3: 423-428 (New World species
of Trachymesopus). —Kempf, 1961. Rev. Brasil. Ent. 10: 89-204 (Brazilian species of
Pachycondyla). —Brown, 1963. Breviora 190: 6-8 (three species groups of
Trachymesopus). —Kempf, 1964. Studia Ent. 7: 49-52 (key to species of Pachycondyla).
—Brown, 1973. In Meggers, et al., Tropical forest ecosystems in Africa and S. Amer., pp.
178-185 (generic synonymy).

harpax (Fabricius). La., Tex. s. to Brazil; W. Indies. Ecology: Colonies of about 150 individuals

are found in rotten logs and stumps or in soil beneath objects. Workers avoid direct
sunlight and forage in the morning and in shade for other insects and myriapods on
which they feed. There are both ergatoid and normal females.

Formica harpax Fabricius, 1804. Systema Piezatorum, p. 401. &.

Pachycondyla Montezumia Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 108. 2, 3.

Pomera(!) amplinoda Buckley, 1866. Ent. Soc. Phila., Proc. 6: 171. %.

Pachycondyla Orizabana Norton, 1868. Amer. Nat. 2: 64. %.

Pachycondyla harpax var. dibullana Forel, 1901. Rev. Suisse Zool. 9: 347. %.

Pachycondyla harpax var. irina Wheeler, 1925. Arkiv for Zool. 17A: 5. ¥.

Pachycondyla harpax var. concinna Wheeler, 1925. Arkiv for Zool. 17A: 5. 8, @.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 401-403 (each caste). —Brown,
1950. Wasmann Jour. Biol. 8: 247-248 (species synonymy). —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 617-618 (larva). —Kempf, 1961. Rev. Brasil. Ent. 10: 194.

Biology: Wheeler, 1900. Biol. Bul. 2: 1-31. —Haskins and Enzmann, 1938. N. Y. Acad. Sci.,
Ann. 37: 150-151 (formation of new colony; in Canal Zone).
stigma (Fabricius). Fla.; W. Indies, Mexico s. to n. Argentina; S. China to Samoa, n.

Queensland. Ecology: They prefer to nest in moist, dead logs or stumps, occasionally
under stones. Apparently a tramp species distributed by commerce outside the New
World. Whether or not it is endemic or adventive to Florida is unknown.

Formica stigma Fabricius, 1804. Systema Piezatorum, p. 400. 2.

Ponera quadridentata Smith, 1859. Linn. Soc. London, Jour. Zool. 3: 143. 2.

Ponera Americana Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 13: 722. &.

Euponera (Trachymesopus) nixoni Donisthorpe, 1943. Ann. and Mag. Nat. Hist., ser. 11,
10: 441. 2.

Euponera (Trachymesopus) brunneus Donisthorpe, 1947. Ann and Mag. Nat. Hist., ser. 11,
14: 300-301. 3, @.

Euponera (Trachymesopus) sexdentatus Donisthorpe, 1949. Ann. and Mag. Nat. Hist., ser.
12, 1: 746. 8.

Taxonomy: Smith, 1934. Ent. Soc. Amer., Ann. 27: 561-564. —Wheeler and Wheeler, 1952.
Amer. Midland Nat. 48: 627 (larva). —Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bul.
119: 355 (synonymy and distribution). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 22
(Polynesia). —Wheeler and Wheeler, 1976. Amer. Ent. Soc., Trans. 102: 55-58 (descriptions
of larvae from different localities; as Mesoponera stigma).

Biology: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 824. —Haskins and Enzmann, 1938.
N. Y. Acad. Sci., Ann. 37: 151 (colony formation; Canal Zone). —Kempf, 1960. Studia Ent.
3: 427-428. —Wilson, 1959. Evolution 13: 128 (distribution; ecology).
villosa (Fabricius). S. Tex. (as far north as San Antonio) s. to n. Argentina. Ecology: Colonies

occur in the soil and in logs and stumps. Workers run rapidly in the bright sun in search
of insects on which they feed. They can sting severely. The largest ponerine ant in the
U.S.

Formica villosa Fabricius, 1804. Systema Piezatorum, p. 409. ¥.

Ponera bicolor Guerin, 1845. Iconogr. Regne Anim., Ins., v. 7, p. 242. 3.

Ponera pilosa Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 95. 3.

Superfamily FORMICOIDEA 1341

Ponera pedunculata Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 96. 8.

Taxonomy: Roger, 1861. Berlin Ent. Ztschr. 5: 1 (worker, male). —Wheeler and Wheeler,
1952. Amer. Midland Nat. 48: 615 (larva). —Wheeler and Wheeler, 1971. Ent. Soc. Amer.,
Ann. 64: 1205 (larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 404.

Genus BRACHYPONERA Emery

Euponera subg. Brachyponera Emery, 1901. Soc. Ent. Belg., Ann. 45: 43.
Type-species: Ponera sennaarensis Mayr. Orig. desig.

A small genus found in the Old World. The one species in North America is adventive.

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 119: 346 (not a subgenus of
Euponera). —Brown, 1958. Acta Hym. 1: 21.
solitaria (Smith). Va., N. C., Ga.; Japan, China, and adjacent areas. Ecology: The small colonies
are found in moist, rotten wood or in soil beneath objects; they prefer dark, damp
places. Food consists of small arthropods. Accidentally introduced into N. Amer.
Ponera solitaria Smith, 1874. Ent. Soc. London, Trans., p. 404. 8.

Taxonomy: Smith, 1934. Ent. Soc. Amer., Ann. 27: 558-561 (description; first record for N.
Amer.). —Wheeler and Wheeler, 1952. Amer. Midland Nat. 49: 629 (larva). —Wheeler and
Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1207 (larva).

Biology: Smith, 1934. Ent. Soc. Amer., Ann. 27: 560-561.
Genus CRYPTOPONE Emery

Cryptopone Emery, 1892. Soc. Ent. France, Bul. 61: CCLXXV.
Type-species: Amblyopone? testacea Motschulsky. Monotypic.

Most species of this genus are found in Southeast Asia and vicinity; only one, apparently en-
demic species occurs in the United States.

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 119: 357-361. —Brown, 1963.
Breviora 190: 6.
gilva (Roger). Ga., Tenn., Ala., Miss., Ark., La., Tex. Ecology: The small colonies nest in moist
dead logs or stumps, preferring loose frass under bark.
Ponera gilva Roger, 1863. Berlin. Ent. Ztschr. 7: 170. %.
Euponera (Trachymesopus) gilva harnedi Smith, 1929. Ent. Soc. Amer., Ann. 22: 543. 9.

Taxonomy: Smith, 1934. Ent. Soc. Amer., Ann. 27: 561-563. —Creighton and Tulloch, 1930.
Psyche 37: 73-79 (each caste). —Wheeler and Wheeler, 1952. Amer. Midland Nat. 48:
625-627 (larva). —Brown, 1963. Breviora 190: 6.

Biology: Haskins, 1931. N. Y. Ent. Soc., Jour. 39: 507-521. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31: 277, 304.

Genus PONERA Latreille

Ponera Latreille, 1804. Nouv. Dict. Hist. Nat. 24: 179.
Type-species: Formica contracta Latreille. Desig. by Latreille, 1805.
Pseudocryptopone Wheeler, 1933. Amer. Mus. Novitates 672: 12-13.
Type-species: Cryptopone tenuis Emery. Orig. desig.
Selenopone Wheeler, 1933. Amer. Mus. Novitates 672: 19.
Type-species: Ponera selenophora Emery. Orig. desig.

Taylor (1967) treated 28 world species and divided the genus into several species groups. Most
species are found in the Indo-Australian area. The only two New World species are North
American. Most forms are found in forested areas where they nest in small colonies in rotten
wood or stumps or in the soil beneath cover. The workers are carnivorous.

Revision: Smith, 1936. Ent. Soc. Amer., Ann. 29: 420-430 (in part; U. S.). —Taylor, 1967.
Pacific Ins. Monog. 13: 1-112 (world).

1342 Hymenoptera in America North of Mexico

Taxonomy: Wilson, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116: 356-357. —Brown, 1958.
Acta Hym. 1: 22-23. —Wheeler and Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1208 (larvae).

SPECIES GROUP LEAE

exotica Smith. N. C., Okla. Ecology: Specimens are all from Berlese samples of leaf litter or
leaf mold. Possibly introduced. Affinities are with the Indo-Australian fauna.
Ponera exotica Smith, 1962. Acta Hym. 1: 378-382. 8, 9°.

Taxonomy: Taylor, 1967. Pacific Ins. Monog. 13: 96-97.

SPECIES GROUP COARCTATA

pennsylvanica Buckley. N. S., Que. s. to Fla. w. to Ont., N. Dak., Colo., Utah, N. Mex. Ecology:
Most abundant in the eastern deciduous forests, east of the 97th meridian, with only
scattered records in the western states. Nests are found under rotting logs, in rotting
stumps, small fragments of wood, acorns and other objects, or in soil or leaf mold. In
drier habitats they may nest under stones. Workers forage in or on the ground and are
carnivorous. Only occasionally a household pest.
Ponera Pennsylvanica Buckley, 1866. Ent. Soc. Phila., Proc. 6: 171. @.

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 631 (larva). —Taylor, 1967.
Pacific Ins. Monog. 13: 29-38 (also biology, ecology). —Wheeler and Wheeler, 1971. Ent.
Soc. Amer., Ann. 64: 1210 (larva).

Biology: Wheeler, 1900. Biol. Bul. 2: 22-23, 43-56. —Wheeler, 1917. Conn. State Geol. and Nat.
Hist. Survey Bul. 22: 581. —Haskins and Enzmann, 1988. N. Y. Acad. Sci., Ann. 37: 152-155
(colony formation). —Headley, 1952. Ent. Soc. Amer., Ann. 45: 436-438. —Kannowski, 1959.
Insectes Sociaux 6: 118 (colony founding). —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 91-92. —Gregg, 1963. Ants of Colo., pp. 281-282, 284.

Morphology: Gotwald, 1969. N. Y. Agr. Expt. Sta. (Cornell Univ.) Mem. 408: 25-42
(mouthparts).

Genus HYPOPONERA Santschi

Ponera subg. Hypoponera Santschi, 1938. Soc. Ent. France, Bul. 43: 79.
Type-species: Ponera abeillei Andre. Orig. desig.

A large cosmopolitan genus whose habits are similar to those of species of Ponera. Prior to
Taylor, 1967, the species below were assigned to the genus Ponera.

Revision: Smith, 1936. Ent. Soc. Amer., Ann. 29: 420-430 (in part, as Ponera; U. S.).

Taxonomy: Taylor, 1967. Pacific Ins. Monog. 13: 9-14 (a distinct genus). —Taylor, 1968. Ent.
News 79: 63-66 (list of N. Amer. species). —Wheeler and Wheeler, 1971. Ent. Soc. Amer.,
Ann. 64: 1210 (larvae).

gleadowi (Forel). Md. (Priest Bridge); Asia. Apparently a widespread tramp species. Possibly
introduced. No other confirmed records from the U. S.
Ponera Gleadowi Forel, 1895. R. Accad. Sci. Bologna, Mem. 5: 292-293. 8.
Ponera japonica formosae Forel, 1913. Arch. f. Naturgesch. 79: 186.
Ponera oblongiceps Smith, 1939. Ent. Soc. Wash., Proc. 41: 76-78. 3, 2, apterous ergatoid
é.

Taxonomy: Wilson, 1958. Harvard Univ., Mus. Comp. Zool., Bul. 119: 326 (Melanesia;
provisional synonymy). —Taylor, 1967. Pacific Ins. Monog. 13: 11, 12, 76. —Wilson and
Taylor, 1967. Pacific Ins. Monog. 14: 29 (unknown from Polynesia; synonymy given for
gleadowi by Wilson, 1958 provisionally transferred to synonymy of punctatissima
(Roger)). —Taylor, 1968. Ent. News 79: 65 (U. S.).

inexorata (Wheeler). S. C. to Fla. w. to Tex., Ariz.; s. to Central Amer.
Ponera inexorata Wheeler, 1903. Psyche 10: 94. %, °.
Taxonomy: Taylor, 1968. Ent. News 79: 65.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 406.

Superfamily FORMICOIDEA 1343

opaciceps (Mayr). S. C. to Fla. w. to Colo., Ariz.; s. to Argentina, W. Indies; s. e. Asia,
Polynesia. Probably spread from New World to Old World by commerce.
Ponera opaciceps Mayr, 1887. Zool.-Bot. Gesell. Wien., Verh. 37: 536. %.
Ponera perkinsi Forel, 1899. Fauna Hawaiiensis, p. 117. 3, 2, d.
Ponera andrei Emery, 1900. Termes. Fuzetek 23: 318. 3.

Taxonomy: Smith, 1929. Ent. Soc. Amer., Ann. 22: 545-546. —Smith and Haug, 1931. Ent.
Soe. Amer., Ann. 24: 507-509 (ergatandrous form). —Wheeler and Wheeler, 1952. Amer.
Midland Nat. 48: 364 (larva). —Kempf, 1962. Studia Ent. 5: 7-9. —Wilson and Taylor, 1967.
Pacific Ins. Monog. 14: 28 (Polynesia). —Taylor, 1968. Ent. News 79: 65. —Wheeler and
Wheeler, 1971. Ent. Soc. Amer., Ann. 64: 1210 (larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 125, 404. —Smith, 1927. Ent. News
38: 308-309. —Kempf, 1960. Studia Ent. 3: 391. —Gregg, 1963. Ants of Colo., pp. 283-284.
opacior (Forel). Va. to Fla. w. to Ohio, Iowa, Colo., Tex.; Oreg., Calif.; Mexico s. to Chile,
Argentina; W. Indies. More sporadically distributed west of Tex. than in eastern states.
Ponera trigona var. opacior Forel, 1893. Ent. Soc. London, Trans., p. 363. 3, °.

Taxonomy: Taylor, 1968. Ent. News 79: 65.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 272, 274, 277, 304 (Tenn.) —Cole, 1940.
Amer. Midland Nat. 24: 37 (Smoky Mts.). —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 63
(Calif.). —Potts, 1948. Pan-Pacifie Ent. 24: 26 (Calif.). —Gregg, 1963. Ants of Colo., pp.
284-286.

punctatissima (Roger). Fla., Tex., N. Mex., Ariz., Calif.; s. to Central Amer., W. Indies;

Europe, N. Africa. Ecology: Nearly cosmopolitan in warmer parts of world. Possibly
introduced; probably of African origin.

Ponera punctatissima Roger, 1859. Berlin. Ent. Ztschr. 3: 246-248. 3, °.

Ponera androgyna Roger, 1859. Berlin. Ent. Ztschr. 3: 254. Ergatoid 6.

Ponera ergatandria Forel, 1893. Ent. Soc. London, Trans., p. 365. 3, 2, apterous ergatoid
3.

Ponera punctatissima schauinslandi Emery, 1899. Zool. Jahrb., Abt. f. System 12: 439. °.

Taxonomy: Smith, 1936. Ent. Soc. Amer., Ann. 29: 422, 425-426 (female, worker, apterous
ergatoid male). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 28-29 (Polynesia;
synonymy under gleadowi (Forel) listed by Wilson, 1958, Harvard Univ., Mus. Comp. Zool.,
Bul. 119: 328-329 provisionally transferred to punctatissima. This synonymy is not listed
above). —Taylor, 1968. Ent. News 79: 65.

Biology: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 825. —Mallis, 1941. South. Calif.
Acad. Sci., Bul. 40: 63.

Genus LEPTOGENYS Roger

Only a single species of this tropicopolitan genus is known to occur in the United States.

Taxonomy: Wheeler and Wheeler, 1976. Amer. Ent. Soc., Trans. 102: 49 (revised
characterization of larvae).

Genus LEPTOGENYS Subgenus LEPTOGENYS Roger

Leptogenys Roger, 1861. Berlin. Ent. Ztschr. 5: 41.

Type-species: Leptogenys falcigera Roger. Desig. by Bingham, 1903.
Dorylozelus Forel, 1915. Arkiv for Zool. 9: 24-25.

Type-species: Dorylozelus joebergi Forel. Monotypic.
Microbolbos Donisthorpe, 1948. Entomologist 81: 170.

Type-species: Microbolbos testaceus Donisthorpe. Orig. desig.

Not known to occur in the Nearctic Region.

Genus LEPTOGENYS Subgenus LOBOPELTA Mayr

Lobopelta Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 733.
Type-species: Ponera diminuta Smith. Desig. by Bingham, 1903.

1344 Hymenoptera in America North of Mexico

Revision: Wheeler, 1923. Amer. Mus. Novitates 90: 1-16.
elongata elongata (Buckley). La., Tex. Ecology: The small colonies are found in the soil. The

workers apparently forage singly and feed largely, if not exclusively, on pillbugs. There
is no typical female as with most ants, reproduction being carried on by a wingless,
slightly modified worker form with an enlarged gaster.

Ponera Texana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 170. %. A questionable synonym.

Ponera elongata Buckley, 1866. Ent. Soc. Phila., Proc. 6: 172. %.

Lobopelta septentrionalis Mayr, 1866. Zool.-Bot. Gesell. Wien, Verh. 36: 438. 3.

Taxonomy: Wheeler, 1904. Biol. Bul. 6: 257-259 (worker, gynaecoid female, male). —Wheeler
and Wheeler, 1952. Amer. Midland Nat. 48: 641 (larva). —Wheeler, 1900. Biol. Bul. 2: 1-31.

Morphology: Hermann, 1969. Kans. Ent. Soc., Jour. 42: 239-243 (poison apparatus).

elongata manni Wheeler. Fla.
Leptogenys (Lobopelta) elongata manni Wheeler, 1923. Amer. Mus. Novitates 90: 14-15. &.

TRIBE ODONTOMACHINI

Genus ODONTOMACHUS Latreille

Odontomachus Latreille, 1804. Nouv. Dict. Hist. Nat. 24: 179.
Type-species: Formica haematoda Linnaeus. Monotypic.

Four forms of this tropicopolitan genus reach the southern portions of the United States. All
of these have previously been considered as subspecies of the Neotropical O. haematodus (L.).
Most colonies are small and are found in soil or in rotting logs and stumps. Workers are
predaceous and carnivorous. Species of this genus have elongated, linear mandibles and long
hairs which arise between the bases of the mandibles and point forward. These hairs act as trig-
gers when the mandibles are open. When the hairs are touched, the mandibles snap shut result-
ing in a clicking sound. If the mandibles close suddenly on a small object, the object may be cut
in two; if they close suddenly on a large object and the mandibles slide over it, the ant is thrown
in a series of leaps by the force of its closing mandibles.

Revision: Smith, 1939. N. Y. Ent. Soc., Jour. 47: 125-130 (U. S.).

Taxonomy: Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 646 (larvae). —Wheeler and
Wheeler, 1964. Ent. Soc. Amér., Ann. 57: 455-456 (larvae). —Wheeler and Wheeler, 1971.
Ent. Soc. Amer., Ann. 64: 1212 (larvae). —Wheeler and Wheeler, 1976. Amer. Ent. Soc.,
Trans. 102: 61 (revised characterization of larvae).

Biology: Wheeler, 1900. Biol. Bul. 2: 1-31. —Wheeler, 1922. Amer. Mus. Nat. Hist., Bul. 45:
99-103. —Weyer, 1930. Zool. Anz. 90: 49-55 (leaping habits).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116: 475-476
(proventriculus).
clarus Roger. La., Tex.; Mexico, Clarion Is., W. Indies. Ecology: Found in semi-desert regions
where colonies occur in coarse, gravelly soil, fully exposed to the sun.
Odontomachus clarus Roger, 1861. Berlin. Ent. Ztschr. 5: 26. 3.
Odontomachus texana Buckley, 1867. Ent. Soc. Phila., Proc. 6: 355. 8.
Odontomachus haematoda clarionensis Wheeler, 1934. Pan-Pacific Ent. 10: 141. 3.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 407 (worker, female, male).
—Wheeler and Wheeler, 1952. Amer. Midland Nat. 48: 648-650 (larva). —Taylor and
Wilson, 1961. Psyche 68: 142.

Biology: Wheeler, 1900. Biol. Bul. 2: 1-31. —Haskins and Enzmann, 1938. N. Y. Acad. Sci.,
Ann. 37: 100-143.
coninodis Wheeler. Ariz. (Huachuca Mtns.). Ecology: Small colonies are in coarse gravelly soil
under stones at high elevations, usually over 5000 ft.
Odontomachus haematoda coninodis Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 391.
topes

Superfamily FORMICOIDEA 1345

desertorum Wheeler. N. Mex., Ariz.; Mexico. Ecology: Small colonies are in coarse gravelly soil
under stones. Found at lower elevations than coninodis.
Odontomachus haematoda desertorum Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 391.
B.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27: 394.
insularis Guerin. Ga., Fla., Ala.; Mexico to Brazil, W. Indies, Cocos Is., Clipperton Is. Ecology:
Colonies may be in soil or in rotting logs and stumps. Possibly adventive in the U. S.,
distributed by commerce.
Odontomachus insularis Guerin, 1844. Iconogr. Regne Anim. Ins., v. 7, p. 423. %.

Taxonomy: Roger, 1861. Berlin. Ent. Ztschr. 5: 26 (worker, male). —Taylor and Wilson, 1961.
Psyche 68: 142 (distribution).

Biology: Haskins and Enzmann, 1938. N. Y. Acad. Sci., Ann. 37: 149 (formation of new
colonies).

SUBFAMILY PSEUDOMYRMECINAE

A small subfamily with one genus in the New World and several genera in the Old World
tropics. M. R. Smith (1951) named this subfamily Leptaleinae based on the genus Leptalea
Erichson, but later (1952) found an earlier valid generic name, Pseudomyrmex Lund, and
changed the subfamily name to Pseudomyrmecinae which has become widely established.

Taxonomy: Smith, 1951. U. S. Dept. Agr., Agr. Monog. 2: 788 (Leptaleinae). —Smith, 1952.
Ent. Soc. Wash., Proc. 54: 97-98 (Pseudomyrmecinae). —Wheeler and Wheeler, 1973.
Psyche 80: 204-211 (larvae).

Genus PSEUDOMYRMEX Lund

Pseudomyrme Lund, 1831 (June). Ann. Sci. Nat. Zool. 23: 137. Latreille ms.; vernacular.
Pseudomyrmex Lund, 1831 (November). Notizen aus dem Gebiete der Natur und
Heilkunde 32(7): 106. Latreille ms.
Type-species: Formica gracilis Fabricius. Desig. by Smith, 1952.
Leptalea Erichson, 1839. Arch. f. Naturgesch. 5: 309. Klug ms.
Type-species: Formica gracilis Fabricius. Desig. by Wheeler, 1911.
Myrmex Guerin, 1844. Iconogr. Regne Anim., Ins., v. 7, p. 427. Preoce. by Sturm, 1826.
Type-species: Formica (Myrmex) perboscii Guerin. Monotypic.
Pseudomyrma Guerin, 1844. Iconogr. Regne Anim., Ins., v. 7, p. 427. emend.
Leptalaea Spinola, 1851. Accad. Sci. Torino, Mem. 13: 68. Emend.

Over 190 forms of Pseudomyrmex have been described, but only five reach the southern por-
tions of the United States. These ants prefer to nest almost exclusively in preformed plant ecavi-
ties such as twigs and branches of trees, stems of plants, acacia thorns, and in culms of sedges
and grasses. Some species may be restricted to one species of plant or even to one part of the
plant. Among these are the acacia nesting species which nest in the swollen bases of the larger
spines. Janzen (1967) studied a case of obligatory mutualism between a species of Psewdomyr-
mex and Acacia in Mexico and found that the ant is dependent on the acacia for food and the
acacia is dependent on the ant for a substantial part of its life in order to produce seeds and
become part of the reproductive population. The ant colony protects the plant from damaging
factors such as defoliators and thereby assures its own survival on the plant which is its only
food source.

Revision: Mayr, 1870. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 61: 406-413.

Taxonomy: Smith, 1952. Ent. Soc. Wash., Proc. 54: 97-98 (correct generic name). —Creighton,
1955. N. Y. Ent. Soc., Jour. 63: 19-20 (key to workers of U.S.). —Wheeler and Wheeler,
1956. Ent. Soc. Amer., Ann. 49: 376-379 (larvae). —Kempf, 1958. Studia Ent. (n. s.) 1: 434
(gracilis (Fabricius) group).

Biology: Wheeler, 1913. 2nd Internatl. Cong. Ent., Oxford, Trans. 2: 109-139 (observations on

Central American Acacia ants). —Wheeler and Bailey, 1920. Amer. Phil. Soc., Trans. (n. s.)
22: 235-279 (feeding habits). —Brown, 1960. Ecology 41: 589-592 (ants, acacias and

1346 Hymenoptera in America North of Mexico

browsing mammals). —Janzen, 1966. Evolution 20: 249-275 (coevolution of mutualism
between ants and acacias in Central America). —Janzen, 1967. Kans. Univ., Sci. Bul. 48:
315-558 (interaction of the bull’s-horn acacia with an ant inhabitant (Psewdomyrmex
ferruginea Smith) in eastern Mexico). —Janzen, 1967. Ecology 48: 26-35 (fire, vegetation
structure and the ant acacia interaction in Central America). —Janzen, 1969. The Condor
71: 240-256 (the ant acacia interaction and birds in Central America.).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116: 452-453
(proventriculus). —Gotwald, 1969. N. Y. Agr. Expt. Sta (Cornell Univ.), Mem. 408: 128
(mouthparts).

apache Creighton. S. Tex., s. N. Mex., s. Ariz., s. Calif.; n. Mexico. Ecology: Most colonies have
been found in sizeable limbs or trunks of species of Quercus and Prosopis.
Pseudomyrmex apache Creighton, 1952. Psyche 59: 134-139. §, 9, d.

Taxonomy: Wheeler and Wheeler, 1956. Ent. Soc. Amer., Ann. 49: 380 (larva).

Biology: Creighton, 1954. Psyche 61: 9-15. —Creighton, 1963. Amer. Mus. Novitates 2156: 14
(high mortality during cold snap).
brunneus (Smith). N. C. to Fla. w. to Tex. s. to Central Amer.; W. Indies. Ecology: Colonies
have been found in dead twigs of various plants and in culms of grasses and sedges.
Pseudomyrma brunnea Smith, 1877. Ent. Soc. London, Trans., p. 63. %.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 420-421. —Wheeler and Wheeler,
1956. Ent. Soc. Amer., Ann. 49: 382 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 69. —Wheeler, 1932. N. Y. Ent.
Soc., Jour. 40: 3.
elongatus (Mayr). Fla., Tex. s. to S. Amer.; W. Indies. Ecology: Colonies have been found in

twigs of trees and in culms of grasses and sedges.

Pseudomyrma elongata Mayr, 1870. Akad. der Wiss. Wien, Math.-Nat. KI. Sitzber. 61: 408,
413. %.

Pseupomyrma elongata var. cubaensis Forel, 1901. Soc. Ent. Belg., Ann. 45: 342 9°.

Pseudomyrma elongata var. tandem Forel, 1906. Soc. Ent. Belg., Ann. 50: 228. 2.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 85-87 (each caste). —Creighton,
1955. N. Y. Ent. Soc., Jour. 63: 17-20. —Wheeler and Wheeler, 1956. Ent. Soc. Amer., Ann.
49: 384 (larva).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 4.

gracilis mexicanus (Roger). Fla., Tex. s. to Central Amer. Ecology: Colonies have been found
in dead limbs of live oaks, in live oak twig galls, in shrubs, hollow stems of composites,
and cavities in other plants. Probably accidentally introduced into Florida. P. gracilis
gracilis (Fabricius) is found in Central and S. Amer.
Pseudomyrma mexicana Roger, 1863. Berlin. Ent. Ztschr. 7: 178. 8.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 421 (worker, female). —Wheeler
and Wheeler, 1956. Ent. Soc. Amer., Ann. 49: 385-386 (larva). —Brown, 1957. Harvard
Univ., Mus. Comp. Zool., Bul. 116: 235.

Biology: Wheeler, 1901. Soc. Ent. Belg., Ann. 45: 204. —Mitchell and Pierce, 1912. Ent. Soc.
Wash., Proc. 14: 69. —Whitcomb, Denmark, Buren, and Carroll, 1972. Fla. Ent. 55: 31-33
(in Florida).
pallidus (Smith). N. C. to Fla. w. to Ariz., Calif. s. to S. Amer.; W. Indies. Ecology: Colonies
have been found in twigs and branches of various plants and in culms of grasses and
sedges. Some of the early western records of this species may actually refer to apache
Creighton. References to P. flavidulus (Smith) for the U.S. pertain to this species;
flavidulus was described from S. Amer. and its correct status is not known (see
Creighton, 1950).
Pseudomyrma pallida Smith, 1855. Ent. Soc. London, Trans., 3: 160. .

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 419-420. —Wheeler, 1905. Amer.
Mus. Nat. Hist., Bul. 21: 83-85 (each caste). —Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bul. 104: 80-82. —Wheeler and Wheeler, 1956. Ent. Soc. Amer., Ann. 49: 386 (larva).

Superfamily FORMICOIDEA 1347

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 69. —Wheeler, 1932. N. Y. Ent.
Soc., Jour. 40: 4.

Morphology: Blum and Callahan, 1963. Psyche 70: 69-74 (morphology and physiology of
poison glands and venom).

UNPLACED TAXON OF PSEUDOMYRMECINAE
Ponera (Ectatoma(!)) Lincecumii Buckley, 1866. Ent. Soc. Phila., Proc. 6: 172. 3. Cent. Tex.

SUBFAMILY MYRMICINAE

This is the largest subfamily of ants and is found throughout the world. In North America, the
Myrmicinae are better represented in the central and southern United States with their in-
cidence rapidly decreasing northwards where the Formicinae become the dominant subfamily.
Members of this subfamily are recognized by the two-segmented petiole and the frontal carinae

which are distant from each other and each of which usually bears a lobe concealing the antennal
insertions.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Amer., Ann. 53:98-110 (larvae). —Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:1-32 (larvae). —Ettershank, 1966. Austral.
Jour. Zool. 14:73-171 (generic revision of world Myrmicinae related to Solenopsis and
Pheidologeton). —Wheeler and Wheeler, 1973. Psyche 80: 204-211 (supp. studies on
larvae). —Wheeler and Wheeler, 1973. Psyche 80: 70-82 (larvae of four tribes, 2nd supp.;
Leptothoracini, Ocymyrmecini, Tetramoriini, Cryptocerini). —Wheeler and Wheeler, 1973.
Ga. Ent. Soc., Jour. 8: 27-39 (larvae of six tribes, 2nd supp.).

Morphology: Blum, 1974. N. Y. Ent. Soc., Jour. 82: 141-147 (Myrmicine trail pheromones:
specificity, source, and significance). —McCluskey, 1974. N. Y. Ent. Soc., Jour. 82: 93-102
(generic diversity in phase of rhythm in Myrmicine ants).

TRIBE MYRMICINI

Genus MYRMICA Latreille

Myrmica Latreille, 1804. Nouv. Dict. Hist. Nat. 24:179.
Type-species: Formica rubra Linnaeus. Desig. by Latreille, 1810.

This holarctic genus is found as far north as Labrador and Alaska in North America and is
restricted to higher elevations in the southern parts of its range. There are apparently no sub-
tropical or xerophilous representatives. The moderate sized colonies nest in soil, rotten wood, or
under cover of various objects. Workers are carnivorous but also feed on honeydew of Homop-
tera and exudates of plants. Workers of some species differ from each other very slightly and
males are sometimes needed for determination. Some species are closely related to Palearctic
forms and have been regarded as subspecies of them by some authors.

Revision: Weber, 1947. Ent. Soc. Amer., Ann. 40:437-474. —Weber, 1948. Ent. Soc. Amer.,
Ann. 41:267-308. —Weber, 1950. Ent. Soc. Amer., Ann. 43:189-226.

Taxonomy: Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. 5:73-83 (varieties of M. brevinodis).
—Wheeler and Wheeler, 1952. Psyche 59:112-123 (larvae). —Yarrow, 1955. Roy. Ent. Soc.
London, Proc., Ser. B: Taxonomy 24:113-115 (type-species). —Collingwood, 1958. Roy Ent.
Soc. London, Proc., Ser. A 33:65-75 (Britain). —Wheeler and Wheeler, 1963. Ants of N.
Dak., pp. 94-108. —Collingwood, 1974. Soc. Brit. Ent., Trans. 16:96-101 (Britain).

Biology: Brian, 1957. Insectes Sociaux 4:177-190 (growth and development of colonies).
—Weir, 1958. Insectes Sociaux 5:97-128, 316-339 (polyethism in workers). —Weir, 1958.
Jour. Ins. Physiol. 1:352-360 (effect of temperature variation on queen oviposition and
colony formation). —Kannowski, 1959. Insectes Sociaux 6:143-144 (flight activities and
colony founding). —Weir, 1959. Insectes Sociaux 6:271-290 (influence of worker age on
trophogenic larval dormancy). —Weir, 1959. Insectes Sociaux 6:167-201 (egg masses and
early larval growth). —Weir, 1959. Physiol. Zool. 32:63-77 (interrelation of queen and
worker oviposition). —Carr, 1962. Insectes Sociaux 9:177-211 (influence of queen). —Brian
and Hibble, 1963. Insectes Sociaux 10:71-82 (larval size and influence of queen on growth).

1348 Hymenoptera in America North of Mexico

—Kannowski, 1970. Ent. Soe. Amer., N. Central Branch, Proc. 25:119-125 (colony
populations of 5 species).

Morphology: Weir, 1957. Quart. Jour. Micros. Sci. 98:499-506 (functional anatomy of mid-gut
of larvae). —Weir, 1959. Insectes Sociaux 6:375-386 (changes in the rectro-cerebral
endocrine system of larvae and their relation to larval growth and development). —Crewe
and Blum, 1970. Ztschr. f. Vergleich. Physiol. 70:363-373 (alarm pheromones of 9 species).

americana Weber. Que., Maine s. to N. C., Tenn., w. to Man., Colo., Utah, Ariz. Ecology: More
common on east slopes of Rocky Mtns.; records from west of the Rockies are rare.
Commonly found in grasslands where nests are in soil in open or under objects; food is
varied, consisting of animal matter and plant juices. Sometimes considered as a
subspecies of the Palearctic Myrmica sabuleti Meinert.
Myrmica sabuleti americana Weber, 1939. Lloydia 2:144. 3, 2, d.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:116 (larva).

Biology: Wheeler, 1916. Conn. State Geol. and Nat. Hist. Survey Bul. 22:587. —Buren, 1944.
Iowa State Col., Jour. Sci. 18:282-283. —Kannowski, 1956. Amer. Midland Nat. 56:175.
—Kannowski and Kannowski, 1957. Ohio Jour. Sci. 57:371-374 (mating activities).
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 96-99. —Gregg, 1963. Ants of Colo., pp.
308-310. —Ayre, 1963. Canad. Ent. 95:712-715 (feeding habits). —Burns, 1964. Ent. Soc.
Amer., Ann. 57:138 (association with tuliptree scale). —Ayre, 1968. Canad. Ent. 100:165-172
(prey finding, capture, and transport). —Ayre, 1969. Canad. Ent. 101:118-128 (trail
formation and group foraging). —Ayre, 1971. Ztschr. f. Angew. Ent. 68:295-299 (foraging
and nesting habits).

brevispinosa brevispinosa Wheeler. N. Dak., Nebr. s. to Colo., N. Mex., w. to Alta., Idaho.
Ecology: Prefers to nest in stream valleys or on shores of permanent bodies of water.
Myrmica rubra brevinodis var. brevispinosa Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. 5:74.
BRnOErSe
Myrmica rubra brevinodis var. decedens Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. 5:75. 8,
3.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:117 (larva). —Kannowski, 1956. Amer.
Midland Nat. 56:176. —Gregg, 1961. N. Y. Ent. Soc., Jour. 69:211.

Biology: Weber, 1942. Canad. Ent. 74:62. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart.
11:241. —Gregg, 1963. Ants of Colo., pp. 296-297, 299. —Wheeler and Wheeler, 1963. Ants
of N. Dak., pp. 101-103 (subspecies not recognized).

brevispinosa discontinua Weber. Newfoundland, N.S. w. to mtns. of Wyo., Colo., N. Mex.
Ecology: Apparently more tolerant than the typical subspecies of higher altitudes in
Rockies where the two subspecies overlap.
Myrmica brevinodis discontinua Weber, 1939. Lloydia 2:150. 3.

Biology: Cole, 1953. Tenn. Acad. Sci., Jour. 28:243 (N. Mex.). —Gregg, 1963. Ants of Colo., pp.
298-300.
emeryana emeryana Forel. Newfoundland s. to Ga., w. to Man., Idaho, Colo., Ariz. Ecology:

Nests are usually in woodlands in moist, shady situations under stones or other objects.
Sometimes considered as a subspecies of the Palearctic M. schencki Emery.

Myrmica scabrinodis schencki var. emeryana Forel, 1914. Deut. Ent. Ztschr., p. 617. 3, 2,
é.

Myrmica schencki latifrons Starcke, 1927. Tijdschr. v. Ent. 70:84. &.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:114, 117 (larva). —Smith, 1954. Brooklyn
Ent. Soc., Bul. 49:138-140 %, 2 (worker, female; type - locality). —Wheeler and Wheeler,
1972. Ga. Ent. Soc., Jour. 7:233 (larva).

Biology: Wheeler, 1915. Psyche 22:206. —Talbot, 1945. Ent. Soc. Amer., Ann. 38:365-372
(population studies). —Kannowski, 1959. Insectes Sociaux 6:121. —Medler, 1958. Ent. Soc.
Wash., Proc. 60:258 (swarming). —Peterson and Davies, 1960. Canad. Jour. Zool. 38:15
(predation on Simuliwm; behavior). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
103-104. —Gregg, 1963. Ants of Colo., pp. 311-312, 314.

Superfamily FORMICOIDEA 1349

emeryana tahoensis Wheeler. Mont., Wyo., Utah, Ariz. w. to B. C., Oreg., Nev., Calif. Ecology:
Mostly in mountains at higher elevations.
Myrmica scabrinodis schencki var. tahoensis Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:504. §, 2, d.
Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7 (3):3-4 (Nevada Test Site).
hamulata hamulata Weber. Colo., N. Mex., Utah, Ariz. Ecology: Apparently prefers to nest in

upland plateaus from 7,000 to 8,000 ft.
Myrmica sabuleti hamulata Weber, 1939. Lloydia 2:146. 8, 2, d.

Biology: Gregg, 1963. Ants of Colo., pp. 300-301.
hamulata trullicornis Buren. Iowa. The exact status of this form will remain uncertain until a
male can be associated with the workers.
Myrmica sabuleti trullicornis Buren, 1944. Iowa State Col., Jour. Sci. 18:281. 3, 9.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool. Bul. 104:99-100.

incompleta incompleta Provancher. Labrador s. to N. J., w. to Rocky Mtns., Colo., Utah, N.
Mex. Ecology: Widely distributed with wide elevational tolerance. Prefers moist, grassy
habitats where it usually nests under objects. Host of the inquiline Leptothorax
provancheri Emery.

Myrmica incompleta Provancher, 1881. Nat. Canad. 12:359. ¥, 2, d.

Myrmica rubra brevinodis Emery, 1895. Zool. Jahrb., Abt. f. System 8:312. 8, d.

Myrmica rubra brevinodis var. canadensis Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. 5:76.
Hm) Ney

Myrmica rubra brevinodis var. subalpina Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. 5:77. 8,
Qe

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:116 (larva). —Francoeur and Beique, 1966.
Canad. Ent. 98:141 (Provancher types).

Biology: Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. 5:73, 77-83. —Wheeler, 1916. Conn. State
Geol. and Nat. Hist. Survey Bul. 22:587. —Kannowski, 1959. Insectes Sociaux 6:121, 155
(pleometrosis). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 99-101. —Gregg, 1963.
Ants of Colo., pp. 291-292, 294.

incompleta kuschei Wheeler. Alaska.
Myrmica brevinodis var. kuschei Wheeler, 1917. Harvard Univ., Mus. Comp. Zool., Bul.

61:17. 3, @.
Myrmica brevinodis var. alaskensis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.
52:503. 9.
incompleta sulcinodoides Emery. S. Dak., Colo., N. Mex. w. to Alta., Alaska, B. C., Oreg.,
Calif. Some authors do not consider this a valid subspecies.
Myrmica rubra brevinodis var. sulcinodoides Emery, 1895. Zool. Jahrb., Abt. f. System.
8:313. 8.
Myrmica rubra brevinodis var. frigida Forel, 1902. Ent. Soc. London, Trans., p. 699. 9.
Myrmica rubra brevinodis var. whymperi Forel, 1904. Soc. Ent. Belg., Ann. 48:154. 9.
Biology: Wheeler, 1915. Psyche 22:206. —Cole, 1934. Psyche 41:223. —Gregg, 1963. Ants of
Colo., pp. 293-295, 297.
lampra Francoeur. Que. (Pare des Laurentides). Ecology: Apparently parasitic, taken from
nest of another Myrmica. Host: Myrmica sp.
Myrmica lampra Francoeur, 1968. Nat. Canad. 95:729. 2, 3.
lobicornis fracticornis Emery. Newfoundland s. to Tenn., Ohio w. to Rocky Mtns. Colo., N.
Mex., Utah, Ariz. Ecology: Nests are usually near a stream or slough, under stones or
wood. Host of the inquiline Leptothorax provancheri Emery. M. lobicornis lobicornis

Nylander is Palearctic.
Myrmica rubra scabrinodis var. fracticornis Emery, 1895. Zool. Jahrb., Abt. f. System.

8:313. &.
Myrmica rubra scabrinodis var. detrinodis Emery, 1895. Zool. Jahrb., Abt. f. System.

8:316. 3. A questionable synonym.

1350 Hymenoptera in America North of Mexico

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:117 (larva). —Cole, 1953. Tenn. Acad. Sci.,
Jour. 28:242-243.

Biology: Wheeler, 1916. Conn. State Geol. and Nat. Hist. Survey Bul. 22:587-588. —Eidmann,
1933. Zool. Anz. 101:203. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:288, 305. —Weber, 1941.
Canad. Ent. 73:140-141 (effect of drouth on nesting habits in prairie states and provinces).
—Weber, 1942. Canad. Ent. 74: 62. —Kannowski, 1957. Psyche 64: 1-5 (host of Leptothorax
provancheri Emery). —Kannowski, 1959. Insectes Sociaux 6:121-124, 155-156 (flight
activities, colony founding). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 104-106.
—Gregg, 1963. Ants of Colo., pp. 300, 302-303.

lobicornis lobifrons Pergande. Colo., N. Mex., Utah, Ariz., n. w. to Alaska. Ecology: In
mountains at high elevations. Some authors consider this a synonum of fracticornis.

Myrmica sabuleti var. lobifrons Pergande, 1900. Wash. Acad. Sci., Proc. 2: 521. 9.
Myrmica rubra scabrinodis var. glacialis Forel, 1904. Soc. Ent. Belg., Ann. 48: 154. &.

Biology: Gregg, 1963. Ants of Colo., pp. 303-307.

mexicana Wheeler. Ariz., Mexico.
Myrmica mexicana Wheeler, 1914. N. Y. Ent. Soc., Jour. 22: 52. 3, 2, d.

Taxonomy: Gregg, 1961. N. Y. Ent. Soc., Jour. 69: 211.
monticola Wheeler. Que., Mich. w. to Man., N. Dak., Colo. Ecology: Nests in woodlands under
cover of objects.
Myrmica scabrinodis schencki var. monticola Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proce. 52:505. %, d.
Myrmica sabuleti nearctica Weber, 1939. Lloydia 2: 148. 3, @, d.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:117 (larva).

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 243. —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 106-108. —Gregg, 1963. Ants of Colo., pp. 306-309. —Francoeur,
1966. Nat. Canad. 93: 455 (Que.).

pinetorum Wheeler. Mass. s. to S. C., w. to Ohio, Okla., Miss. Ecology: Prefers to nest in sandy
soil.
Myrmica punctiventris pinetorum Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:384. §,
oe

Biology: Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17:10. —Wesson and Wesson,
1940. Amer. Midland Nat. 24:90, 94.
punctiventris Roger. Mass. s. to Ga., w. to Iowa, Nebr., Ark. Ecology: Nests have been found
in soil and rotten logs in woodlands.
Myrmica punctiventris Roger, 1863. Berlin Ent. Ztschr. 7:190. §.
Myrmica punctiventris var. isafahani Forel, 1922. Rev. Suisse de Zool. 30:92. 3, 2.

Taxonomy: Gregg, 1961. N. Y. Ent. Soc., Jour. 69:211.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:383-384. —Cole, 1940. Amer. Midland
Nat. 24:55. —Headley, 1943. Ohio Jour. Sci. 43:25-26. —Buren, 1944. Iowa State Col., Jour.
Sci. 18:283. —Kannowski, 1959. Insectes Sociaux 6:124.
rubra (Linnaeus). Que., Maine, Mass., R. I.; Europe. Ecology: Unlike other species of Myrmica,

rubra is pugnacious and can inflict a painful sting. Probably accidentally introduced
from Europe.

Formica rubra Linnaeus, 1758. Syst. Nat., Ed. 10 1:580.

Myrmica laevinodis Nylander, 1846. Acta Soc. Fenn. 2:927. 8, @, d.

Myrmica rubra champlaini Forel, 1901. Naturhist. Mus. Hamburg Mitt. 18:80. 9.

Myrmica rubra laevinodis var. bruesi Wheeler, 1906. Psyche 13:38. 3, 2, d.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 58:119 (larva). —Picquet, 1958. Dijon Univ.
Lab. de Zool. Trav. 23:28 (larva). —Yarrow, 1955. Roy. Ent. Soc. London, Proc., Ser. B:
Taxonomy. 24:113-115. —Wheeler and Wheeler, 1960. Ent. Soc. Amer., Ann. 53:4-5 (larva).

Biology: Wheeler, 1908. Jour. Econ. Ent. 1:337-339 (introduced into Mass.). —Sturtevant,
1931. Psyche 38:75. —Brian, 1951. Experientia 7:182 (caste determination). —Brian, 1951.

Superfamily FORMICOIDEA 1351

Physiol. Comp. and Oecol. 2:248-262. —Brian, 1953. Physiol. Comp. and Oecol. 3:25-36
(oviposition by workers). —Brian, 1954. Insectes Sociaux 1:101-122 (caste differentiation,
larval dormancy, winter size, vernalization). —Brian, 1955. Insectes Sociaux 2:1-34 (caste
differentiation, growth of workers and intercastes). —Brian, 1955. Insectes Sociaux
2:85-114 (caste differentiation, larval dormancy, winter size, and vernalization). —Brian
and Brian, 1955. Evolution 9: 280-290 (macrogynes and microgynes). —Brian, 1956.
Insectes Sociaux 3:369-394 (caste differentiation, controlled larval nutrition). —Brian, 1956.
Jour. Anim. Ecol. 25:319-337 (segregation of Myrmica spp.). —Brian, 1957. Insectes
Sociaux 4:191-210 (serial organization of brood). —Brian, 1957. Physiol. Comp. and Oecol.
4:329-345 (food distribution and larval size in cultures). —Weir, 1958. Insectes Sociaux
5:315-339 (polyethism in workers). —Brian, 1962. Insectes Sociaux 9:295-310 (social
conditions affecting early larval differentiation). —Brian, 1963. Insectes Sociaux 10:91-102
(caste differentiation). —Plateaux, 1960. Insectes Sociaux 7: 221. —Brian, 1965. Insectes
Sociaux 12:347 (caste differentiation). —Brian, et al., 1967. Insectes Sociaux 14:13-24
(caste differentiation). —Brian, 1969. Insectes Sociaux 16:249-268 (ecological notes).
—Jaisson, 1969. Insectes Sociaux 16:279-312. —Brian, 1970. Animal Behavior 18:467-472
(communication between queens and larvae).

Morphology: Tulloch, 1936. Ent. Soc. Amer., Ann. 29:81-84 (metasternal glands). —Allen,
1957. Ent. Monthly Mag. 93:136-139 (intercastes). —Weir, 1959. Insectes Sociaux 6:375-386
(rectro-cerebral endocrine glands and their relation to larval growth and development).

spatulata Smith. Tenn., Miss., III.
Myrmica schencki var. spatulata Smith, 1930. Ent. Soc. Amer., Ann. 23:566. 3, @.

Biology: Smith, 1931. Ent. News 42:21.

striolagaster Cole. Tex., Colo., N. Mex., Ariz.
Myrmica striolagaster Cole, 1953. Tenn. Acad. Sci., Jour. 28:81. %.

Taxonomy: Cole, 1957. Tenn. Acad. Sci., Jour. 32:208-209 (male). —Gregg, 1961. N.Y. Ent. Soc.,
Jour. 69:211. —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:234 (larva).
Biology: Gregg, 1963. Ants of Colo., pp. 313-314.
wheeleri Weber. Ariz. (Mt. Lemmon and Mt. Stratton in Santa Catalina Mtns.).
Myrmica wheeleri Weber, 1939. Lloydia 2:150. %, 2, 3.

UNPLACED TAXON OF MYRMICA

rubra neolaevinodis Forel. “From New York with iris roots”. The iris from which the ants
were taken may not have come from New York; consequently, this form may not belong
in the Nearctic fauna.
Myrmica rubra neolaevinodis Forel, 1901. Naturhist. Mus. Hamburg Mitt. 18: 80. %.

Genus PARAMYRMICA Cole

Paramyrmica Cole, 1957. Tenn. Acad. Sci., Jour. 32:37-42.
Type-species: Paramyrmica colax Cole. Monotypic.

Little is known of the habits of the two species included in this genus. The genus is close to
Myrmica, but differences in the larvae support the distinctness of Paramyrmica.
Taxonomy: Gregg, 1961. N. Y. Ent. Soc., Jour. 69:209-220.
colax Cole. Tex. (Limpia Canyon, Davis Mtns., Jeff Davis Co.). Ecology: An inquiline. Host:
Myrmica striolagaster Cole.
Paramyrmica colax Cole, 1957. Tenn. Acad. Sci., Jour. 32:37-41. %, °.
Taxonomy: Wheeler and Wheeler, 1959. Tenn. Acad. Sci., Jour. 34:291-220 (larva).
rugiventris (Smith). Colo., Ariz., s. Calif.
Tetramorium rugiventris Smith, 1943. Ent. Soc. Wash., Proc. 45:2, 4. 9.
Taxonomy: Brown, 1957. Breviora 72:5-7 (an endemic species; in Myrmica). —Gregg, 1961.
N. Y. Ent. Soc., Jour. 69:209 (in Paramyrmica). —Francoeur, 1968. Nat. Canad. 95:728
(probably in Myrmica).

1352 Hymenoptera in America North of Mexico

Genus MANICA Jurine

Manica Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 276.

Type-species: Formica rubida Latreille. Desig. by Wheeler, 1911.
Aphaenogaster subg. Neomyrma Forel, 1914. Rev. Suisse Zool. 22:275.

Type-species: Aphaenogaster (Neomyrma) calderoni Forel. Monotypic.
Myrmica subg. Oreomyrma Wheeler, 1914. Psyche 21:118.

Type-species: Formica rubida Latreille. Orig. desig.

Four of the five species in this genus are found in western North America, west of the hun-
dredth meridian; the other is Palearctic. Colonies are small and nests are usually found in
openings in coniferous forests commonly under stones in creek or river bottoms. For an excel-
lent account of each species, see Wheeler and Wheeler (1970).

Revision: Wheeler, 1914. Psyche 21:118-122. —Wheeler and Wheeler, 1970. Kans. Ent. Soc.,
Jour. 43:129-162.

Taxonomy: Weber, 1947. Ent. Soc. Amer., Ann. 40:439. —Wheeler and Wheeler, 1952. Psyche
59:123 (larvae). —Cole, 1957. Tenn. Acad. Sci., Jour. 32:213 (key to males). —Wheeler and
Wheeler, 1960. Ent. Soc. Wash., Proc. 62:15 (larva). —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 108-110. —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:234-236 (larvae).

Biology: Wheeler and Wheeler, 1970. Kans. Ent. Soc., Jour. 43:129-162. —Wheeler and
Wheeler, 1970. Kans. Ent. Soc., Jour. 43:363 (additions to natural history of Manica).
—Went, Wheeler, and Wheeler, 1972. BioScience 22:82-88 (feeding and digestion).

Morphology: Fales, et al., 1972. Jour. Ins. Physiol. 18: 1077-1088 (alarm pheromones derived
from the mandibular galnd).
bradleyi (Wheeler). W. Nev., Calif. Ecology: Typically found in openings in coniferous forests
in the Sierra Nevada Mtns. of w. Nev. and Calif., and the Transverse Ranges in s. Calif.
Host of Manica parasitica (Creighton), but the exact relationship is not known.
Myrmica bradleyi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:77. 8.
Aphaenogaster (Neomyrma) calderoni Forel, 1914. Rev. Suisse de Zool. 22:275. 9.

Taxonomy: Cole, 1957. Tenn. Acad. Sci., Jour. 32:210-212 (female, male). —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62:5-6 (larva). —Wheeler and Wheeler, 1972. Ga. Ent. Soc.,
Jour. 7:234-235 (larva).

Biology: Creighton, 1934. Psyche 41:188-189. —Mallis, 1941. South. Calif. Acad. Sci., Bul.
40:65.
hunteri (Wheeler). Alta., Mont., Wyo., Idaho, Utah, B. C., Wash., Oreg., Nev., Calif. Ecology:
Typically found in openings in coniferous forests.
Myrmica (Oreomyrma) hunteri Wheeler, 1914. Psyche 21:119, 121. &.
Myrmica (Oreomyrma) aldrichi Wheeler, 1914. Psyche 21:119, 120. %.

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31:262. —Cole, 1957. Tenn. Acad. Sci., Jour.
32:212-213 (male). —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:235 (larva).
mutica (Emery). N. Dak., S. Dak., Colo., N. Mex. w. to Alaska, B. C., Wash., Oreg., Calif.
Ecology: The most widely distributed ant of the genus in N. Amer. It is more
xerophilous and occurs in a greater variety of habitats. Host of the inquiline
Symmyrmica chamberlini Wheeler.
Myrmica mutica Emery, 1895. Zool. Jahrb., Abt. f. System. 8:311. 9.

Taxonomy: Cole, 1957. Tenn. Acad. Sci., Jour. 32: 213 (male). —Wheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:5-6 (larva). —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour.
7:235 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:6-7. —Wheeler, 1910. Ants, pp.
432-434. —Cole, 1953. Tenn. Acad. Sci., Jour. 28:243. —Gregg, 1963. Ants of Colo., pp.
314-317. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 109-110.

parasitica (Creighton). Calif. (Alpine Co.; Yosemite Nat. Pk., Mariposa Co.). Ecology:
Collections made from ants nests. Host: Manica bradleyi (Wheeler).
Myrmica (Manica) parasitica Creighton, 1934. Psyche 41:185. %.

Superfamily FORMICOIDEA 1353

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:66. —Wheeler and Wheeler, 1968.
Pan-Pacific Ent. 44:71-72 (rediscovery).

Genus POGONOMYRMEX Mayr

Members of this genus are collectively known as harvester ants and some species are among
the most conspicuous ants in the arid regions of the western United States and Mexico because
of their mound building habits. The workers collect seeds for food, harvesting plants in their
nesting areas by snipping off seeds with their mandibles. The seeds are stored and are the main
food source though the ants are also scavengers. Nests are in the soil in areas fully exposed to
the sun; some are under stones and others are surmounted by soil craters or by small to huge
mounds with or without coverings of gravel. Some species alter the area surrounding their nest
by clearing away the vegetation. Some species have a painful sting, procure seeds from cul-
tivated crops and damage rangelands.

Revision: Wheeler, 1902. Amer. Nat. 36:85-100. —Wheeler, 1902. Psyche 9:387-393.
—Wheeler, 1914. Psyche 21:151-157. —Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bul.
77:493-514. —Cole. 1968. Pogonomyrmex Harvester Ants, 222 pp.

Taxonomy: Wheeler and Wheeler, 1952. Psyche 59:106-111 (larvae). —Cole, 1954. Tenn. Acad.
Sci., Jour. 29:117-119. —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:2 (larva).
—Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:236-237 (larvae).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116:477-478
(proventriculus). —McGurk, et al., 1966. Jour. Ins. Physiol. 12:1435-1441 (identification of
4-methyl-3-heptanone).

Genus POGONOMYRMEX Subgenus POGONOMYRMEX Mayr

Pogonomyrmex Mayr, 1868. Soc. Nat. Modena, Ann. 3:169.
Type-species: Formica badia Latreille. Desig. by Wheeler, 1911.

SPECIES GROUP BARBATUS

anergismus Cole. N. Mex. (15 mi. e. Silver City, 6900 ft.). Ecology: Possibly a social parasite;
found in nest of P. rugosus. Host: Pogonomyrmex rugosus Emery.
Pogonomyrmex (Pogonomyrmex) anergismus Cole, 1954. iene Acad. Sci., Jour.
29:115-116. 2, d.
apache Wheeler. Southwest. Kans. (?), w. Tex., s. Colo., N. Mex., s. Ariz., s. Nev.; Mexico.
Ecology: Colonies are small and the usually obscure nests are in stony soils with no
superstructure though sometimes in sandy soil with small circular craters.
Pogonomyrmex apache Wheeler, 1902. Psyche 9:392. &.
Pogonomyrmex sancti-hyacinthi Wheeler, 1902. Psyche 9:388. &.

Biology: Wheeler, 1910. Ants, p. 283. —Cole, 1954. Tenn. Acad. Sci., Jour. 29:266-267.
—Gregg, 1963. Ants of Colo., pp. 318-319.
barbatus (Smith). E. Ark., e. La., s. and w. Kans., Okla., Tex., e. Colo., s. and central N. Mex.,

s.e. and central Ariz., Nev. (Clark Co.); Mexico. Ecology: The usual nest is a low to high
gravel mound, frequently with a scooped-out center, though the nest structure varies.
The most conspicuous ant of this genus in its range. Red harvester ant.

Myrmica barbata Smith, 1858. Cat. Hym. Brit. Mus. 6:130. 2.

Myrmica (Atta) molefaciens Buckley, 1860. Acad. Nat. Sci. Phila., Proc. 12:445. 9, °.

Pogonomyrmex barbatus var. nigrescens Wheeler, 1902. Psyche 9:389, 391. 9.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:308 (each caste). —Wheeler, 1914. N.
Y. Ent. Soc., Jour. 22:51-52 (each caste). —Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bul. 104:116-119. —Wheeler and Wheeler, 1952. Psyche 59:107 (larva). —Wheeler
and Wheeler, 1960. Ent. Soe. Wash., Proc. 62:2 (larva).

Biology: McCook, 1879. The Natural History of the Agricultural Ant of Texas, 311 pp.
—Wheeler, 1901. Soc. Ent. de Belg., Ann. 45:202. —Wheeler, 1901. Amer. Nat. 35:723-724.
—Wheeler, 1910. Ants, pp. 11, 85, 179, 197, 202-203, 222, 264, 284, 286-288, 290-293.
—Hunter, 1921. U. S. Dept. Agr. Bur. Ent. Cir. 148:4-7. —Gordon, 1943. Jour. Econ. Ent.

1354 Hymenoptera in America North of Mexico

36:354 (damage to airplane runways). —Michener, 1948. N. Y. Ent. Soc., Jour. 56:239-242.
—Cassidy, et al., 1950. U. S. Dept. Agr. Cir. 842:15. —Barnes and Nerney, 1953. U. S.
Dept. Agr. Farmers’ Bul. 1668:1-11. —Young and Howell, 1954. Okla. Acad. Sci., Proc.
35:60-62 (mating swarms). — Young and Gonzalez, 1957. Mex. Secretaria de Agr. y
Ganaderia, Fol. Tec. 23:1-20. —Michener, 1960. Kans. Ent. Soc., Jour. 33:46 (treetop mating
aggregations). —Box, 1960. Ecology 41:381-382. —Gregg, 1963. Ants of Colo., pp. 319-321.
—Johnson, et al., 1969. Biochemical Genetics 3:429-450 (isozyme genotype-environment
relationships in natural populations).

Morphology: Williams and Williams, 1965. Soc. Exp. Biol. Med., Proc. 119:344-346 (venom).
—Benthuysen and Blum, 1974. Ga. Ent. Soc., Jour. 9: 235-238 (quantitative sensitivity of
ant to enantiomers of its alarm pheromone).

bicolor Cole. S. Ariz.; Mexico. Ecology: Nests are in gravelly soil with a low mound of sand
and with a single entrance. Apparently does not clear plants from nest periphery.
Pogonomyrmesx (Pogonomyrmex) bicolor Cole, 1968. Pogonomyrmex Harvester Ants, pp.
59-64. 3, 9, d.
desertorum Wheeler. W. Tex., s. N. Mex., s. Ariz., s.e. Calif.; Mexico. Ecology: Nests are in
sandy soil in open areas and have circular craters.
Pogonomyrmex desertorum Wheeler, 1902: Psyche 9:387, 390. &.
Pogonomyrmesx desertorum var. ferrugineus Olsen, 1934. Harvard Univ., Mus. Comp.
Zool., Bul. 77:496. %.

Taxonomy: Cole, 1954. Tenn. Acad. Sci., Jour. 29:119.

Biology: Wheeler, 1910. Ants, p. 283. —Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Mallis,
1941. South. Calif. Acad. Sci., Bul. 40:64. —Whitford and Ettershank, 1975. Environ. Ent.
4: 689-696 (factors affecting foraging activity).

rugosus Emery. W. Okla., w. Tex., s.e. and s.w. Colo., N. Mex., s. Utah, Ariz., s. and central

Nev., s. Calif.; Mexico. Ecology: Typically, the nest is a flattened gravel disc, though
there is variation in structure.

Pogonomyrmesx barbatus rugosus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:309. 3, d.

Pogonomyrmex barbatus var. fuscatus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:309. 9.

Pogonomyrmex barbatus var. marfensis Wheeler, 1902. Amer. Nat. 36:98. 8.

Pogonomyrmex similis Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bul. 77:497, 512. &.

Pogonomyrmex barbatus curvispinosus Cole, 1936. Ent. News 47:120. &.

Pogonomyrmex barbatus spadix Cook, 1953. The Ants of Calif., pp. 98-99, 3 figs. 3. No
description.

Taxonomy: Wheeler, 1902. Psyche 9:391, 393 (each caste).

Biology: Wheeler, 1902. Psyche 9:393. —Wheeler, 1910. Ants, pp. 284, 290. —Wheeler, 1917.
Psyche 24:178. —Cole, 1937. Ent. News 48:134. —Cole, 1942. Amer. Midland Nat. 28:367.
—Lindquist, 1942. Jour. Econ. Ent. 35:850-853. —Gregg, 1963. Ants of Colo., pp. 322-324,
325, 327. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 50. —Whitford and
Ettershank, 1975. Environ. Ent. 4: 689-696 (factors affecting foraging activity).

tenuispina Forel. S. Calif. (Deep Canyon); Mexico (Baja Calif.).
Pogonomyrmex desertorum var. tenuispina Forel, 1914. Soc. Vaud. des Sci. Nat., Bul.
50:269. 8.
Pogonomyrmex dentatus Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bul. 77:505. &.

Taxonomy: Cole, 1968. Pogonomyrmex Harvester Ants, pp. 77-81. —Wheeler and Wheeler,
1973. Ants of Deep Canyon, pp. 50-51 (first U. S. record).

SPECIES GROUP OCCIDENTALIS

anzensis Cole. Calif. (Split Mtn., Anza Desert State Park).
Pogonomyrmex (Pogonomyrmex) anzensis Cole, 1968. Pogonomyrmex Harvester Ants,
pp. 87-89. &.
brevispinosus Cole. S. Calif., Nev. (Lyon Co.). Ecology: Nests are in compact sandy soil, each
marked by a low circular crater 3 to 8 inches in diameter with a single entrance.
Pogonomyrmex (Pogonomyrmex) brevispinosus Cole, 1968. Pogonomyrmex Harvester
Ants, pp. 89-94. %, 9, d.

Superfamily FORMICOIDEA 1355

occidentalis (Cresson). Southwest. N. Dak., w. S. Dak., w. Nebr., central and w. Kans., central

and w. Okla., n. Tex., s.e. Mont., Wyo., Colo., N. Mex., s.e. Idaho, Utah, e. and n. Ariz.,
Nev., e. Calif. Ecology: The populous colonies build conical pebble mounds with basal
entrances and peripheral clearings. Workers are pugnacious and are active harvesters,
feeding on and storing large quantities of seeds. Western harvester ant.

Myrmica occidentalis Cresson, 1895. Ent. Soc. Phila., Proc. 4:426. 3, °.

Myrmica seminigra Cresson, 1865. Ent. Soc. Phila., Proc. 4:427. 3.

Pogonomyrmex opaciceps Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:971. 8.

Pogonomyrmex occidentalis ruthveni Gaige, 1914. Biol. Soc. Wash., Proce. 27:93. %, 2, 3.

Pogonomyrmex occidentalis var. utahensis Olsen, 1934. Harvard Univ., Mus. Comp. Zool.,
Bul. 77:498, 509. %, 2, d.

Taxonomy: Smith, 1953. Brooklyn Ent. Soc., Bul. 48:131. —Wheeler and Wheeler, 1952.
Psyche 59:110 (larva).

Biology: McCook, 1882. The Honey Ants of the Garden of the Gods, and the Occident Ants
of the American Plains, pp. 123-160. —Dean, 1905. Kans. Acad. Sci., Trans. 19:164-170.
—Headlee and Dean, 1908. Kans. State Col., Bul. 154:165-180. —Wheeler, 1910. Ants, pp.
145, 200, 202-205, 222, 283-284, 290, 291, 426. —Herrick, 1914. Insects Injurious to the
Household and Annoying to Man, pp. 172-173. —Cole, 1934. Ent. News 45:170. —Cole,
1942. Amer. Midland Nat. 28:366. —Knowlton, 1951. Brooklyn Ent. Soc., Bul. 46:75 (loss of
surface area on western rangeland). —Weber, 1959. Ent. News 70:85 (stings; note on
populations). —Chew, 1960. N. Y. Ent. Soc., Jour. 68:81 (colony size and activity).
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 118-120. —Gregg, 1963. Ants of Colo.,
pp. 328-334 (as occidentalis comanche). —Spangler, 1968. Kans. Ent. Soc., Jour. 41:318-323
(behavior). —Lavigne, 1969. Ent. Soc. Amer., Ann. 62:1166-1175 (bionomics and nest
structure). —Clark and Camanor, 1973. Biol. Soc. Nev., Occ. Papers 34:1-6 (use of seed
stores by Heteromyid rodents). —Lavigne and Rogers, 1974. Wyo. Univ. Agr. Exp. Sta.
Sci. Monog. 26: 1-14 (annotated bibliography).

owyheei Cole. Sask., central and w. Mont., n.w. Wyo., Alta., Idaho, n.w. Utah, B. C., Wash.,
Oreg., n. Nev., n.e. Calif. Ecology: Nest structure and habits are similar to those of P.
occidentalis.
Pogonomyrmex occidentalis owyheei Cole, 1938. Amer. Midland Nat. 19:240. %, 9.

Biology: Cole, 1932. Ohio Jour. Sci. 32:17-20, 133-146, 245-246, 533-538 (as occidentalis).
—Cole, 1933. Ent. News 44:16-19 (as occidentalis). —Cole, 1934. Canad. Ent. 66:193-198
(as occidentalis). —Cole, 1934. Ent. News 45:96-101 (as occidentalis). —Willard and
Crowell, 1965. Jour. Econ. Ent. 58:484-489 (biology; e. Oreg.). —Lavigne and Rogers, 1974.
Wyo. Univ., Agr. Exp. Sta. Sci. Monog. 26: 14-18 (annotated bibliography).

salinus Olsen. Southeast. Oreg., Nev., e. Calif. Ecology: Found mostly in pinyon-juniper areas;

nests are a low bed of gravel with a cleared area surmounted by one or more craterlike
depressions each with one entrance. A docile ant.

Pogonomyrmex salinus Olsen, 1934. Harvard Univ., Mus. Comp. Zool., Bul. 77:498, 510. &.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soe., Jour. 7:237 (larva).
subdentatus Mayr. Southwest. Oreg., w. Nev., Calif. Ecology: A timid, inoffensive ant whose
nests are low, irregular beds of sand and gravel with more than one entrance. Mayr’s

citation of this form from Conn. is an error.
Pogonomyrmex subdentatus Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:971. &.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Mallis, 1941. South. Calif. Acad. Sci., Bul.
40:65.
subnitidus Emery. Nev. (?) (Nye Co., Lyon Co.). coastal central and s. Calif.; Mexico. Ecology:
Nests are in sand or loose, sandy soil surmounted by semicircular or circular craters or
marked by irregular beds of sand.
Pogonomyrmesx occidentalis var. subnitidus Emery, 1895. Zool. Jahrb., Abt. f. System.
8:310. 8.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Mallis, 1941. South. Calif. Acad. Sci., Bul.
40:65. —Wheeler and Wheeler, 1973. Auts of Deep Canyon, p. 52.

1356 Hymenoptera in America North of Mexico

SPECIES GROUP MARICOPA

californicus (Buckley). W. Tex., s. N. Mex., s. Utah., Ariz., Nev., s. Calif.; Mexico. Ecology:
Nests are surmounted by a circular or semicircular crater of loose sand and have a
single entrance. California harvester ant.
Myrmica californica Buckley, 1867. Ent. Soc. Phila., Proc. 6:336. 3.
Pogonomyrmex badius var. estebanius Pergande, 1893. Calif. Acad. Sci., Proc. 4:33. %, .
Pogonomyrmex californicus longinodis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:311.
3

Pogonomyrmex californicus nitratus Cook, 1953. The Ants of Calif., pp. 99-100, 3 figs. %.

Biology: Wheeler, 1910. Ants, pp. 188-190, 200-201, 284-286, 290-291. —Essig, 1926. Ins. of
West. N. Amer., p. 861. —Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Michener, 1942. Sci.
Monthly 55:248-258 (history and behavior of a colony). —Michener, 1960. Kans. Ent. Soc.,
Jour. 33:46 (treetop mating habit). —McCluskey, 1969. Amer. Zool. 9:566 (flights).
—Erickson, 1972. Ent. Soc. Amer., Ann. 65:57-61 (mark-recapture techniques for
population estimates). —Wheeler and Wheeler, 1973. Ants of Deep Canyon pp. 53-60.
—Whitford and Ettershank, 1975. Environ. Ent. 4: 689-696 (factors affecting foraging
activity).

Morphology: Shapley, 1921. Natl. Acad. Sci., Proc. 6:687-690 (pterergates). —Tullock, 1930.
Psyche 37:61-70.
comanche Wheeler. Southwest. Ark., w. La., s. Kans., Okla., e. Tex. Ecology: Nests are in
sandy areas in close proximity to post-oak groves and each is marked by a crescentric
or circular crater of sand or sandy soil 3 inches to 2 feet in diameter with a single
entrance.
Pogonomyrmex occidentalis comanche Wheeler, 1902. Psyche 9:392. %.

Biology: Wheeler, 1910. Ants, pp. 201, 284-285, 292. —Strandtmann, 1942. Ent. Soc. Amer.,
Ann. 38:140.

magnacanthus Cole. S. Nev., w. Ariz., s. Calif.; Mexico. Ecology: Nests are in loose, sandy soil
of open deserts with a circular, shallow crater 4 to 6 inches in diameter.
Pogonomyrmex (Pogonomyrmex) magnacanthus Cole, 1968. Pogonomyrmex Harvester
Ants, pp. 133-137. 8, 2, d.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 60-61.
maricopa Wheeler. W. Tex., s. Colo., N. Mex., s. Utah., s. Nev., Ariz., s.e. Calif.; Mexico.
Ecology: Nests and habits are similar to those of P. californicus.
Pogonomyrmex californicus maricopa Wheeler, 1914. Psyche 21:155. 8, @.
Pogonomyrmex californicus barnesi Smith, 1929. Ent. Soc. Amer., Ann. 22:546. 8.
Pogonomyrmex californicus sinaloanus Olsen, 1934. Harvard Univ., Mus. Comp. Zool.,
Bul. 77:504. 3.

Taxonomy: Cole, 1954. Tenn. Acad. Sci., Jour. 29:120.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:399. —Gregg, 1963. Ants of Colo., pp. 326-328.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 61-62.

SPECIES GROUP BADIUS

badius (Latreille). N. C., S. C., Ga., Fla., Ala., Miss., La. Ecology: Nests are in sand or sandy

soil usually in open woodlands and grassy fields; they construct single or multiple,
flattened, circular sand craters or dome-shaped sand mounds with depressed tops. The
only species of Pogonomyrmex found east of the Mississippi River and the only N.
Amer. polymorphic species. Florida harvester ant.

Formica badia Latreille, 1802. Hist. Nat. Fourmis, p. 238. 8, °.

Myrmica transversa Smith, 1858. Cat. Hym. Brit. Mus. 6:129. &.

Atta crudelis Smith, 1858. Cat. Hym. Brit. Mus. 6:170. %, °.

Myrmica brevipennis Smith, 1858. Cat. Hym. Brit. Mus. 6:130. 5. A questionable syn.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:2 (larva).

Superfamily FORMICOIDEA 1357

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:384-385. —Turner, 1909. Biol. Bul.
17:161-170. —Wheeler, 1910. Ants, pp. 131, 152, 201, 280, 283-285, 292. —Wray, 1938. Ent.
Soe. Amer., Ann. 31:196-200. —Van Pelt, 1953. Tenn. Acad. Sci., Jour. 28:164-168. —Van
Pelt, 1958. Amer. Midland Nat. 59:11-12. —Carter, 1962. Elisha Mitchell Sci. Soc., Jour.
78:172. —Golley and Gentry, 1964. Ecology 45:217-225. —Hangartner, Reichson, and
Wilson, 1970. Animal Behavior 18:331-334 (orientation to nest material). —Gentry and
Carlson, 1970. Assoc. Southeast. Biol., Bul. 17:44 (formation and early development).
—Holldobler and Wilson, 1970. Psyche 77:385-399 (recruitment trails). —Holldobler, 1971.
Science 171:1149-1151 (homing). —Nickle and Neal, 1972. Fla. Ent. 55:65-66 (foraging
behavior). —Morrill, 1972. Fla. Ent. 55:59-60 (tool using behavior). —Morrill, 1975. Ga. Ent.
Soc., Jour. 10: 50-51 (a predator, Apiomerus crassipes crassipes (F.) (Reduviidae)).

Morphology: Wilson, Durlach, and Roth, 1958. Psyche 65:108-114 (chemical releasers of
necrophoric behavior). —Wilson, 1958. Psyche 65:41-51 (chemical releasers of alarm and
digging behavior). —Hermann and Blum, 1967. Ent. Soc. Amer., Ann. 60:661-668
(morphology and histology of poison apparatus). —Blum and Hermann, 1969. Ga. Ent. Soc.,
Jour. 4:23-28 (poison gland, functions of the main glandular elements).

UNPLACED TAXON OF POGONOMYRMEX SUBGENUS POGONOMYRMEX

Pogonomyrmex (Pogonomyrmex) californicus var. Hindleyi Forel, 1914. Soc. Vaud. des Sci.
Nat., Bul. 50:270. 2. Calif. (Escondido).

Genus POGONOMYRMEX Subgenus EPHEBOMYRMEX Wheeler

Pogonomyrmesx subg. Ephebomyrmex Wheeler, 1902. Psyche 9:390.
Type-species: Pogonomyrmex naegelii Forel. Desig. by Wheeler, 1911.

Biology: Creighton, 1956. Psyche 63:54-56.
huachucanus Wheeler. N. Mex., Ariz. Ecology: Nests are in stony soil, sometimes under
stones, in the open exposed to the sun; there is little or no excavated ‘soil.
Pogonomyrmex huachucanus Wheeler, 1914. Psyche 21:151. 8.

Taxonomy: Creighton, 1952. Psyche 59:76-81 (each caste). —Wheeler and Wheeler, 1972. Ga.
Ent. Soc., Jour. 7:237 (larva).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 26:119.
imberbiculus Wheeler. Southwest. Okla., central and w. Tex., s.w. Colo., N. Mex., s. Ariz., s.
Nev., s. Calif.; Mexico. Ecology: Nests are in open or under stones, sometimes with a

small crater.
Pogonomyrmesx imberbiculus Wheeler, 1902. Amer. Nat. 36:87, 97. %.
Pogonomyrmex (Ephebomyrmex) townsendi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:80. %.

Taxonomy: Creighton, 1956. Psyche 63:63-64 ?. —Wheeler and Wheeler, 1952. Psyche 59:111
(larva). —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:237 (larva).

Biology: Wheeler, 1910. Ants, pp. 283-284, 290, 292. —Wheeler, 1917. Psyche 24:178-179.
—Creighton, 1956. Psyche 63:54-56. —Gregg, 1963. Ants of Colo., pp. 334-336. —Wheeler
and Wheeler, 1973. Ants of Deep Canyon, pp. 63-64.

pima Wheeler. S. Ariz.; Mexico. Ecology: Habits are similar to those of P. imberbiculus.
Pogonomyrmex (Ephebomyrmex) pima Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:79. §.

Biology: Wheeler, 1910. Ants, p. 283. —Cole, 1934. Ent. Soc. Amer., Ann. 27:399-400.
—Creighton, 1956. Psyche 63:61.

TRIBE PHEIDOLINI
Genus STENAMMA Westwood

Stenamma Westwood, 1840. Introduct. Mod. Class. Ins., Sup. 2:83.
Type-species: Stenamma westwoodii Westwood. Monotypic.

Asemorhoptrum Mayr, 1861. Die Europaischen Formiciden, p. 76.
Type-species: Myrmica lippula Nylander. Monotypic.

Theryella Santschi, 1921. Soc. d’Hist. Nat. l’Afrique de Nord, Bul. 12:68.

1358 Hymenoptera in America North of Mexico

Type-species: Theryella myops Santschi. Monotypic.

This is a holarctic genus which also extends into the Neotropical Region. Nests are usually in
wooded areas where the small colonies are found in logs, stumps, branches, in soil beneath rocks,
moss, debris, or humus. Workers are apparently predatory, feeding mostly on other arthropods.
Snelling (1973) set up five species groups for the 12 western species of this genus; other North
American species may belong to one of these groups or form new groups.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:297-301. —Forel, 1901. Soc. Ent. de
Belg., Ann. 45:347-348. —Wheeler, 1903. Psyche 10:164-168. —Smith, 1957. Amer. Midland
Nat. 57:133-174 (N. Amer.). —Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:1-38
(western N. Amer.).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:50-52. (larvae).

Biology: Francoeur, 1966. Ent. Soc. Quebec, Ann. 11:115-119 (4 species in Que.).
brevicorne (Mayr). N.S., Que. s. to Va., w. to Ont., Minn., Nebr., Colo. (?). Ecology: Prefers to
nest in wooded areas, though sometimes found in meadows, in soil under stones, in
debris, rotting wood, etc.
Aphaenogaster brevicornis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:443, 447-448. 8,

OF
Stenamma neoarcticum (!) Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:454. % (female,

male misdet.).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:373, 382. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bul. 22:585. —Kannowski, 1958. Ent. News 69:231-233
(swarming of males). —Gregg, 1963. Ants of Colo., pp. 346-348. —Francoeur, 1966. Ent.
Soe. Quebec, Ann. 11:115-119.

californicum Snelling. Calif. Ecology: Found in fern humus and oak leaf litter.
Stenamma californicum Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:16-18. %,

oF
carolinense Smith. N. C. (Hoffman, Richmond Co.). Ecology: Collected in sparsely vegetated
sandy soil.
Stenamma carolinense Smith, 1951. Ent. Soc. Wash., Proc. 53:156-158. &.
chiricahua Snelling. S. Ariz. Ecology: Found in mountains in shaded creek bed.
Stenamma chiricahua Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:7-10. 8, 9,
é.
diecki Emery. Que., Maine s. to N. C., w. to B. C., Wash., Oreg., Calif.; Mexico. Ecology: A
widespread species adapted to various habitats but apparently absent in the southern
states from S. C. to Fla., w. to Tex. Nests are usually in wooded areas in rotting wood
or under objects.
Stenamma westwoodi diecki Emery, 1895. Zool. Jahrb., Abt. f. System. 8:300. &, °.
Stenamma westwoodi diecki var. impressum Emery, 1895. Zool. Jahrb., Abt. f. System.
8:301. 8.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:50 (larva). —Snelling, 1973.
Los Angeles Co. Mus., Contrib. Sci. 245:18-21.

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11:244. —Dennis, 1938. Ent. Soc.
Amer., Ann. 31:284, 304. —Cole, 1950. Tenn. Acad. Sci., Jour. 25:297. —Wheeler and
Wheeler, 1963. Ants of N. Dak., pp. 121-122. —Gregg, 1963. Ants of Colo., pp. 350-351, 353.
—Francoeur, 1966. Ent. Soc. Quebec, Ann. 11:115-119. —Lettendre and Pilon, 1972. Nat.
Canad. 99:73-82 (in Que.).

dyscheres Snelling. Calif. Ecology: Found at low to moderate elevations in the Sierra Nevada
foothills. Specimens have been found in pine duff.
Stenamma dyscheres Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:21-25. 3, 9,
3.
exasperatum Snelling. Calif. (Calaveras Co. and Yosemite Nat. Pk.). Ecology: Found under

stones in Sequoia and pine forests.
Stenamma exasperatum Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:28-30. %.

Superfamily FORMICOIDEA 1359

foveolocephalum Smith. Miss. (2 mi. s. of Ackerman). Ecology: Specimens collected from
sandy soil on a thinly wooded hillside.
Stenamma fovolocephala (!) Smith, 1930. Ent. Soc. Amer., Ann. 23:564. 9.

Biology: Smith, 1931. Ent. News 42:17.

heathi Wheeler. Nev., Calif.; Mexico. Ecology: Collected under rocks in pine forests.
Stenamma brevicorne heathi Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:410. 8.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:30-34.
huachucanum Smith. Colo., Ariz. Ecology: Collected from under rocks.
Stenamma huachucanum Smith, 1957. Amer. Midland Nat. 57:153. %.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sei. 245:34.

Biology: Gregg, 1963. Ants of Colo., pp. 350-351, 353.
impar Forel. Que., Mass. s. to N. C., w. to N. Dak., IIl., Mo. Ecology: Nests in soil or rotten
wood.
Stenamma brevicorne impar Forel, 1901. Soc. Ent. de Belg., Ann. 45:347. 8, @.

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., p. 122. —Francoeur, 1966. Ent. Soc.
Quebec, Ann. 11:115-119.
meridionale Smith. Va. s. to Ga., w. to IIl., Ark. Ecology: Found in wooded areas in soil.
Stenamma meridionale Smith, 1957. Amer. Midland Nat. 57:169-171. %, @.
occidentale Smith. S. Dak., Colo., N. Mex., Idaho, Utah, Ariz., B. C., Wash., Oreg., Calif.;
Mexico. Ecology: Nests in soil under rocks.
Stenamma occidentale Smith, 1957. Amer. Midland Nat. 57:146. %, 2, o.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:25-28.

Biology: Gregg, 1963. Ants of Colo., pp. 352-354.

punctatoventre Snelling. Calif. Ecology: Specimens collected under a decayed limb in litter.
Stenamma punctatoventre Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:10-12. &,
2s
schmittii Wheeler. Que., Maine s. to N. C., w. to Minn., Iowa, Mo., Tenn. Ecology: Nests in
wooded areas under stones, rotten wood and litter.
Stenamma brevicorne schmittii Wheeler, 1903. Psyche 10:167. 9.

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:93. —Francoeur, 1966. Ent. Soc.
Quebec, Ann. 11:115-119.
sequoiarum Wheeler. Calif. (coastal region from Siskiyou Co. to San Luis Obispo Co.).
Ecology: Specimens have been found under stones in redwood forests and in redwood
litter.
Stenamma brevicorne sequoiarum Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:520.
eS.
Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:28.
smithi Cole. Idaho, Utah, Nev. Ecology: Collected from sagebrush and juniper duff.

Stenamma smithi Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7 (3):7. 9.
Stenamma knowltoni Gregg, 1972. Great Basin Nat. 32:35-39. 3, 9.

Taxonomy: Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:12-16.
wheelerorum Snelling. Nev. (Mt. Rose, Washoe Co., 8800 ft.).
Stenamma wheelerorum Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 245:34-35. &,
2

Genus APHAENOGASTER Mayr

Aphaenogaster Mayr, 1853. Zool.-Bot. Gesell. Wien, Verh. 3:107.

Type-species: Aphaenogaster sardous Mayr. Desig. by Bingham, 1903.
Deromyrma Forel, 1913. Zool. Jahrb., Abt. f. System. 36:772.

Type-species: Aphaenogaster (Ischnomyrmex) swammerdami Forel. Monotypic.
Planimyrma Viehmeyer, 1914. Zool. Jahrb., Abt. f. System. 37:604.

Type-species: Stenamma (Ischnomyrmezx) loriai Emery. Orig. desig.

1360 Hymenoptera in America North of Mexico

Aphaenogaster subg. Attomyrma Emery, 1915. Accad. delle Sci. dell’Ist. Bologna, Rend. (n.
s.) 19: 70.
Type-species: Formica subterranea Latreille. Orig. desig.
Novomessor Emery, 1915. Accad. delle Sci. dell’Ist. Bologna, Rend. (n. s.) 19:73.
Type-species: Aphaenogaster (Ischnomyrmex) cockerelli Andre. Orig. desig.
Nystalomyrma Wheeler, 1916. Roy. Soc. So. Austral., Trans. 40:215.
Type-species: Myrmica longiceps Smith. Orig. desig.

Most species of this genus nest in soil beneath a covering object but some may be in rotten
logs or arboreal in dead branches. Two species, mariae and tennesseensis are believed to be tem-
porary social parasites but more investigation is needed. The genus is worldwide. The North
American fauna is in need of intensive study.

Revision: Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:443-446. —Emery, 1895. Zool. Jahrb.,
Abt. f. System. 8:301-306. —Wheeler and Wheeler, 1934. Psyche 41:6-12 (treatae and
forms). —Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci., Proc. 69:343-354
(Novomessor). —Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:147-151 (Novomessor).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:54-62 (larvae). —Brown,
1973. In Meggers, et al., Tropical forest ecosystems in Afr. and S. Amer., pp. 178-185
(generic syn.). —Brown, 1974. Ent. News 85: 45-47 (justification of synonymy of
Novomessor).

Biology: Fellers and Fellers, 1976. Science 192: 70-72 (tool use in 4 species).

albisetosa Mayr. Tex., N. Mex., Ariz.; Mexico. Ecology: Found in arid plateaus, at elevations of
2,500 to 5,000 feet; makes small crater nests, often under large stones. Omnivorous, but
has a preference for fruits and seeds.
Aphaenogaster albisetosa Mayr, 1886. Zool.-Bot. Gesell. Wien., Verh. 36:443, 446. 9.
Novomessor cockerelli var. minor Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:148. %.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soe. Wash., Proc. 55:70 (larva). —Wheeler and
Wheeler, 1960. Ent. Soc. Wash., Proc. 62:8 (larva).

Biology: Wheeler, 1910. Ants, pp. 280-282. —Cole, 1934. Ent. Soc. Amer., Ann. 27:398.
—Creighton, 1955. Psyche 62:89-97.
ashmeadi (Emery). N. C., Tenn. s. to Fla., w. to Mo., Tex.
Stenamma (Aphaenogaster) treatae var. ashmeadi Emery, 1985. Zool. Jahrb., Abt. f.
System. 8:302. 8.
Aphaenogaster treatae harnedi Wheeler, 1919. Psyche 26:50. &.

Taxonomy: Wheeler, 1919. Psyche 26:50. —Smith, 1924. Ent. News 35:50.

boulderensis boulderensis Smith. Ariz. (Horseshoe Is., Mead Lake); Nev., Tex. (?). Ecology:
Probably nests in small colonies beneath stones. Possibly confused with A. mutica
Pergande.
Aphaenogaster (Attomyrma) boulderensis Smith, 1941. Great Basin Nat. 2:118, 120. 8.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7:9.
boulderensis smithi Gregg. N. Mex. (Carrizozo).
Aphaenogaster (Attomyrma) boulderensis smithi Gregg, 1949. Ent. Soc. Wash., Proc.
51:171. 8.

cockerelli Andre. Tex., N. Mex., Ariz., Nev., s. Calif.; Mexico. Ecology: Found in arid plateaus
as high as 7,000 ft. Constructs large craters of pebbles under four inches in height with
a single entrance.
Aphaenogaster (Ischnomyrmex) cockerelli Andre, 1893. Rev. de. Ent. 12:150. &.
Aphaenogaster sonorae Pergande, 1893. Calif. Acad. Sci., Proc. 4:34. 8.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:10 (larva). —Wheeler and
Wheeler, 1972. Ga. Ent. Soc., Jour. 7:239 (larva).

Biology: Wheeler, 1910. Ants, pp. 69, 178, 201, 280-282. —Cole, 1934. Ent. Soc. Amer., Ann.
27:397-398. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:67. —Cole, 1953. Tenn. Acad.
Sci., Jour. 28:243. —Creighton, 1955. Psyche 62:89-97. —Whitford and Ettershank, 1975.
Environ. Ent. 4: 689-696 (factors affecting foraging activity).

Superfamily FORMICOIDEA 1361

Morphology: Vick, Drew, et al., 1969. Ent. Soc. Amer., Ann. 62:723-725 (identification of
hydrocarbons).
flemingi Smith. N. C. s. to Fla., w. to Ky., La.
Aphaenogaster texana flemingi Smith, 1928. Ent. News 39:275. &%.
Aphaenogaster texana macrospina Smith, 1934. Ent. Soc. Amer., Ann. 27:386. 9.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:238 (larva). —Smith, 1957.
Brooklyn Ent. Soc., Bul. 52:113.
floridana Smith. N. C. s. to Fla., Ala.
Aphaenogaster (Attomyrma) floridana Smith, 1941. Great Basin Nat. 2:118. &.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:7 (larva).
fulva Roger. Vt. s. to Fla., w. to Nebr., Colo., La. Ecology: Nests in rotting wood such as logs
and stumps or in soil under stones or other objects. Food consists mostly of live and
dead insects. Temporary host of A. tennesseensis (Mayr) and possibly of A. mariae
Forel.
Aphaenogaster fulva Roger, 1863. Berlin Ent. Ztschr. 7:190. %.
Aphaenogaster fulva var. rubida Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:147. 8.

Biology: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22:586. —Wheeler,
1910. Ants, pp. 81, 83, 206, 448, 453. —Gregg, 1963. Ants of Colo., pp. 336-337, 339. —Smith,
1965. U.S. Dept. Agr., Tech. Bul. 1326:24-25.

huachucana crinimera Cole. N. Mex. (Bandelier Natl. Mon.).
Aphaenogaster (Allomyrma(!)) huachucana crinimera Cole, 1953. Tenn. Acad. Sci., Jour.
28:82. 3, 2, d.
huachucana huachucana Creighton. Colo., Ariz.
Aphaenogaster (Attomyrma) huachucana Creighton, 1934. Psyche 41:189. §.

Taxonomy: Creighton, 1951. Psyche 58:89-99 (each caste).

Biology: Gregg, 1963. Ants of Colo., pp. 338-340.
lamellidens Mayr. N. Y. s. to Fla., w. to Ill., Mo., Tex. Ecology: Typically nests in stumps and
logs and feeds on live and dead insects.
Aphaenogaster lamellidens Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:443, 445. 3, 2, d.
Aphaenogaster lamellidens var. nigripes Smith, 1923. Ent. News 34:308. 8.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24:50, 52.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:383. —Smith, 1918. Ent. News 29:21.
—Smith, 1924. Ent. News 35:51. —Smith, 1928. Ent. News 39:246. —Dennis, 1938. Ent.
Soe. Amer., Ann. 31:285, 304. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:20-21.

mariae Forel. N. Y. s. to Fla., w. to Iowa, Kans. Ecology: Probably a temporary parasite. Host:
Aphaenogaster fulva Roger.
Aphaenogaster mariae Forel, 1886. Soc. Ent. de Belg., Bul. 30:41. 8.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22:585. —Buren,
1944. Iowa State Col., Jour. Sci. 18:284 (female).

Biology: Wheeler, 1910. Ants, pp. 151, 448. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:93.
megommata Smith. Oreg., Nev., Ariz., Calif. Ecology: A crepuscular and nocturnal forager.
Aphaenogaster (Attomyrma) megommatus Smith, 1963. N. Y. Ent. Soc., Jour. 71:244-246.
3.

Taxonomy: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(3):9-11 (female, male).
—Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:238 (larva).
miamiana Wheeler. Fla., Ala., Miss. Ecology: Possibly arboreal, nesting in branches.
Aphaenogaster (Attomyrma) texana var. miamiana Wheeler, 1932. N. Y. Ent. Soc., Jour.
40:5. 3, 2, do.
mutica Pergande. Tex. (?); Mexico (Lower Calif.). Ecology: Recorded from Brownsville, Tex.,
but may not occur in U. S.
Aphaenogaster mutica Pergande, 1895. Calif. Acad. Sci., Proc. 5:891. &.

1362 Hymenoptera in America North of Mexico

patruelis Forel. Calif. (Coastal Islands); Mexico (Lower Calif., Guadeloupe Is., St. Nicholas Is.).
Aphaenogaster patruelis Forel, 1886. Soc. Ent. de Belg., Ann. 30:41. 9.
Stenamma (Aphaenogaster) patruelis bakeri Wheeler, 1904. Amer. Mus. Nat. Hist., Bul.
20:270. &.
Aphaenogaster patruelis willowsi Wheeler, 1933. Calif. Acad. Sci., Proc. 21:64. %.

Taxonomy: Wheeler, 1934. Pan-Pacific Ent. 10:133.

rudis picea (Emery). N. S. s. to N. C. Ecology: Habits are similar to rudis rudis (Emery), but
rudis picea is normally found at higher elevations. Temporary host of A. tennesseensis
(Mayr).
Stenamma (Aphaenogaster) fulvum aquia var. piceum Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:305. %, 2, d.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:57-60 (larva).

Biology: Wheeler, 1910. Ants, pp. 195, 282, 447-448, 453. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31:286-287, 305. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:94. —Haskins,
1960. N. Y. Ent. Soc., Jour. 68:66-67 (longevity of fertile females).

rudis rudis (Emery). Mass. s. to Fla., w. to Wyo., Colo. Ecology: A common and highly
adaptable species, nesting in soil, under stones or logs, in decaying wood, leaf litter,
hollow stems of plants, or under bark at bases of trees. Food consists of insects, seeds,
and pollen of ground nesting bees. Host of temporary ant parasite Aphaenogaster
tennesseensis (Mayr).
Stenamma (Aphaenogaster) fulva aquia var. rude Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:305. %, °. ’

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:56 (larva).

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:90, 94. —Talbot, 1951. Ent. Soc.
Amer., Ann. 44:302-307. —Brown, 1958. Psyche 65:39-40 (nest raided by Formica
subintegra Emery). —Gregg, 1963. Ants of Colo., pp. 340-342. —Bobb, 1965, Jour. Econ
Ent. 58:925 (predator of Neodiprion pratti pratti (Dyar)). —Smith, 1965. U. S. Dept., Agr.,
Tech. Bul. 1326:21-22.

Morphology: Gotwald, 1969. N. Y. (Cornell) Agr. Expt. Sta., Mem. 408:99-112 (mouthparts).

subterranea occidentalis (Emery). B. C. s. to Nev., Calif. A. subterranea subterranea
(Latreille) occurs in the Mediterranean region and central Europe.
Stenamma (Aphaenogaster) subterraneum occidentale Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:301. %.
Aphaenogaster subterranea borealis Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:412. %.
Aphaenogaster subterranea valida var. manni Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:516. 8.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:363. —Wheeler and Wheeler, 1953. Ent. Soc.
Wash., Proc. 55:61 (larva).

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul 40:7. —Bruniquel, 1970. Insectes Sociaux
17:245-252 (biology of A. subterranea in Europe).
subterranea valida Wheeler. S. Dak. to B. C., s. to Colo., Utah. Ecology: Prefers to nest under
stones in moist, shady foothill canyons. Intergrades in northern part of range with
subsp. occidentalis.
Aphaenogaster subterranea valida Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:411. &,
Qe Be

Biology: Gregg, 1947. Colo. Univ., Studies (ser. D) 2:393. —Gregg, 1963. Ants of Colo., pp.
342-344.

tennesseensis (Mayr). Que., Ont. s. to Fla., w. to Minn., S. Dak., Nebr., Kans., Okla. Ecology: In
early stages of colony formation, probably a temporary parasite in ground nests of other
Aphaenogaster species. Lives in ground nests only when females occur in nests of rudis
and picea, otherwise it is exclusively a wood nesting ant. Host: Aphaenogaster fulva
Roger, A. rudis picea (Emery), A. rudis rudis (Emery).
Atta Tenneseensis (!) Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12:743. &.
Atta laevis Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12:743. @.

Superfamily FORMICOIDEA 1363

Myrmica subruba Buckley, 1867. Ent. Soc. Phila., Proc. 6:336. 3, “2” = d.
Stenamma (Aphaenogaster) tenesseense (!) var. ecalcaratum Emery, 1895. Zool. Jahrb.,
Abt. f. System. 8:301. .

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22:585-586.
—Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:61-62 (larva). —Francoeur and
Beique, 1968. Nat. Canad. 95:227 (distribution).

Biology: Wheeler, 1910. Ants, pp. 114, 447-448, 450. —Dennis, 1938. Ent. Soc. Amer., Ann.
31:287-288. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:90, 94. —Smith, 1965. U. S.
Dept. Agr., Tech. Bul. 1326:23-24.

texana carolinensis Wheeler. S. Va. to n. Ala. Ecology: Nests are in both wood and soil.
Aphaenogaster texana var. carolinensis Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:414.

’

Taxonomy: Smith, 1931. Ent News 42:17.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:287, 305.

texana texana (Emery). N. C., Tenn. s. to Fla., w. to Kans., Ariz.
Stenamma (Aphaenogaster) fulvuum aquia var. texanum Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:306. &.
Aphaenogaster texana var. furvescens Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:413.
Cee
Aphaenogaster (Deromyrma) Silvestrii Menozzi, 1929. Bol. Lab. Zool. Gen. e Agr. Portici
22:2825 95 9

Taxonomy: Creighton, 1934. Psyche 41:192. —Wheeler and Wheeler, 1953. Ent. Soc. Wash.,
Proce. 55:62 (larva).
treatae pluteicornis Wheeler and Wheeler. Ala. w. to Okla., Tex.
Aphaenogaster treatae pluteicornis Wheeler and Wheeler, 1934. Psyche 41:7, 12. 8, 2, d.
Aphaenogaster treatae pluteicornis var. oklahomensis Wheeler and Wheeler, 1934. Psyche
41:10, 12. 3, 2.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:62 (larva).
treatae treatae Forel. Ont. s. to Fla., w. to Mich., Ohio, IIl., Ala. Ecology: Nests are in soil
usually in open areas.
Aphaenogaster treatae Forel, 1886. Soc. Ent. de Belg., Bul. 30:40-41. %, 2, d.
Aphaenogaster treatae wheeleri Mann, 1915. Psyche 22:51. 8, °.
Aphaenogaster treatae pluteicornis var. alabamensis Wheeler and Wheeler, 1934. Psyche
41:10. 8, 2.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22:585. —Wheeler,
1919. Psyche 26:50.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:303. —Wheeler, 1910. Ants, pp. 151,
200. —Talbot, 1954. Mich. Univ., Contrib. Lab. Vertebr. Biol. 69:1-9. —Talbot, 1966. Kans.
Ent. Soc., Jour. 39:67-77 (flights).

uinta Wheeler. Colo., Idaho, Utah, Nev. Ecology: Nests are in fully exposed areas of great
aridity.
Aphaenogaster winta Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:517. %, 2, d.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:363-364.

Biology: Cole, 1934. Psyche 41:223. —Gregg, 1963. Ants of Colo., pp. 344-345, 347.

UNPLACED TAXA OF APHAENOGASTER

According to Creighton (1950, Harvard Univ., Mus. Comp. Zool., Bul.) the following forms
were described from minims only, and it is not possible to determine what they actually
represent.

aquia (Buckley). N. Y., Va. In the early literature, this and fulva aquia (Buckley) were used
for what is currently called rudis rudis (Emery).
Myrmica (Monomarium (!)) aquia Buckley, 1867. Ent. Soc. Phila., Proc. 6: 341. DOs

1364 Hymenoptera in America North of Mexico

fulva aquia var. pusilla (Emery). D. C.
Stenamma (Aphaenogaster) fulvum aquia var. pusillum Emery, 1895. Zool. Jahrb., Abt. f.

System. 8: 306. $.
texana nana Wheeler. Fla. (Gainesville).
Aphaenogaster (Attomyrma) texana nana Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 6. &.
texana punctithorax Cole. Tenn. (Gregory’s Bald, Great Smoky Mtn. Natl. Pk.).
Aphaenogaster texana punctithorax Cole, 1938. Amer. Midland Nat. 19: 239. &.

Genus VEROMESSOR Forel

Novomessor subg. Veromessor Forel, 1917. Soc. Vaud. des Sci. Nat., Bul. 51:235.
Type-species: Aphaenogaster andrei Mayr. Desig. by Emery, 1921.
Veromessor subg. Lobognathus Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:152. Nomen
nudum.

These are harvester ants, most common in the arid Southwest. They feed on seeds and other
Arthropods and usually nest in exposed soil where the nests may or may not be marked with a
small crater or piles of chaff from seeds stored in the nest. They are docile ants which seldom
sting.

Revision: Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci., Proc. 69:354-387.

Taxonomy: Smith, 1956. Pan-Pacific Ent. 32:36-37 (key to workers of U. S. species).
—Wheeler and Wheeler, 1956. Psyche 63:142-143 (larvae). —Wheeler and Wheeler, 1972.
Ga. Ent. Soe., Jour. 7:240 (larvae).

Biology: Creighton, 1953. Amer. Mus. Novitates 1612:1-18. —Went, Wheeler, and Wheeler,
1972. BioScience 22:82-88 (feeding and digestion).
andrei (Mayr). Oreg., Nev., Ariz., Calif.; Mexico. Ecology: Found from sea level up to 3500
feet. Nests are in various types of soil and openings are surrounded by a circular disc or
low, obscure crater.
Aphaenogaster andrei Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:443, 448. 8.
. Veromessor andrei flavus Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci., Proc.
69:361, 366. 9.
Veromessor andrei castaneus Wheeler and Creighton, 1934. Amer. Acad. Arts and Sci.,
Proc. 69:361, 365. 8.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:306-307 (worker, female). —Wheeler
and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 (larva).

Biology: Wheeler, 1910. Ants, p. 280. —Cole, 1934. Ent. Soc. Amer., Ann. 27:398. —Mallis,
1941. South. Calif. Acad. Sci., Bul. 40:7. —Eckert and Mallis, 1937. Calif. Agr. Expt. Sta.
Cir. 342:24 (economics). —Creighton, 1953. Amer. Mus. Novitates 1612:2-4. —McCluskey,
1958. Science 128:536-537 (daily rhythm of male). —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 65-67.

chamberlini (Wheeler). Calif. (coastal islands and mainland).
Messor chamberlini Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:410. 8.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 (larva).

lariversi Smith. Nev., Calif. Ecology: Nests are in exposed soil or gravel, and may be marked
by one or more small circular craters several inches in diameter.
Veromessor lariversi Smith, 1951. Great Basin Nat. 11:94-96. &.

Taxonomy: Cole, 1955. Tenn. Acad. Sci., Jour. 30:51-52 (female).

Biology: Creighton, 1953. Amer. Mus. Novitates 1612:5-6. —Cole, 1963. Ent. Soc. Amer., Ann.
56:680-682. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(3):12.

lobognathus (Andrews). Southwest. N. Dak., Colo., Nev. Ecology: Colonies are populous and
nests are under large stones.

Messor lobognathus Andrews, 1916. Psyche 23:82. 8.

Taxonomy: Wheeler and Wheeler, 1956. Psyche 63:143-145 (larva). —Cole, 1963. Ent. Soc.
Amer., Ann. 56:680-682 (each caste).

Superfamily FORMICOIDEA 1365

Biology: Gregg, 1955. Psyche 62:45-52 (rediscovery). —Wheeler and Wheeler, 1956. Psyche
63:140-145 (in N. Dak.). —Wheeler and Wheeler, 1959. Ent. Soc. Amer., Ann. 52:176-179
(behavior, nest structure). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 123-125.
—Gregg, 1963. Ants of Colo., pp. 354-356. —Wheeler and Wheeler, 1965, Kans. Ent. Soc.,
Jour. 38:55-61. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(3):12-13.

pergandei (Mayr). Ariz., Nev., Calif.; Mexico. Ecology: Nests are in exposed soil, usually with a
single entrance which is surrounded by a crater of excavated soil. Workers are diurnal

foragers and can remain active during periods of intense heat.
Aphaenogaster pergandei Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:444, 448. 9.

Taxonomy: Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342:8, 25. —Cole, 1963. Ent.
Soc. Amer., Ann. 56:681. —Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 (larva).

Biology: Wheeler, 1910. Ants, pp. 16, 280. —Cole, 1937. Ent. News 48:101. —Mallis, 1941.
South. Calif. Acad. Sci., Bul. 40:68. —Creighton, 1953. Amer. Mus. Novitates 1612:7-17.
—Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(3):13-14. —Wheeler and Wheeler,
1965. Kans. Ent. Soc., Jour. 38:55 (habitat, harvesting habits, defense). —Wheeler and
Wheeler, 1967. Kans. Ent. Soc. Jour. 40:238 (distribution and habitat). —Tevis, 1958.
Ecology 39:695-704 (interrelations with some desert ephemerals). —Wheeler and Wheeler,
1973. Ants of Deep Canyon, pp. 67-72. —Clark and Comanor, 1973. Amer. Midland Nat. 90:
467-474 (quantitative examination of spring foraging). —Wheeler and Rissing, 1975.
Pan-Pacific Ent. 51: 205-216 (nest). —Wheeler and Rissing, 1975. Pan-Pacifie Ent. 51:
303-314 (behavior). —Rissing and Wheeler, 1976. Pan-Pacific Ent. 52: 63-72 (foraging
responses to changes in seed production).

smithi Cole. Nev. (Mercury, Nye Co.), Ariz. Ecology: Nests are in fully exposed areas, each
with a small circular crater of soil about their entrance.
Veromessor smithi Cole, 1963. Ent. Soc. Amer., Ann. 56:678-680. %, 2, od.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:240 (larva).

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(3):14.
stoddardi chicoensis Smith. Calif. (Chico). Ecology: The type series was found beneath a large
stone.
Veromessor stoddardi chicoensis Smith, 1956. Pan-Pacifie Ent. 32:36-38. 8.
stoddardi stoddardi (Emery). S. Calif.; Mexico. Ecology: Nests have been found in hard clay
soil, each with several openings situated at the bottom of a shallow depression and
without excavated soil.
Stenamma (Messor) stoddardi Emery, 1895. Zool. Jahrb., Abt. f. System. 8:307. 9.

Biology: Wheeler, 1910. Ants, p. 280. —Creighton, 1953. Amer. Mus. Novitates 1612:17-18.

Genus PHEIDOLE Westwood

Pheidole Westwood, 1841. Ann. and Mag. Nat. Hist. 6:87.
Type-species: Atta providens Sykes. Monotypic.

Oecophthora Heer, 1852. Naturf. Gesell. in Zurich, Neujahrsbl. 54:11, 15.
Type-species: Oecophthora pusilla Heer. Monotypic.

Ischnomyrmex Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12:738.
Type-species: Myrmica longipes Smith. Monotypic.

Leptomyrma Motschulsky, 1863. Soc. Imp. Nat. Moscou, Bul. 36:17.
Type-species: Leptomyrma gracilipes Motschulsky. Monotypic.

Pheidolacanthinus Smith, 1864. Linn. Soc. London, Jour., Zool. 8:75.
Type-species: Pheidolacanthinus armatus Smith. Monotypic.

Pheidole subg. Ceratopheidole Pergande, 1895. Calif. Acad. Sci., Proc. 5:889.
Type-species: Pheidole (Ceratopheidole) granulata Pergande. Monotypic.

Phidole Bingham, 1903. Fauna Brit. India, Hym. 2:220. Variant spelling.

Epipheidole Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19:664.
Type-species: E'pipheidole inquilina Wheeler. Monotypic.

Sympheidole Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:1, 7.
Type-species: Sympheidole elecebra Wheeler. Monotypic.

Phidola Schulz, 1906. Spolia Hym., p. 155. Emend.

1366 Hymenoptera in America North of Mexico

Allopheidole Forel, 1912. Soc. Ent. de Belg., Mem. 19:237.
Type-species: Pheidole kingi Andre. Desig. by Wheeler, 1913.
Decapheidole Forel, 1912. Soc. Ent. de Belg., Mem. 19:237.
Type-species: Pheidole perpusilla Emery. Desig. by Wheeler, 1913.
Tsopheidole Forel, 1912. Rev. Suisse de Zool. 20:765.
Type-species: Myrmica longipes Smith. Monotypic.
Elasmopheidole Forel, 1913. Zool. Jahrb., Abt. f. System. 36:43.
Type-species: Pheidole aberans Mayr. Desig. by Emery, 1922.
Cardiopheidole Wheeler, 1914. N. Y. Ent. Soc., Jour. 22:48-51.
Type-species: Pheidole vasliti Pergande. Orig. desig.
Anergatides Wasmann, 1915. Ent. Mitt. 4:281. Uncertain syn.
Type-species: Anergatides kohli Wasmann. Monotypic.
Macropheidole Emery, 1915. Soc. Ent. de France, Bul. p. 190.
Type-species: Pheidole fimbriata Roger. Monotypic.
Scrobopheidole Emery, 1915. Soc. Ent. de France, Bul. p. 190.
Type-species: Pheidole scrobifera Emery. Monotypic.
Stegopheidole Emery, 1915. Soc. Ent. de France, Bul. p. 190.
Type-species: Pheidole (Elasmopheidole) wpeneci Forel. Monotypic.
Trachypheidole Emery, 1915. Soc. Ent. de France, Bul. p. 190.
Type-species: Pheidole bicornis Forel. Orig. desig.
Parapheidole Emery, 1915. R. Accad. delle Sci. dell’Ist. Bologna, p. 68.
Type-species: Aphaenogaster oculata Emery. Monotypic.
Pheidole subg. Electropheidole Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bul. 64:438.
Type-species: Pheidole (Electropheidole) roosevelti Mann. Desig. by Donisthorpe,
1943.
Bruchomyrma Santschi, 1922. Soe. Cient. Argentina, An. 94:248. Uncertain syn.
Type-species: Bruchomyrma, acutidens Santschi. Monotypic.
Cephalomorium Forel, 1922. Rev. Suisse de Zool. 30:91.
Type-species: Tetramorium (Cephalomorium) bahai Forel. Monotypic.
Hendecapheidole Wheeler, 1922. Amer. Mus. Novitates 96:3.
Type-species: Pheidole tachigaliae Wheeler. Orig. desig.
Gallardomyrma Bruch, 1932. La Plata Mus., Rev. 33:271.
Type-species: Gallardomyrma argentina Bruch. Orig. desig.
Conothorax Karawajew, 1935. Treubia 15:75. Preoce. by Jekel, 1854.
Type-species: Conothorax bilobum Karawajew. Monotypic.
Conothoracoides Strand, 1935. Folia Zool. Hydrobiol. 8:176. N. name for Conothorax
Karavaiev.
Eriopheidole Kusnezov, 1952. Mus. de Entre Rios, Mem. 29:5.
Type-species: Eriopheidole symbiotica Kusnezov. Orig. desig.

A large genus of harvesting ants for which most species are found in the arid Southwest. Most
species have a dimorphic worker caste with majors or soldiers and minors which are not con-
nected by intermediate forms. Only a few species are polymorphic with intermediate sized wor-
kers. The soldier is characteristic in having an enormous head, way out of proportion to the rest
of the body, and may function as a seed husker.

Most species harvest seeds for food and chaff piles are sometimes found around the entrances
to the nests. Nests may have craterlike excavations when in exposed soil, or nests may be under
stones or other objects and sometimes in rotten wood. Though the main diet normally consists of
seeds, most species also feed on other animal food. Most colonies are small with 300 or fewer in-
dividuals.

This is a huge cosmopolitan genus for which few satisfactory subdivisions have been made. A
few species had been segregated into subgenera or closely related genera, but the bulk of the
species always remained in the subgenus Pheidole. Emery (1921) attempted a group arrange-
ment for this large subgenus, but this has met with little acceptance in recent years. It is now
the trend to thrust all species into a single unit until satisfactory divisions can be made. The ar-
rangement of this genus essentially follows the generic synonomy by Brown (1973) and the
treatment of the genus by Gregg (1958).

Superfamily FORMICOIDEA 1367

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:979-989. —Mayr, 1887. Zool.-Bot.
Gesell. Wien, Verh. 37:582-608. —Emery, 1895. Zool. Jahrb., Abt. f. System. 8:288-297.
—Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:431-478. —Gregg, 1958. N. Y. Ent. Soc.,
Jour. 66:7-48 (key to and list of species).

Taxonomy: Emery, 1921. Jn Wytsman, Gen. Ins., fase. 174:77-111 (groups). —Wheeler and
Wheeler, 1953. Ent. Soc. Wash., Proc. 55:70-80 (larvae). —Smith, 1953. N. Y. Ent. Soc.,
Jour. 61:143 (Ceratopheidole). —Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser.
3:1-46. —Creighton, 1957. N. Y. Ent. Soc., Jour. 65:203-212 (vasliti complex). —Cole, 1965.
Ent. Soe. Amer., Ann. 58:173-175 (Epipheidole). —Brown, 1973. In Meggers, et al.,
Tropical forest ecosystems in Afr. and S. Amer., pp. 178-185 (generic syn.).

Biology: Weber, 1948. Ent. News 59:31-35 (food of larvae and adults). —Szlep-Fessel, 1970.
Insectes Sociaux 17:233-244 (regulatory mechanism in mass foraging and recruitment of
soldiers).

anastasii Emery. S. Fla.; Mexico, Central Amer. Probably introduced. Reported from
greenhouses in various parts of the East, but only established in southern Fla.
Pheidole anastasii Emery, 1896. Soc. Ent. Ital., Bol. 28:76. 3, 2f.

Taxonomy: Forel, 1901. Naturhist. Mus. Hamburg Mitt. 18:78 (female).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:6. —Smith, 1933. Fla. Ent. 17:23.
bahai (Forel). N. C. (Faisons).
Tetramorium (Cephalomorium) bahai Forel, 1922. Rev. Suisse de Zool. 30:91. 8.

Taxonomy: Smith, 1955. Brooklyn Ent. Soc., Bul. 50:99 (correct taxonomic placement).

barbata Wheeler. W. Ariz., s. Nev., s.e. Calif. Ecology: In small crater nests in desert areas.
Pheidole barbata Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:448. %, 2f.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:68. —Creighton and Gregg, 1955. Colo.
Univ. Studies, Biol. Ser. 3:1-3. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 75.
bicarinata bicarinata Mayr. Mich. to Tenn., w. to N. Dak., Wyo., Colo. Ecology: Found in
grasslands where nests are usually under objects.
Pheidole bicarinata Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:982, 989. 2.
Pheidole hayesi Smith, 1924. Ent. News 35:251. 8, 2.

Biology: Hayes, 1925. Ent. News 36:42. —Talbot, 1934. Ecology 15:418, 420, 422. —Buren,
1944. Iowa State Col., Jour. Sci. 18:286. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
126-127. —Gregg, 1963. Ants of Colo., pp. 408-409.

bicarinata longula Emery. N. Y., N. Dak., w. Tex., Colo., N. Mex. Ecology: Nests may be in
sand or under stones.
Pheidole vinelandica var. longula Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 292. %,
Y.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:453-454. —Cole, 1956. Tenn. Acad.
Sci., Jour. 31:112. —Gregg, 1958. N. Y. Ent. Soc., Jour. 65:37.

Biology: Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17:9. —Wheeler and Wheeler,
1963. Ants of N. Dak., p. 127. —Gregg, 1963. Ants of Colo., pp. 408, 410.

bicarinata paiute Gregg. Nev., s. Calif. Ecology: Nests are beneath stones or with craters.
Pheidole bicarinata paiute Gregg, 1958. N. Y. Ent. Soc., Jour. 66:17-18. 9, 2.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(8):15. —Wheeler and
Wheeler, 1973. Ants of Deep Canyon, pp. 80-81
bicarinata vinelandica Forel. N. Y. to Fla. w. to N. Dak., Utah, Ariz. Ecology: Nests in rotten

logs, exposed soil, or under objects in grassy areas.

Pheidole bicarinata race vinelandica Forel, 1886. Soc. Ent. de Belg., Ann. 30:45. %, 2f, 2,
é.

Pheidole vinelandica laeviuscula Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 292. 9,
2.

Pheidole vinelandica buccalis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:454. 9, 2, @.

Pheidole vinelandica longula var. castanea Wheeler, 1915. Amer. Mus. Nat. Hist., Bul.
34:405. %, 2. Preoce. by Smith, 1858.

1368 Hymenoptera in America North of Mexico

Pheidole (Allopheidole) vinelandica var. nebrascensis Forel, 1922. Rev. Suisse de Zool.
30:902595 20.

Pheidole vinelandica longula var. huachucana Smith, 1951. In Muesebeck, et al., U.S.
Dept. Agr., Agr. Monog. 2:805. N. name for castanea Wheeler.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:18, 36.

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:336-337. —Cole, 1937. Ent. News
48:100. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:92. —Cole, 1953. Tenn. Acad.
Sci., Jour. 28:297. —Reid and Nugara, 1961. Jour. Parasitology 47:885-889 (as intermediate
hosts of a turkey tapeworm). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 127-128.
—Gregg, 1963. Ants of Colo., pp. 410, 412-413. —Smith, 1965. U. S. Dept. Agr., Tech. Bul.
1326:25-26.

californica californica Mayr. Calif. (San Francisco region southwards). Ecology: Nests are
found under objects, in oak galls, or in sandy soil.
Pheidole californica Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:981, 984, 987. 9, 2.
Pheidole californica var. incenata Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:407. 3, 2.
Pheidole californica var. satura Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:407. 8, 2.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:243 (larva).

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:395-397. —Mallis, 1941. South. Calif. Acad. Sci.,
Bul. 40:68-69.
californica oregonica Emery. Idaho, Wash., Oreg., n. Calif.
Pheidole oregonica Emery, 1895. Zool. Jahrb., Abt. f. System. 8:288, 291. %, 2.
Pheidole californica var. shoshoni Cole, 1933. Ent. Soc. Amer., Ann. 26:618. 3, 2.
Pheidole californica var. hagermani Cole, 1936. Canad. Ent. 68:35. 9, 2.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:19.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:69. —Cole, 1942. Amer. Midland Nat.
28:362.
californica pyramidensis Emery. Nev. (Pyramid Lake).
Pheidole californica nevadensis Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:408. Creat
Q. Preoce. by Forel, 1901.
Pheidole californica pyramidensis Emery, 1922. In Wytsman, Gen. Ins., fase. 174:105. N.
name for nevadensis Wheeler.
casta Wheeler. Tex. (Canyon of the Rio Grande, Langtry).
Pheidole casta Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:454. 9, 2.
cerebrosior Wheeler. N. Mex., Ariz., Calif.; Mexico. Ecology: Nests more frequently in
mountan canyons than on open desert and the species seems to prefer the evergreen
oak association.
Pheidole vinelandica cerebrosior Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:405. 8, 2.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:3-5.
ceres Wheeler. Tex., Colo., N. Mex., Ariz. Ecology: Host of the permanently parasitic ant P.
elecebra. Parasitized colonies have been found to contain only soldiers and workers of
ceres. Ceres nests under stones in rather dry, sunny localities at altitudes of 5,000 to
9,000 ft. and is a seed-storing form. Parasite: Pheidole elecebra (Wheeler).
Pheidole ceres Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:10. %, 2f, 2, d.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:20, 35.

Biology: Wheeler, 1919. Amer. Phil. Soc., Proc. 58:25. —Wheeler, 1923. Social Life Among the
Insects, p. 219. —Wheeler, 1910. Ants, pp. 279, 497. —Cole, 1953. Tenn. Acad. Sci., Jour.
28:297. —Gregg, 1963. Ants of Colo., pp. 413-415.

clementensis Gregg. Calif. (San Clemente Is., Los Angeles Co.; Orange Co.; San Diego Co.).
Pheidole clementensis Gregg, 1969. Ent. News 80: 93-97. %, 2.
clydei Gregg. N. Mex., Ariz., Nev., s. Calif. Ecology: Nests are found in and among rocks and
boulders.
Pheidole (Ceratopheidole) clydei Gregg, 1950. N. Y. Ent. Soc., Jour. 58: 89. 9.

Taxonomy: Gregg, 1953. Amer. Mus. Novitates 1673: 1 (soldier, distribution).

Superfamily FORMICOIDEA 1369

Biology: Creighton, 1964. Psyche 71: 169. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, pp. 84-86.

cockerelli Wheeler. Okla., Tex., Colo., N. Mex., Ariz.
Pheidole cockerelli Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 464. %, 2.

Biology: Gregg, 1963. Ants of Colo., pp. 415-417.
crassicornis crassicornis Emery. N. C., Tenn., Ga. w. to Tex.
Pheidole crassicornis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 296. 2.
Pheidole crassicornis var. diversipilosa Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:467.
ORL enr

Taxonomy: Forel, 1901. Soc. Ent. de Belg., Ann. 45:350 (male).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:302. —Smith, 1918. Ent. News 29:22.
crassicornis tetra Wheeler. Tex. w. to Ariz.
Pheidole crassicornis porcula var. tetra Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:467.
Cri
creightoni Gregg. Oreg., Nev., n. Calif.
Pheidole creightoni Gregg, 1955. Psyche 62:19-28. %, 9, 6d.

Taxonomy: Cole, 1957. N. Y. Ent. Soc., Jour. 65:131 (distribution).

davisi Wheeler. N. J. s. to N. C., n. Ala.
Pheidole davisi Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:380. %, 2f.

Biology: Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17:8-9. —Bequaert, 1928. N. Y.
(Cornell) Agr. Expt. Sta., Mem. 101:996.
dentata Mayr. Va. to Fla., w. to Ill, Kans, Tex. Ecology: Nests of small to large colonies are in
exposed soil with a mound of excavated earth above it, under the cover of objects, or in
rotting wood. Occasionally a house-infesting ant.
Pheidole Morrisi var. dentata Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:457. %, 2, do.
Pheidole commutata Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:459. %, 2.
Pheidole dentata var. faisonsica Forel, 1901. Soc. Ent. de Belg., Ann. 45:352. 9, 2.
Leptothorax tennesseensis Cole, 1938. Amer. Midland Nat. 19:238. 3.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:460-461. —Wheeler and Wheeler,
1953. Ent. Soc. Wash., Proc. 55:71 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14:71. —Smith, 1924. Ent. News
35:77. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:281, 304. —Cole, 1940. Amer. Midland Nat.
24:29, 44. —Van Pelt, 1950. Ent. News 61:161-163 (parasitism by an Orasema chalcid).
—Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:27-28. —Wilson, 1975. Science 190: 798-800
(enemy specification in the alarm-recruitment system).

dentigula Smith. N. C., Tenn. s. to Fla., w. to La. Ecology: Nests in soil or in well-rotted

stumps.
Pheidole dentigula Smith, 1927. Ent. News 38:310. 8, 2.

Taxonomy: Smith, 1928. Ent. News 39:245-246 (female). —Cole, 1940. Amer. Midland Nat.
24:42, 45. —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:12 (larva).

Biology: Smith, 1944. Fla. Ent. 27:14.
desertorum Wheeler. W. Okla., w. Tex., N. Mex., Utah, Ariz., Nev. Ecology: Nests have been

found beneath stones and as small crater mounds in sand.

Pheidole desertorum Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:337. %, 2, 2, d.

Pheidole desertorum var. comanche Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:339. 8,
at 2.

Pheidole desertorum var. maricopa Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:339. 8,
Y.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:362.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:397. —Cole, 1937. Ent. News 48:100. —Gregg,
1963. Ants of Colo., pp. 417-419.

1370 Hymenoptera in America North of Mexico

elecebra (Wheeler). Colo. Ecology: A permanent, workerless parasite in colonies of its host.
Host: Pheidole ceres Wheeler.
Sympheidole elecebra Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:8. 2, 3.
Biology: Wheeler, 1910. Ants, p. 497. —Wheeler, 1923. Social Life Among the Insects, p. 219.
—Gregg, 1963. Ants of Colo., pp. 429-431.
flavens Roger. Fla.; W. Indies, Central Amer. Possibly an adventive.
Pheidole flavens Roger, 1863. Berlin. Ent. Ztschr. 7:198. %, 2.
Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:45-46 (Fla. populations regarded as not
quite typical representatives of Pheidole flavens).
floridana constipata Wheeler. Tex. (Austin and New Braunfels).
Pheidole constipata Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:468. 9, 2, 2, do.
floridana floridana Emery. N. C. to Fla., w. to Tex.
Pheidole flavens floridana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 293. %, UY, °.
Pheidole lauta Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:470. 3, 2, 9, od.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:35.
gilvescens Wheeler. Ariz., s. Nev., Calif. Ecology: Nests in soil in deserts, usually forms small

craters.
Pheidole xerophila tucsonica var. gilvescens Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.

24:448. 3, UY.
Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:5-7, 40.

Biology: Cole, 1934. Ent. Soe. Amer., Ann. 27:397. —Cole, 1956. Tenn. Acad. Sci., Jour. 31:113.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 76.

grundmanni Smith. Utah (Ashley Cr. near Vernal).
Pheidole (Ceratopheidole) grundmanni Smith, 1953. N. Y. Ent. Soe., Jour. 61:144-146. %.

Biology: Gregg, 1963. Ants of Colo., pp. 426-427.
humeralis Wheeler. Tex. (Corsicana).
Pheidole humeralis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:456. 9, 2.
hyatti hyatti Emery. Okla., Tex., Colo. w. to Nev., s. Calif.; Mexico.
Pheidole hyatti Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289-290, 295. %, 2.
Pheidole hyatti var. ecitonodora Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:463. 3, 2,
OCLs
Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:74 (larva). —Wheeler and
Wheeler, 1972. Ga. Ent. Soc., Jour. 7:244 (larva).

Biology: Essig, 1926. Ins. of West. N. Amer., p. 859. —Eckert and Mallis, 1937. Calif. Agr.
Expt. Sta. Cir. 342:26. —Gregg, 1963. Ants of Colo., pp. 419-420. —Wheeler and Wheeler,
1973. Ants of Deep Canyon, pp. 79-80.

hyatti solitanea Wheeler. Calif. (coastal area in San Diego region).
Pheidole hyatti solitanea Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:409. %, Uf, 9.
inquilina (Wheeler). Nebr., Colo., Nev. Ecology: An inquiline in nests of other ants. Host:
Pheidole pilifera pilifera (Roger), P. pilifera coloradensis Emery.
Epipheidole inquilina Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19:664. Gynandromorph.

Taxonomy: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:15-17 (female, male). —Smith, 1940.
Ent. Soe. Wash., Proc. 42:106-109 (worker). —Cole, 1965. Ent. Soc. Amer., Ann. 58:173-175
(worker, soldier; placement in Pheidole; biological notes).

Biology: Wheeler, 1910. Ants, pp. 107, 113, 150, 156, 497-498. —Wheeler, 1923. Social Life
Among the Insects, pp. 215-219. —Gregg, 1963. Ants of Colo., pp. 427-429. —Cole, 1966.
Brigham Young Univ., Sci. Bul., Biol. Ser. 7(3):15.

lamia Wheeler. Ga., Miss., Tex. Ecology: Apparently hypogaeic, living under stones in small
colonies. The phragmotic head of the soldier is a striking feature of this species.
Pheidole lamia Wheeler, 1901. Amer. Nat. 35:534. %, 2.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:477-478 (soldier, female.).

Superfamily FORMICOIDEA 1371

Biology: Wheeler, 1910. Ants, pp. 212, 248. —Smith, 1931. Ent. News 42:21-22. —Gregg, 1956.
Ent. News 67:37-39 (also distribution).
macclendoni Wheeler. S. Tex. w. to Ariz.
Pheidole macclendoni Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:450. %, 2f.

Taxonomy: Cole, 1957. N. Y. Ent. Soc., Jour. 65:130-131 (worker, soldier). —Gregg, 1958. N.
Y. Ent. Soc., Jour. 66:46-47.
marcidula Wheeler. Tex. (Barton Cr., Austin).
Pheidole marcidula Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:457. %, 2.
megacephala (Fabricius). Fla.; W. Indies, Hawaii, and probably all tropical regions of the
world. Introduced, probably native to Africa but spread by commerce throughout the
world. Bigheaded ant.
Formica megacephala Fabricius, 1793. Ent. System. 2:361. 2.
Phidola megalocephala Schulz, 1906. Spolia Hym., p. 155. Emend.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:843-844. —Wheeler and Wheeler,
1953. Ent. Soc. Wash., Proc. 55:75 (larva). —Wilson and Taylor, 1967. Pacific Insects
Monog. 14:46-48.

Biology: Illingsworth, 1916. Hawaii. Ent. Soc., Proc. 3:349-368. —Phillips, 1934. (Hawaii
Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bul. 15:5-12. —Broekhuysen, 1948. Union
So. Africa Dept. Agr. Bul. 266:1-40. —Brown, 1958. Acta Hym. 1:47. —Brown, 1959. Bul.
Ent. Res. 50:523. —Weber, 1960. Ent. Soc. Wash., Proc. 62:232. —Taylor and Wilson, 1961.
Psyche 68:143. —Kempf, 1962. Studia Ent. 5:18-19. —Haskins and Haskins, 1965. Ecology
46:737 (competition between P. megacephala and Iridomyrmex humilis (Mayr) in
Bermuda). —Fluker, Huddleston, and Beardsley, 1968. Jour. Econ. Ent. 61:474. —Fluker
and Beardsley, 1970. Ent. Soc. Amer., Ann. 63:1290-1296 (sympatric association with
Iridomyrmex humilis and Anoplolepis longipes in Hawaii).

metallescens metallescens Emery. Fla. w. to Tex.
Pheidole metallescens Emery, 1895. Zool. Jahrb., Abt. f. System. 8:289, 294. &.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:476-477. —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62:12 (larva).
Biology: Smith, 1924. Ent. News 34:78 (as splendidula).
metallescens splendidula Wheeler. Southwest. Tex.
Pheidole metallescens splendidula Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:474. &,
2 ORGE
micula Wheeler. Ariz., s. Calif. Ecology: Nest was found under a stone.
Pheidole californica micula Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:408. %, 2f.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:33. —Wheeler and Wheeler, 1972. Ga. Ent.
Soc., Jour. 7:244 (larva).
Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 80.
militicida Wheeler. N. Mex., Ariz.
Pheidole militicida Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:398. %, 2f.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:9-12 (female, male;
biological notes).
Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:395. —Cole, 1953. Tenn. Acad. Sci., Jour. 28:298.
—Creighton and Creighton, 1959. Psyche 66:1-12 (observations on habits in s. Ariz.).
moerens Wheeler. Fla., Ala. (Mobile); Puerto Rico. Apparently introduced.
Pheidole moerens Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:136-138. %, 2.

Taxonomy: Wheeler and Wheeler, 1972. Ga. Ent. Soc., Jour. 7:244 (larva). —Wojcik, Banks,
and Buren, 1975. Coop. Econ. Ins. Rpt. 25 (49-52): 906 (first report in Fla.).

Biology: Smith, 1937. Puerto Rico Univ., Jour. Agr. 20:842-843.

morrisi impexa Wheeler. Okla., Tex.
Pheidole morrisi var. impexa Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:461. 8, 2, 2,
é.

1372 Hymenoptera in America North of Mexico

Biology: Lindquist, 1942. Jour. Econ. Ent. 35:850-852.
morrisi morrisi Forel. N. Y. to Fla., w. to Ill., La., Tex.
Pheidole Morrisii Forel, 1886. Soc. Ent. de Belg, Bul. 30:46. 8, 2f.
Pheidole morrisi var. Vanceae Forel, 1901. Soc. Ent. de Belg., ‘Ann. 45:351. 8, U, Q, é.
Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:461. —Smith, 1924. Ent. News
35:53. —Wheeler and Wheeler, 1960. Ent. Soc. Amer., Ann. 53:12 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:302. —Wheeler, 1905. Amer. Mus.
Nat. Hist., Bul. 21:380. —Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17:8-9.
nuculiceps Wheeler. Tex. (Canal River, New Braunfels).
Pheidole nuculiceps Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:473. 8, 2.

pilifera artemisia Cole. Utah, Ariz.
Pheidole pilifera artemisia Cole, 1933. Ent. Soc. Amer., Ann. 26:616. %, 2f.
Taxonomy: Cole, 1938. Amer. Midland Nat. 20:372 (female). —Cole, 1952. Tenn. Acad. Sci.,
Jour. 27:280. —Gregg, 1955. Psyche 62:22-23.
Biology: Cole, 1942. Amer. Midland Nat. 28:362.

pilifera coloradensis Emery. N. Dak., Colo., N. Mex., Nev. Ecology: A harvester of seeds of
various grasses; nests are usually under stones. Parasite: Pheidole inquilina (Wheeler).

Pheidole pilifera var. coloradensis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:288, 291. %,

Y.
Pheidole pilifera coloradensis var. neomexicana Wheeler, 1908. Amer. Mus. Nat. Hist.,

Bul. 24:436. 2.
Taxonomy: Cole, 1952. Tenn. Acad. Sci., Jour. 27:279. —Gregg, 1955. Psyche 62:22-23.
Biology: Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19:666. —Wheeler, 1904. Amer. Mus.
Nat. Hist., Bul. 20:16-17. —Wheeler, 1910. Ants, pp. 279, 498. —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 128-130. —Gregg, 1963. Ants of Colo., pp. 419, 421-422. —Cole,
1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7(8):15.
pilifera pacifica Wheeler. Nev., Calif.
Pheidole xerophila pacifica Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:404. 3, 2f, 2, d.
Taxonomy: Cole, 1952. Tenn. Acad. Sci., Jour. 27:279. —Gregg, 1955. Psyche 62:22-23.
pilifera pilifera (Roger). N. Y., Mass. s. to Fla., w. to N. Dak., Nebr., Kans. Ecology: A
harvester of seeds; most nests have been found in exposed soil and have craterlike
excavations. Parasite: Pheidole inquilina (Wheeler).
Leptothorax pilifer Roger, 1863. Berlin. Ent. Ztschr. 7:180. %.
Pheidole pennsylvanica Roger, 1863. Berlin. Ent. Ztschr. 7:199. °.
Pheidole pilifera septentrionalis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:436. 2.
Pheidole pilifera var. simulans Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:436. 2.
Taxonomy: Cole, 1952. Tenn. Acad. Sci., Jour. 27:280. —Wheeler and Wheeler, 1953. Ent. Soc.
Wash., Proc. 55:79 (larva). —Gregg, 1955. Psyche 62:22-23.
Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:379-380. —Wheeler, 1910. Ants, pp.
152, 278. —Wesson and Wesson, 1940. Amer. Midland Nat. 24:92. —Wheeler and Wheeler,

1963. Ants of N. Dak., pp. 130-131.
pinealis Wheeler. Tex.; Mexico. Ecology: Nests have been found under stones.
Pheidole pinealis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:459. , 2.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:12-15 (also biological

notes).

porcula Wheeler. W. Tex., Colo. (7).
Pheidole crassicornis porcula Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:466. %, 2.

psammophila Creighton and Gregg. Ariz., s. Calif.; Mexico. Ecology: Found in sandy areas at

low elevations.
Pheidole psammophila Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:15-19.

Oreee
Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 81.

Superfamily FORMICOIDEA 1373

rhea Wheeler. S. Ariz.; Mexico. Ecology: Prefers to nest on plateaus or in foothills at the base
of mountains, 3700 to 7000 ft.
Pheidole rhea Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:452. °.

Taxonomy: Smith, 1943. Ent. Soc. Wash., Proc. 45:5-9 (worker, soldier). —Gregg, 1949.
Psyche 56:70-73.
Biology: Gregg, 1949. Psyche 56:70-73.
ridicula Wheeler. Tex. (Brownsville area).
Pheidole ridicula Wheeler, 1916. New England Zool. Club, Proce. 6:29. YU.
Biology: Creighton, 1966. Psyche 73:1.
rugulosa Gregg. Tex., Ariz.
Pheidole rugulosa Gregg, 1958. N. Y. Ent. Soe., Jour. 66:26-29. 3, 2f, 2, d.
sciara Cole. Tex., N. Mex.
Pheidole sciara Cole, 1955. Tenn. Acad. Sci., Jour. 30:47-49. 3, 2.
Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31:116.

sciophila Wheeler. Tex. to s. Ariz.; Mexico.
Pheidole sciophila Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:443. 8, 2, °, or
Pheidole proserpina Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:437. 8, 2.
Pheidole sciophila var. semilaevicephala Smith, 1934. Ent. Soc. Amer., Ann. 27:385. 2.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:19-22.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:397.

senex Gregg. Tex., Colo., N. Mex.
Pheidole senex Gregg, 1952. Amer. Mus. Novitates 1557:1-4. 8, 2.
Pheidole pilifera anfracta Cole, 1952. Tenn. Acad. Sci., Jour. 27:278. 9, 2.

Taxonomy: Gregg, 1955. Psyche 62:22-23.

Biology: Gregg, 1963. Ants of Colo., pp. 422-424.

sitarches campestris Wheeler. Miss., Mo. w. to Colo., central Tex.
Pheidole sitarches rufescens var. campestris Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.
24:443. 9, 2.
Pheidole sitarches rufescens Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:443. 3, 2, °.

Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:37.

Biology: Smith, 1924. Ent. News 35:53. —Gregg, 1963. Ants of Colo., pp. 424-425.
sitarches littoralis Cole. Fla. (Lido Beach, Sarasota).
Pheidole sitarches littoralis Cole, 1952. Ent. Soc. Amer., Ann. 45:443-444. &, 2.
Taxonomy: Gregg, 1958. N. Y. Ent. Soc., Jour. 66:39.

sitarches sitarches Wheeler. Tex. (Brownsville area); Mexico.
Pheidole sitarches Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:440. 8, 2, 2.
Pheidole sitarches var. transvarians Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:442. &,
ae

Taxonomy: Cole, 1952. Ent. Soc. Amer., Ann. 45:444. —Gregg, 1958. N. Y. Ent. Soe., Jour.
66:39.
Biology: Wheeler, 1910. Ants, p. 279. —Wilson, 1957. Psyche 64:46-50 (nuptial flights).
sitarches soritis Wheeler. W. Tex., N. Mex., Utah, Ariz.; Mexico.
Pheidole soritis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:439. &, aie
Pheidole tepicana cavigenis Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:403. 2.
Taxonomy: Cole, 1952. Ent. Soc. Amer., Ann. 45:444. —Gregg, 1958. N. Y. Ent. Soc., Jour.
66:37.
spadonia Wheeler. S. Ariz.; Mexico.
Pheidole spadonia Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:400. %, 2.
Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:22-24 (also biological
notes).

1374 Hymenoptera in America North of Mexico

subdentata Pergande. Ariz.; Mexico.
Pheidole subdentata Pergande, 1895. Calif. Acad. Sci., Proc. 5:888. 9.
Pheidole obtusospinosa Pergande, 1895. Calif. Acad. Sci., Proc. 5:889. 2.
Pheidole arizonica Santschi, 1909. Soe. Ent. Ital., Bol. 41:3. 2.

Taxonomy: Creighton, 1957. N. Y. Ent. Soc., Jour. 65:211.
tepicana Pergande. Tex., Ariz.; Mexico. A polymorphic species with the largest soldiers and

smallest workers connected by intermediates, unlike the dimorphic worker castes of
most Pheidole.

Pheidole tepicana Pergande, 1895. Calif. Acad. Sci., Proc. 5:878. %, 2.

Pheidole rugifrons Pergande, 1895. Calif. Acad. Sci., Proc. 5:880. 2.

Pheidole carbonaria Pergande, 1895. Calif. Acad. Sci., Proc. 5:881. %, 2.

Pheidole Kingi Andre, 1898. Soc. Ent. de France, Bul. p. 244. %, 2.

Pheidole Townsendi Andre, 1898. Soc. Ent. de France, Bul., p. 246. %, 2.

Pheidole kingi instabilis Emery, 1901. Soe. Ent. de France, Bul., p. 129. %, 2.

Pheidole kingi torpescens Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:404. 9, 2.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Wash., Proc. 55:74 (larva). —Creighton and
Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:24-35.
Biology: Wheeler, 1901. Soc. Ent. de Belg., Ann. 45:203. —Wheeler, 1907. Amer. Mus. Nat.
Hist., Bul. 23:2-12. —Cole, 1956. Tenn. Acad. Sci., Jour. 31:113.
Morphology: Wheeler, 1910. Ants, p. 56.
texana Wheeler. Tex. (Travis Co.).
Pheidole texana Wheeler, 1903. Psyche 10:95. 8, 2.
Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:464.
titanis Wheeler. W. Tex. to s. Ariz. Ecology: Diet may be restricted to termites.
Pheidole titanis Wheeler, 1903. Psyche 10:95. 8, 2.
Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:461462. —Creighton and Gregg,
1955. Colo. Univ. Studies, Biol. Ser. 3:35-37 (also biological notes; termite feeding).
tysoni Forel. Va., N. C., Ohio, Tenn.
Pheidole tysoni Forel, 1901. Soc. Ent. de Belg., Ann. 45:348. 3, 2, 3.
Taxonomy: Cole, 1940. Amer. Midland Nat. 24:42, 45.
Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:282, 304. —Wesson and Wesson, 1940. Amer.
Midland Nat. 24:92.

vallicola Wheeler. S. Ariz.
Pheidole crassicornis vallicola Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:409. %, 2f.
virago Wheeler. Tex., Ariz.
Pheidole virago Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:401. %, 2.
vistana Forel. S. Calif.; Mexico. Ecology: Found in deserts in small crater nests in the soil.
Pheidole susannae longipes Pergande, 1895. Calif. Acad. Sci., Proc. 5:885. %, 2{. Preoce. by
Latreille, 1802; Smith, 1858.
Pheidole longipes var. vistana Forel, 1914. Soc. Vaud. des Sci. Nat., Bul. 50:272. °.
Pheidole grallipes Wheeler, 1916. Psyche 23:40. N. name for longipes Pergande.

Taxonomy: Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:397-398.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:69. —Wheeler and Wheeler, 1973. Ants
of Deep Canyon, pp. 77-79.
xerophila tucsonica Wheeler. Tex., N. Mex., Ariz., Calif.; Mexico. Ecology: Nests in soil may
be craterlike or with small mounds.
Pheidole xerophila tucsonica Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:448. 8, 2.
Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:40-42.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27:397. —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 82-83.

Superfamily FORMICOIDEA 1375

xerophila xerophila Wheeler. Tex., N. Mex., s. Calif. Ecology: Found in small crater nests in
the desert soil. The range of this subspecies lies slightly to the south of that of
tucsonica in Tex. and N. Mex.
Pheidole xerophila Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:446. 3, 2, d.

Taxonomy: Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:40-42.
Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 82.
yaqui Creighton and Gregg. S. Calif.; Mexico. Ecology: Found in deserts in small crater nests
in the soil.
Pheidole yaqui Creighton and Gregg, 1955. Colo. Univ. Studies, Biol. Ser. 3:43-46. %, 2.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 83-84.
TRIBE CARDIOCONDYLINI

Genus CARDIOCONDYLA Emery

Cardiocondyla Emery, 1869. Accad. degli Aspiranti Naples, Ann. 2:20.
Type-species: Cardiocondyla elegans Emery. Monotypic.
Emeryia Forel, 1890. Soc. Ent. de Belg. Ann. (C. R.) 34:110.
Type-species: Emeryia wroughtoni Forel. Monotypic.
Xenometra Emery, 1917. Soc. Ent. de France, Bul., p. 96.
Type-species: Xenometra monilicornis Emery. Monotypic.
Cardiocondyla subg. Dyclona Santschi, 1930. Rev. Suisse de Zool. 37:70. Syn. questionable.
Type-species: Monomorium cristatum Santschi. Orig. desig.
Cardiocondyla subg. Loncyda Santschi, 1930. Rev. Suisse de Zool. 37:70. Syn. questionable.
Type-species: Cardiocondyla (Loncyda) monardi Santschi. Monotypic.
Cardiocondyla subg. Prosopidris Wheeler, 1935. Psyche 42:40.
Type-species: Cardiocondyla (Prosopidris) sima Wheeler. Orig. desig.

The majority of species of this genus are found in the warmer parts of Europe, Asia, and
Africa. In the United States, members of this genus are found in the southern tier of states
from Florida to California. The colonies are small and nests are constructed in soil and in plant

cavities. They are easily transported by commerce and all the species below may have been in-
troduced.

Revision: Smith, 1944. Ent. Soc. Wash., Proc. 46:30-41 (U.S. species).

Taxonomy: Weber, 1952. Amer. Mus. Novitates 1548:5. —Brown, 1973. In Meggers, et al.,
Tropical forest ecosystems in Afr. and S. Amer., pp. 161-185 (generic syn.).

Biology: Wilson, 1960. Psyche 66:29-34 (tandem running).
ectopia Snelling. Ariz., s. Calif. Possibly of Old World origin.
Cardiocondyla ectopia Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 76-81, figs. 3, 2, d.

Biology: Creighton and Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 87-91 (behavior).
emeryi Forel. Fla., Tex.; W. Indies, Africa, Asia, Pacific Islands. Probably introduced. A tramp
species distributed by commerce throughout the tropics.
Cardiocondyla emeryi Forel, 1881. Munchen Ent. Ver., Mitt. 5:5. 9.

Cardiocondyla nuda subsp. nereis Wheeler, 1927. Amer. Acad. Arts and Sci., Proc. 62:140.
Gaon

Taxonomy: Borgmeier, 1937. Rev. de Ent. 7:133 (ergatoid male). —Weber, 1952. Amer. Mus.
Novitates 1548:5. —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:53 (Polynesia).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:89. —Smith, 1936. Puerto Rico Univ.,
Jour. Agr. 20:835. —Creighton and Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 82-86
(behavior).

nuda (Mayr). Fla, Ga. La, Tex.; Pacific Islands. Probably introduced into U. S. from the
Oriental Region.
Leptothorax nudus Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. K1., Sitzber. 53:508. °.
Cardiocondyla nuda var. minutior Forel, 1899. Fauna Hawaiiensis, 1:120. 9.

1376 Hymenoptera in America North of Mexico

Taxonomy: Wilson and Taylor, 1967. Pacific Ins. Monog. 15:55 (Polynesia). —Wheeler and
Wheeler, 1973. Ga. Ent. Soce., Jour. 8:27 (larva).

Biology: Phillips, 1934. (Hawaii Univ.) Expt. Sta., Pineapple Prod. Coop. Assoc. Bul. 15:22.
—Creighton and Snelling, 1974. N. Y. Ent. Soc., Jour. 82: 86-87 (behavior).
venustula Wheeler. Fla., La.; Puerto Rico, Haiti. Probably introduced.
Cardiocondyla venustula Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:128. 3, @.

Biology: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:836. —Smith, 1944. Fla. Ent. 27:15.
wroughtonii (Forel). Fla., Ga.; Oriental Region. Probably introduced.
Emeryia wroughtonii Forel, 1890. Soc. Ent. de Belg., Ann. (C. R.) 34:110. “3” =
ergatomorphic d.
Cardiocondyla wroughtonii var. hawaiensis (!) Forel, 1899. Fauna Hawaiiensis 1:119. &.
Cardiocondyla wroughtoni var. bimaculata Wheeler, 1929. Lab. Zool. Gen. e Agr. Portici,
Bol. 24:43. 8, @.

Taxonomy: Wilson and Taylor, 1967. Pacific Ins. Monog. 14:56 (Polynesia).
Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:7. —Smith, 1933. Fla. Ent. 17:24.

TRIBE CREMATOGASTRINI

Genus CREMATOGASTER Lund

Crematogaster Lund, 1831. Ann. des Sci. Nat., Zool. 23:132.

Type-species: Formica scutellaris Olivier. Desig. by Bingham, 1903.
Cremastogaster Agassiz, 1846. Nomencl. Zool., Index Univ., p. 103. Emend.
Acrocoelia Mayr, 1852. Zool.-Bot. Gesell. Wien, Verh. 2:147.

Type-species: Acrocoelia ruficeps Mayr. Desig. by Wheeler, 1911.
Crematogaster subg. Oxygyne Forel, 1901. Soc. Ent. de Belg., Ann. 45:375.

Type-species: Crematogaster (Oxygyne) daisyi Forel. Desig. by Wheeler, 1911.
Decacrema Forel, 1910. Soc. Ent. de Belg., Ann. 54:18.

Type-species: Crematogaster schencki Forel. Desig. by Wheeler, 1911.
Crematogaster subg. Atopogyne Forel, 1911. Soc. Vaud. des Sci. Nat., Bul. 47:342.

Type-species: Crematogaster (Atopogyne) hellanica Forel. Desig. by Wheeler, 1911.
Physocrema Forel, 1912. Soc. Ent. de Belg., Mem. 14:220.

Type-species: Crematogaster inflatus Smith. Desig. by Wheeler, 1911.
Xiphocrema Forel, 1913. Zool. Jahrb., Abt. f. System. 36:80.

Type-species: Crematogaster tetracantha Emery. Desig. by Emery, 1922.
Eucrema Santschi, 1918. Soc. Ent. de France, Bul., p. 182.

Type-species: Formica acuta Fabricius. Orig. desig.

Nematocrema Santschi, 1918. Soc. Ent. de France, Bul. p. 182.

Type-species: Crematogaster stadelmanni Mayr. Orig. desig.
Neocrema Santschi, 1918. Soc. Ent. de France, Bul., p. 182.

Type-species: Crematogaster distans Mayr. Orig. desig.
Orthocrema Santschi, 1918. Soc. Ent. de France, Bul. p. 182.

Type-species: Myrmica sordidula Nylander. Orig. desig.
Paracrema Santschi, 1918. Soc. Ent. de France, Bul. p. 182.

Type-species: Crematogaster spengeli Forel. Orig. desig.
Sphaerocrema Santschi, 1918. Soc. Ent. de France, Bul. p. 182.

Type-species: Crematogaster kneri Mayr. Orig. desig.

Crematogaster subg. Rhachiocrema Mann, 1919. Harvard Univ., Mus. Comp. Zool., Bul.
63:318.

Type-species: Crematogaster (Rhachiocrema) wheeleri Mann. Orig. desig.
Tranopeltoides Wheeler, 1922. Amer. Mus. Novitates 48:10.

Type-species: Tranopelta huberi Forel. Orig. desig.

Crematogaster subg. Colobocrema Wheeler, 1927. Quart. Rev. Biol. 2:31.

Type-species: Crematogaster (Colobocrema) cylindriceps Wheeler. Monotypic.
Mesocrema Santschi, 1928. Soc. Ent. de Belg., Bul. 68:33.

Type-species: Crematogaster rasoherini Forel. Desig. by Donisthorpe, 1943.
Apterocrema Wheeler, 1936. Psyche 43:45.

Superfamily FORMICOIDEA 1377

Type-species: Apterocrema atillani Wheeler. Monotypic.

These ants nest in moderately large colonies under objects in the soil, in wood, in cavities of
plants, in insect galls, in carton nests of their own making, or even in the woodwork of buildings.
Most are omnivorous but show a preference for sweets and some tend honeydew excreting in-
sects. Some are well known house-infesting forms; others have been reported to kill young birds
or gnaw rubber insulation from telephone wires. North American species of this genus had been
separated into two subgenera, subgenus Orthocrema for arizonensis and minutissima and its
subspecies and subgenus Crematogaster for the remaining species. Subgenera are not recog-
nized here.

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:989-996. —Emery, 1895. Zool. Jahrb.,
Abt. f. System. 8:280-288. —Wheeler, 1919. Psyche 26:111. —Creighton, 1939. Psyche
46:137-140 (subg. Orthocrema). —Buren, 1968. Ga. Ent. Soc., Jour. 3:91-121 (subg.
Crematogaster).

Taxonomy: Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:258-261 (larvae). —Buren,
1958. N. Y. Ent. Soe., Jour. 66:119-134 (also biological notes). —Brown, 1973. In Meggers,
et al., Tropical Forest Ecosystems in Afr. and S. Amer., pp. 161-185 (generic syn.).
—Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:27-30 (larvae).

Biology: Soulie, 1962. Insectes Sociaux 9:181-195 (colony foundation and development).
arizonensis Wheeler. S. Ariz. Ecology: Apparently arboreal; found nesting in mistletoe on

oaks.
Crematogaster arizonensis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:482. 9.

Biology: Wheeler, 1912. N. Y. Ent. Soc., Jour. 20:130-133 (also description of female, male).
ashmeadi Mayr. Va. to Fla., w. to Tex. Ecology: Strictly arboreal; nests in twigs and branches.
Crematogaster Ashmeadi Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:463. %, d.
Crematogaster (Acrocoelia) ashmeadi var. matura Wheeler, 1932. N. Y. Ent. Soc., Jour.
40:8. 8.

Taxonomy: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:8.

Biology: Smith, 1924. Ent. News 35:79. —Cole, 1940. Amer. Midland Nat. 24:46. —Smith,
1965. U. S. Dept. Agr., Tech. Bul. 1326:28-29 (economic importance). —Leuthold, 1968.
Psyche 75:233-248 (tibial gland scent-trail and trail-laying behavior). —Leuthold, 1968.
Psyche 75:334-350 (recruitment to food).

atkinsoni Wheeler. N. C. s. to Fla., w. to Miss. Ecology: Common in coastal salt-grass marshes
where they often build large carton nests on sedges or bushes. Nests are sometimes a
foot and a half or more above the ground and range in size from the diameter of an egg
to that of a human head.
Crematogaster atkinsoni Wheeler, 1919. Psyche 26:108. &.
Crematogaster atkinsoni var. helveola Wheeler, 1919. Psyche 26:109. % (?, d misdet.).
The female and male are C. ashmeadi.

Biology: Atkinson, 1887. Amer. Nat. 21:770-771. —Smith, 1930. Fla. Ent. 14:4-5,. —Wheeler,
1932. N. Y. Ent. Soc., Jour. 40:8-9.
browni Buren. W. Tex., N. Mex., s. Ariz. Ecology: In mountains, usually over 5000 ft.; nests

have been found under rocks.
Crematogaster browni Buren, 1968. Ga. Ent. Soc., Jour. 3:100. 8, °.

californica Emery. S. Calif.; Mexico. Ecology: Nests in soil in desert or semidesert habitats;
found at bases and on roots of various plants cultivating aphids and coccids. Males are

unusually large for this genus.
Crematogaster lineolata laeviuscula var. californica Emery, 1895. Zool. Jahrb., Abt. f.

System. 8:285. 8.
Taxonomy: Wheeler, 1934. Pan-Pacifie Ent. 10:135-136.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:70. —Wheeler and Wheeler, 1973. Ants
of Deep Canyon, pp. 87-88.

1378 Hymenoptera in America North of Mexico

cerasi (Fitch). Que. s. to Ga., w. to S. Dak., Ark., N. Mex. Ecology: Nests under rocks and logs;
sometimes does minor damage when nesting in woodwork and timbers in houses.
Myrmica cerasi Fitch, 1855 (1854). N. Y. State Agr. Soc., Trans. 14:835. %.
Crematogaster (Acrocoelia) kennedyi Wheeler, 1930. Psyche 37:58. 2, d.
Crematogaster (Acrocoelia) lineolata cerasi var. punctinodis Enzmann, 1946. N. Y. Ent.
Soce., Jour. 54:91, 93, 96. 3, 2, 3.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:282-283. —Wheeler, 1933. Psyche
40:83-84. —Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:250-252 (larva, as
lineolata). —Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:28 (larva).

Biology: Gaige, 1914. Mich. Univ., Mus. Zool., Occas. Papers 5:8-9. —Headley, 1943. Ohio

Jour. Sci. 43:25. —Morris, 1943. Ind. Acad. Sci., Proce. 52:208. —Wheeler and Wheeler, 1944.

N. Dak. Hist. Quart. 11:245. —Kannowski, 1959. Insectes Sociaux 6:125. —Kulman, 1965.

Jour. Econ. Ent. 58:865 (in cocoons of Thyridopteryx ephemeraeformis). —Smith, 1965. U.

S. Dept. Agr., Tech. Bul. 1326:30-31 (economic importance).

clara Mayr. N. J. s. to Fla., w. to Ind., Mo., Tex. Ecology: Nests in a variety of semi-arboreal

situations in or near swamps, salt marshes, rivers, and streams.

Oecodoma (Atta) bicolor Buckley, 1867. Ent. Soc. Phila., Proc. 6:350. 3. Preoce. by Smith,
1860.

Crematogaster clara Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:990, 993. &.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:481 (each caste). —Smith, 1924.
Ent. News 35:80. —Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:255-256 (larva,
as laeviuscula).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:283. —Smith, 1965. U. S. Dept. Agr., Tech.
Bul. 1326:31-32 (economic importance).

Morphology: Pasteels, Crews, and Blum, 1970. Acad. Soc. Paris, Compt. Rend., Ser. D
271:835-838 (histology of gland secreting the trail pheromone).
coarctata Mayr. Oreg.(?), Nev., Calif. Ecology: Nests under rocks and boulders. Suspected of
now being a rather rare species in coastal Calif. due to displacement by the Argentine
ant.
Crematogaster coarctata Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:990, 992. 9.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:482. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bul. 104:207-208.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:70-71. —Essig, 1926. Ins. of West. N.
Amer., p. 859.
colei Buren. W. Tex., N. Mex., Ariz. Ecology: Nests under rocks in grassland situations at
elevations over 5000 ft.
Crematogaster colei Buren, 1968. Ga. Ent. Soc., Jour. 3:108. %, 2.
dentinodis Forel. Ariz.; Mexico. Ecology: Nests under stones in open grasslands and chaparral,
or grass and mesquite, or among shrubs in succulent desert areas.
Crematogaster opaca var. dentinodis Forel, 1901. Soc. Ent. de Belg., Ann. 45:130. %
depilis Wheeler. Tex., N. Mex., Ariz., Nev., s. Calif.; Mexico. Ecology: Nests in.and among
roots and lower stems of various plants throughout desert and semi-desert regions.
Crematogaster lineolata opaca var. depilis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.
24:478. %.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104:209 (worker, not
female). —Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:28-30 (larva).

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7:16. —Wheeler and Wheeler,
1973. Ants of Deep Canyon, p. 88.
emeryana Creighton. W. Tex., Colo., N. Mex., Utah, Ariz.; Mexico. Ecology: Usually in
mountains at elevations of over 6000 ft.; often constructs carton structures under rocks.
Crematogaster lineolata lineolata var. Emery, 1895. Zool. Jahrb., Abt. f. System. 8:282. %,
?, 6. Variety described but not named.

Superfamily FORMICOIDEA 1379

Crematogaster (Acrocoelia) lineolata emeryana Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bul. 104:213-214. &.

Biology: Gregg, 1963. Ants of Colo., pp. 358-360.
hespera Buren. W. Tex., N. Mex., Utah, Ariz., Calif.; Mexico. Ecology: Usually arboreal in
cottonwood trees along rivers and streams; also in other trees and rotten logs.
Crematogaster hespera Buren, 1968. Ga. Ent. Soc., Jour. 3:98. 3, °.
isolata Buren. W. Tex., N. Mex., s. Ariz. Ecology: Arboreal in oaks in mountain ranges at
altitudes over 5000 ft.
Crematogaster isolata Buren, 1968. Ga. Ent. Soc., Jour. 3:106. %.
laeviuscula Mayr. La., Tex., Okla.; Mexico. Ecology: Incipient colonies are often found in twigs
and oak galls, larger colonies in hollow trees and logs. No authentic records east of
Mississippi River.
Oecodoma (Atta) arborea Buckley, 1867. Ent. Soc. Phila., Proc. 6:349-350. 3, 2. Preoce. by
Smith, 1858.
Crematogaster laeviuscula Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:990, 993. &.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:481 (each caste). —Cole, 1940.
Amer. Midland Nat. 24:46.
larreae Buren. W. Tex., Ariz., Calif.; Mexico. Ecology: Nests in lower stems and among roots
of the creosote bush. Host: Larrea divaricata.
Crematogaster larreae Buren, 1968. Ga. Ent. Soc., Jour. 3:117. %, @, d.

lineolata (Say). Que., Ont. s. to Fla., w. to N. Dak., Colo., Tex. Ecology: Nests in ground under

objects or in logs or stumps; infests houses and often nests within buildings.

Myrmica lineolata Say, 1836. Boston Jour. Nat. Hist. 1:290. %, 2, d.

Myrmica (Monomarium (!)) marylandica Buckley, 1867. Ent. Soc. Phila., Proc. 6:339. %.
Syn. uncertain.

Myrmica (Monomarium (!)) columbiana Buckley, 1867. Ent. Soc. Phila., Proc. 6:340. %, 9.
Syn. uncertain.

Crematogaster lineolata lineolata var. lutescens Emery, 1895. Zool. Jahrb., Abt. f. System.
8:282. &.

Crematogaster (Acrocoelia) lineolata cerasi var. wheldeni Enzmann, 1946. N. Y. Ent. Soc.,
Jour. 54:92-93, 96. %.

Taxonomy: Smith, 1918. Ent. News 29:19. —Buren, 1944. Iowa State Col., Jour. Sci. 18:288.
—Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:252 (larva as lineolata subopaca).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:1-18. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bul. 22:585. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:93. —Wallace, 1945. Conn. State Ent., 44th Rpt., Bul. 488:389 (economics). —Wheeler
and Wheeler, 1963. Ants of N. Dak., pp. 132-133. —Gregg, 1963. Ants of Colo., pp. 357, 359.
—Ayre, 1963. Canad. Ent. 95:712-715 (feeding habits). —Smith, 1965. U. S. Dept. Agr.,
Tech. Bul. 1326:32-33 (economic importance). —Ayre, 1968. Canad. Ent. 100:165-172 (prey
finding, capture, transport). —Ayre, 1969. Canad. Ent. 101:118-128 (trail formation and
group foraging).

marioni Buren. S. Calif.; Mexico. Ecology: Arboreal species in manzanita and live oaks.
Crematogaster marioni Buren, 1968. Ga. Ent. Soc., Jour. 3:105. &.
minutissima minutissima Mayr. N. C. s. to Fla., w. to Tex. Ecology: Nests have been found in
soil at bases of stumps.
Crematogaster minutissima Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:991, 995. , 2.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:484. —Wheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:14 (larva).

Biology: Smith, 1928. Ent. News 39:277.

Morphology: Pasteels, Crewe, and Blum, 1970. Paris Acad. Soc., Compt. Rend., Ser. D
271:835-838 (histology of gland secreting the trail pheromone).

1380 Hymenoptera in America North of Mexico

minutissima missouriensis Emery. Nebr., Mo., Tex., Colo., N. Mex.
Crematogaster victima missuriensis (!) Emery, 1895. Zool. Jahrb., Abt. f. System. 8:287.

%.
Taxonomy: Wheeler and Wheeler, 1952. Wash. Acad. Sci., Jour. 42:260-261 (larva).

Biology: Gregg, 1963. Ants of Colo., pp. 363-364.
minutissima smithi Creighton. Ariz. (Huachuca Mtns.).
Crematogaster (Orthocrema) minutissima thoracica Creighton, 1939. Psyche 46:138. &.
Preocc. by Santschi, 1921.
Crematogaster (Orthocrema) minutissima smithi Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bul. 104:205. N. name.
mormonum Emery. Utah, Idaho, e. Oreg. (?), Nev., Calif.; Mexico. Ecology: Most common in
the Great Basin; nests under rocks.
Crematogaster lineolata coarctata var. mormonum Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:284. &.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:482 (each caste).

Biology: Cole, 1942. Amer. Midland Nat. 28:363. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, pp. 88-89.
mutans Buren. Nev., Calif. Ecology: The only North American species of this genus suspected
of being parasitic.
Crematogaster mutans Buren, 1968. Ga. Ent. Soc., Jour. 3:115. 8, @.
navajoa Buren. N. Mex., Ariz. in the Colorado Plateau region. Ecology: Found under rocks and
at bases of various plants.
Crematogaster navajoa Buren, 1968. Ga. Ent. Soc., Jour. 3:102. %, ?.
nocturna Buren. S. Utah, n. Ariz.
Crematogaster nocturna Buren, 1968. Ga. Ent. Soc., Jour. 3:112. 2, d, possible &.
opaca Mayr. Ariz. (Tumacacori Mts., under stones among oaks, 3900 ft.); Mexico. Only a single
Arizona record for the U. S.
Crematogaster opaca Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:989. §.
opuntiae Buren. N. Mex., s. Ariz. Ecology: Found principally at bases of and foraging on
Opuntia fulgida but also at bases of various other plants.
Crematogaster opuntiae Buren, 1968. Ga. Ent. Soc., Jour. 3:120. 8.
pilosa Emery. N. J. s. to Ga. Ecology: Often nests in logs and fallen branches in marshy
situations.
Crematogaster lineolata pilosa Emery, 1895. Zool. Jahrb., Abt. f. System. 8:285. &.
Crematogaster lineolata subpilosa Wheeler, 1913. Psyche 20:115. Syn. uncertain. Nomen
nudum.
Crematogaster (Acrocoelia) creightoni Wheeler, 1933. Psyche 40:86. °.

Taxonomy: Wheeler, 1933. Psyche 40:85.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:379. —Wheeler, 1906. Amer. Mus.
Nat. Hist., Bul. 22:6-7.
punctulata Emery. Va. s. to Fla., w. to Colo., Ariz.; Mexico. Ecology: Abundant in southern

Great Plains region where they nest in the ground and tend aphids and coccids.

Crematogaster punctulata Emery, 1895. Zool. Jahrb., Abt. f. System. 8:287. %.

Crematogaster lineolata lineolata var. subopaca Emery, 1895. Zool. Jahrb., Abt. f. System.
8:2334 Geer

Crematogaster (Acrocoelia) opaca var. texana Santschi, 1929. Wien Ent. Ztg. 46:91. 9.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:479-480.

Biology: Smith, 1927. Ent. News 38:310. —Gregg, 1963. Ants of Colo., pp. 361-362, 364.
rifelna Buren. S. Tex.;Mexico. Ecology: Arboreal, found nesting in live oaks and other trees.
Crematogaster rifelna Buren, 1968. Ga. Ent. Soc., Jour. 3:96. 8, 2, d.

Superfamily FORMICOIDEA 1381

vermiculata Emery. N. C. s. to Fla., w. to Ark., La. Ecology: Arboreal, found only in cypress
swamps.
Crematogaster vermiculata Emery, 1895. Zool. Jahrb., Abt. f. System. 8:286. %. Types
mislabeled, not found in Calif. or other western states.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:13 (larva).
TRIBE SOLENOPSIDINI

Genus MONOMORIUM Mayr

Monomorium Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:452.
Type-species: Monomorium minutum Mayr. Monotypic.
Phacota Roger, 1862. Berlin. Ent. Ztschr. 6:260. Uncertain syn.
Type-species: Phacota sicheli Roger. Monotypic.
Trichomyrmex Mayr, 1865. Novara Exped., Zool., Formicidae, pt. 2, v. 1, p. 19.
Type-species: Trichomyrmex rogeri Mayr. Monotypic.
Lampromyrmex Mayr, 1868. Beitr. Naturk. Preuss. 1:92.
Type-species: Lampromyrmesx gracillimus Mayr. Monotypic.
Holcomyrmex Mayr, 1878. Zool.-Bot. Gesell. Wien, Verh. 28:671.
Type-species: Holocomyrmesx scabriceps Mayr. Desig. by Bingham, 1903.
Epoecus Emery, 1892. Soc. Ent. de France, Bul. 61: CCLXXVI.
Type-species: Epoecus pergandei Emery. Monotypic.
Wheeleria Forel, 1905. Soc. Ent. de Belg., Ann. 49:171. Preoce. by Tutt, 1895.
Type-species: Wheeleria sanischii Forel. Monotypic.
Wheeleriella Forel, 1907. Inst. Sci. Revue 4:145. N. name for Wheeleria Forel.
Epixenus Emery, 1908. Deut. Ent. Ztschr., p. 556.
Type-species: Epixenus andrei Emery. Desig. by Wheeler, 1911.
Monomorium subg. Xeromyrmex Emery, 1915. Ent. Soc. de France, Bul. p. 190.
Type-species: Formica salomonis Linnaeus. Orig. desig.
Monomorium subg. Parholcomyrmex Emery, 1915. Ent. Soc. de France, Bul. p. 190.
Type-species: Myrmica gracillima Smith. Orig. desig.
Monomorium subg. Paraholcomyrmex Emery, 1915. Soe. Ent. de France, Bul. p. 191.
Misspelling ?.
Mitara Emery, 1913. Soc. Ent. de Belg., Ann. 57:261.
Type-species: Monomorium laeve Mayr. Orig. desig.
Monomorium subg. Corynomyrmex Viehmeyer, 1916. Arch. f. Naturgesch. 18:134.
Type-species: Monomorium (Corynomyrmex) hospitum Viehmeyer. Monotypic.
Monomorium subg. Isolcomyrmex Santschi, 1917. Soc. Cient. Argentina, An. 84:296.
Type-species: Holcomyrmex santschii Forel. Orig. desig.
Paraphacota Santschi, 1919. Soc. Ent. de France, Bul., p. 91.
; Type-species: Phacota noualhieri Emery. Orig. desig.
Monomorium subg. Equestrimessor Santschi, 1919. Soc. Ent. de France, Bul. p. 92.
Type-species: Holcomyrmesx chobauti Emery. Desig. by Donisthorpe, 1943.
Xenhyboma, Santschi, 1919. Soc. Espan. de Hist. Nat., Bol. 19:405. Uncertain syn.
Type-species: Xenhyboma mystes Santschi. Monotypic.
Equessimessor Santschi, 1936. Bul. Soc. Sci. Nat. Maroc. 16:32. Emend.
Isholcomyrmex Santschi, 1936. Bul. Soc. Sci. Nat. Maroc. 16:32. Variant spelling.
Ireneidris Donisthorpe, 1943. Ent. Monthly Mag. 79:81.
Type-species: Ireneidris myops Donisthorpe. Monotypic.
Monomorium subg. Pharaophanes Bernard, 1952. Inst. France. d’Afr. Noire, Mem. 19:238.
Nomen nudum.
Xenoaphaenogaster Baroni Urbani, 1964. Atti della Accad. Gioenia di Sci. Nat. Catania
16:50. Uncertain syn.
Type-species: Xenoaphaenogaster inquilina Baroni Urbani. Orig. desig.

Members of this genus are small in size and are particularly adaptable with respect to nesting
sites. They may nest in various preformed cavities or in soil. Most species are Old World, but
because of their adaptability and size, they are especially susceptible to distribution by com-

1382 Hymenoptera in America North of Mexico

merce. Three of the North American species have been introduced and are common tramp spe-
cies and house pests.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:121-122 (larvae).
—Ettershank, 1966. Austral. Jour. Zool. 14:82-93 (generic syn., list of world species).
—Brown, 1973. In Meggers, et al., Tropical Forest Ecosystems in Afr. and S. Amer., pp.
161-185 (generic syn.). —Wheeler and Wheeler, 1973. Ga. Ent. Soc., Jour. 8:30-31 (larvae).

Morphology: Blum, 1966. Roy. Ent. Soc. London, Proc. (A) 41: 155-160 (source and specificity
of trail pheromones).
destructor (Jerdon). Tenn., Fla.; throughout tropical regions of world. Ecology: A pantropical
tramp and common house-infesting ant that may nest in soil or in buildings. They are
omnivorous and may feed on various household foods. They have been reported to gnaw
holes in fabrics, rubber goods, remove rubber insulation from electric or telephone
wires, and damage polyethylene cable. Introduced, probably originating from Africa or
the Oriental region.
Atta Destructor Jerdon, 1851. Madras Jour. Lit. and Sci. 17:105. 3.
Myrmica vastator Smith, 1857. Linn. Soe. London, Jour., Zool. 2:71.
Myrmica basalis Smith, 1858. Cat. Hym. Brit. Mus. 6:125. &.

Taxonomy: Bingham, 1903. Fauna British India 2:209 (each caste). —Emery, 1908. Deut. Ent.
Ztschr. pp. 665-666, 671. —Phillips, 1934. (Hawaii Univ.) Expt. Sta., Pineapple Prod. Coop.
Assoc., Bul. 15:2. —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:64.

Biology: Wheeler, 1910. Ants, pp. 10, 153, 221. —Wheeler, 1914. Amer. Jour. Trop. Dis. and
Prev. Med. 2:160-168 (economics). —Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:833.
—Kalshoven, 1937. Ent. Meded. van Nederland. Indie 3:65-71. —Marlatt, 1928. U. S. Dept.
Agr., Farmers’ Bul. 740:9. —Kempf, 1960. Studia Ent. 3:506-507 (economics). —Smith,
1965. U. S. Dept. Agr., Tech. Bul. 1326:37-38 (economic importance).

floricola (Jerdon). Fla., Ala; throughout tropical regions of world. Ecology: An arboreal
species, nesting in twigs and branches or under bark of trees or other plants. Commonly
infests houses and feeds on household foods. Introduced, probably originating from
tropical Asia.
Atta floricola Jerdon, 1851. Madras Jour. Lit. and Sci. 17:107. 8.
Monomorium speculare Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. K1., Sitzber. 53:509.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:88 (female). —Smith, 1936. Puerto
Rico Univ., Jour. Agr. 20:831, 834. —Wheeler and Wheeler, 1955. Amer. Midland Nat.
54:121 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:64-65.

Biology: Wheeler, 1910. Ants, pp. 153, 426. —Smith, 1930. Fla. Ent. 14:3. —Wheeler, 1932. N.
Y. Ent. Soc., Jour. 40:9. —Plank and Smith, 1940. Puerto Rico Univ., Jour. Agr. 24:49-75
(association with pineapple mealybug, Pseudococcus brevipes (Ckll.)). —Brown, 1964. Ent.
News 75:15. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:36-37 (economic importance).

minimum (Buckley). Que., Ont. s. to Fla., w. to Mont., Colo., Calif.; Mexico. Rare or absent in

the Pacific Northwest. Ecology: Nests in exposed soil, under cover of objects, or in
rotting or faulty wood. Sometimes invades houses from outdoors or nests in woodwork.
Parasite: Monomorium pergandei (Emery). Little black ant.

Myrmica (Monomarium (!)) minima Buckley, 1867. Ent. Soc. Phila., Proc. 6:338. 3, °.

Myrmica (Monomarium (!)) atra Buckley, 1867. Ent. Soc. Phila., Proc. 6:342. % (?).

Monomorium minutum ergatogyna Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:269. &,
apterous ergatoid °.

Monomorium minimum emersoni Gregg, 1945. Psyche 52:66. 8, °.

Monomorium metoecus Brown and Wilson, 1957. Ent. News 68: 239-244. Ergatogyne.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:274-275 (each caste). —Wheeler,
1905. South. Calif. Acad. Sci., Bul. 40:60. —Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.
24:423. —Buren, 1944. Iowa State Col., Jour. Sci. 18:289. —Wheeler and Wheeler, 1955.
Amer. Midland Nat. 54:122 (larva). —Wilson and Brown, 1958. Ent. News 69:33-38 (worker
of metoecus; also biology). —Ettershank, 1966. Austral. Jour. Zool. 14:90 (syn. of

metoecus).

Superfamily FORMICOIDEA 1383

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:274, 377. —Marlatt, 1922. U. S. Dept.
Agr., Farmers’ Bul. 740:4, 10. —Eckert and Mallis, 1937. Calif. Agr. Expt. Sta., Cir. 342:23.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31:271, 272, 274, 279-280. —Metcalf and Flint, 1939.
Destructive and Useful Insects, p. 770. —Gregg, 1944. Ent. Soc. Amer., Ann. 37:454,456,
466. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 133-135. —Gregg, 1963. Ants of
Colo., pp. 366-368. —Burns, 1964. Ent. Soc. Amer., Ann. 57:138 (association with tuliptree
scale). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:33-34 (economic importance).
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 89-90.

pergandei (Emery). D. C. Ecology: Supposedly workerless and parasitic. The host colony
contained only winged males and winged females. Host: Monomoriwm minimum
(Buckley).
Epoecus pergandei Emery, 1892. Soc. Ent. de France, Bul. 61: CCLXXVI. 2, d.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:272-274. —Wheeler, 1910. Ants, p.
498. —Smith, 1947. Amer. Midland Nat. 37:569 (female). —Ettershank, 1966. Austral. Jour.
Zool. 14:82, 91.

pharaonis (Linnaeus). Fla., throughout U. S. and Canada in larger cities; cosmopolitan.
Ecology: Not uniformly distributed; adapted to field conditions only in Florida but
probably found in every town or city of commercial importance especially in hotels,
large apartment buildings, groceries, or other places where food is commercially
handled. Colonies are populous and are commonly found in the most inaccessible places
in buildings. They breed continuously through the year and take 38 to 42 days to
develop. They may feed on various household foods, damage silk, rayon and rubber
goods and are also known to damage insect collections. This is the most persistent and
difficult of all our house-infesting ants to control or eradicate. Introduced, probably
originating from Africa or tropical Asia. Pharaoh ant.

Formica Pharaonis Linnaeus, 1758. Syst. Nat., Ed. 10:580.

Taxonomy: Emery, 1908. Deut. Ent. Ztschr., pp. 664-665, 667, 684 (each caste).
—Donisthorpe, 1927. British Ants, p. 104. —Buren, 1944. Iowa State Col., Jour. Sci. 18:289.
—Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:121 (larva). —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62: 16 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog.
14:68-60.

Biology: Bellevoye, 1889. U. S. Dept. Agr., Insect Life 2:230-233 (in 2 years, 1,360,000
workers, 1,900 females, and 560 males taken on same premises). —Herrick, 1914. Insects
Injurious to the Household and Annoying to Man, pp. 174-176. —Essig, 1926. Insects of
West. N. Amer., p. 857. —Metcalf and Flint, 1939. Destructive and Useful Insects, p. 770.
—Smith, 1934. Kans. Acad. Sci., Trans. 37:140-142. —Armand, 1942. Pests 10:18-19
(thermal preference as guide to control work). —Peacock and Baxter, 1949. Ent. Monthly
Mag. 85:256. —Peacock, 1950. Ent. Monthly Mag. 86:129. —Peacock and Baxter, 1950. Ent.
Monthly Mag. 86:171-178 (life history). —Peacock, 1950. Ent. Monthly Mag. 86:294.
—Peacock, et al., 1950. Seot. Dept. Agr., Misc. Pub. 17:1-50 (biology and control).
—Peacock, 1951. Ent. Monthly Mag. 87:185-191. —Sudd, 1953. Advancement of Sci. (Brit.)
10:17-18 (colony foundation). —Peacock, et al., 1954. Ent. Monthly Mag. 90:154-158 (male
production by parthenogenesis). —Peacock, et al., 1955. Ent. Monthly Mag. 91:125, 130.
—Peacock, et al., 1955. Ent. Monthly Mag. 91:37-42 (viability in regard to temperature
and moisture). —Brown, 1958. Acta Hym. 1:36. —Wheeler and Wheeler, 1963. Ants of N.
Dak., pp. 135-136. —Brown, 1964. Ent. News 75:14-15. —Eichler, 1963. Deut. Ent. Ztschr.
10:207-215. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:34-36 (economic importance).
—Wisniewski, Myjak, and Przyborowski, 1971. Polskie Pismo Ent. 61:459-474 (biology, in
Poland). —Petersen and Buschinger, 1971. Anz. f. Schadlingsk. u. Pflanzensch. 44:121-127
(behavior). —Petersen and Buschinger, 1971. Ztschr. f. Angew. Ent. 68:168-175 (behavior
of sexuals).

Morphology: Tanquary, 1913. Ill. State Lab. Nat. Hist., Bul. 9:443-453 (physiology). — Hall
and Smith, 1951. Ent. Monthly Mag. 87:217: —Hall and Smith, 1952. Ent. Monthly Mag.
88:97-102 (thoracic structure). —Hall and Smith, 1953. Evolution 7:127. —Hall and Smith,
1954. Ent. Monthly Mag. 90:176-182 (somatic mosaics). —Smith and Peacock, 1957. Roy.
Soc. Edinb., Proc., Section B, 66:235-261 (cytology).

1384 Hymenoptera in America North of Mexico

viridum peninsulatum Gregg. Ga., Fla. w. to Colo., Ariz., s. Calif. Ecology: Most commonly
found in’soil under objects.
Monomorium peninsulatum Gregg, 1945. Psyche 52:62. %, 9.

Biology: Gregg, 1963. Ants of Colo., pp. 368-370.
viridum viridum Brown. N. Y. (Coram, Selden, L. I.), N. J. (Lakehurst). Ecology: The types
from N. J. were taken from nests with 5 to 8 inch craters in yellow and white sand
among scanty, low weeds of a roadside strip.
Monomorium viridum Brown, 1943. Ent. News 54:243. 3, 9°.

Genus XENOMYRMEX Forel

Xenomyrmesx Forel, 1885. Soc. Vaud. des Sci.Nat., Bul. 20:369.
Type-species: Xenomyrmezx stollii Forel. Monotypic.

Myrmecinella Wheeler, 1922. Amer. Mus. Novitates 46:1.
Type-species: Myrmecinella panamana Wheeler. Monotypic.

A small New World genus of three species. The ants are arboreal, nesting in small colonies in
plant cavities such as twigs, galls, and thorns.

Revision: Wheeler, 1931. Rev. de Ent. 1:129-139. —Creighton, 1957. Amer. Mus. Novitates
1843:1-14.

Taxonomy: Creighton, 1953. Amer. Mus. Novitates 1634:2. —Wheeler and Wheeler, 1960.
Ent. Soc. Wash., Proc. 62:19-20 (larva). —Ettershank, 1966. Austral. Jour. Zool. 14:150-152.

floridanus floridanus Emery. Fla.; Bahamas, Cuba. Another subspecies is found in Mexico.
Xenomyrmex stolli floridanus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:275. 8, d.
Xenomyrmex stolli floridanus var. lucayanus Wheeler, 1905. Amer. Mus. Nat. Hist., Bul.
21:87. &.
Xenomyrmesx stolli rufescens Wheeler, 1931. Rev. de Ent. 1:133,137. °.
Xenomyrmex stolli cubanus Wheeler, 1931. Rev. de Ent. 1:134-135. 8.

Biology: Wheeler, 1901. Amer. Nat. 35:538. —Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:6.
Morphology: Holldobler, 1971. Jour. Ins. Physiol. 17:1497-1499 (sex pheromone).

Genus SOLENOPSIS Westwood

Solenopsis Westwood, 1841. Ann. and Mag. Nat. Hist. 6:86.
Type-species: Solenopsis mandibularis Westwood. Monotypic.

Diplorhoptrum Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:449.
Type-species: Formica fugax Latreille. Monotypic.

Solenopsis subg. Synsolenopsis Forel, 1918. Soc. Vaud. des Sci. Nat., Bul. 52:155.
Type-species: Solenopsis bruchiella Emery. Monotypic.

Solenopsis subg. Diagyne Santschi, 1923. Rev. Suisse de Zool. 30:267.
Type-species: Solenopsis succinea Emery. Monotypic.

Labauchena Santschi, 1930. Soc. Ent. Argent., Rev. 13:81.
Type-species: Labauchena daguerrei Santschi. Monotypic.

Solenopsis subg. Euophthalma Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66: 43.
Type-species: Myrmica globularia Smith. Orig. desig.

Solenopsis subg. Oedaleocerus Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66:43.
Type-species: Solenopsis argulata Emery. Orig. desig.

Bisolenopsis Kusnezov, 1953. Acta. Zool. Lilloana [Tucuman] 13:1.
Type-species: Bisolenopsis sea Kusnezov. Monotypic.

Paranamyrma Kusnezov, 1954. Mus. Entre Rios, Mem. 30:9.
Type-species: Paranamyrma solenopsidis Kusnezov. Monotypic.

Lilidris Kusnezov, 1957. Zool. Anz. 158:268, 274. Uncertain syn.
Type-species: Lilidris metatarsalis Kusnezov. Monotypic.

Solenopsis subg. Granisolenopsis Kusnezov, 1957. Zool. Anz. 158:270, 277.
Type-species: Solenopsis (Granisolenopsis) granivora Kusnezov. Monotypic.

Superfamily FORMICOIDEA — 1385

Previously, this genus was divided into several subgenera, the subgenus Solenopsis included
the larger, polymorphic species aurea, geminata, invicta, richteri, and xyloni, the subgenus
Euophthalma included globularia littoralis and huachucana, and the subgenus Diplorhoptrum
included the minute “thief ants”, the remaining species listed here. Those previously placed in
the subgenus Solenopsis are commonly known as “fire ants”, the colonies are populous, nests are
usually mounded in exposed situations, and workers are agressive and possess a painful sting.
They are of the most economically important of ants. Most species in the other subgenera are
minute and cryptic, often lestobiotic in the nests of other ants.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:277-279 (Diplorhoptrum). —Wheeler,
1915. Amer. Mus. Nat. Hist., Bul. 34:397. —Creighton, 1930. Amer. Acad. Arts and Sci.,
Proc. 66:39-139.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:131-136 (larvae). —Snelling,
1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3, 10 pp. (keys to all castes of subg.
Solenopsis). —Ettershank, 1966. Austral. Jour. Zool. 14:134-144 (generic syn. and list of
world species). —Buren, 1972. Ga. Ent. Soe., Jour. 7:1-27 (revisionary studies of the
imported fire ants).

Morphology: Blum, et al., 1958. Science 128:306-307 (chemical, insecticidal and antibiotic
properties of fire ant venom). —MacConnel, Blum and Fales, 1971. Tetrahedron
26:1129-1139 (chemistry of fire ant venom). —San Martin, 1971. In Bucheil and Buckley,
Venomous Animals and Their Venoms v. 3, pp. 95-101 (venomous ants of Solenopsis).

aurea Wheeler. Tex., N. Mex., Ariz., s. Calif.; Mexico. Ecology: Nests are in fully exposed
situations, in dry, coarse, gravelly soil, and are without a mound.
Solenopsis geminata var. aurea Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:336. 3, °,
3

Solenopsis aurea amblychila Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:394. 8, 2, d.
Taxonomy: Snelling, 1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3: 7.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:425-426.

carolinensis Forel. Mass. s. to N. C., Tenn., Ohio. Ecology: Probably lestobiotic.
Solenopsis tenana race carolinensis Forel, 1901. Soc. Ent. de Belg., Ann. 45:345. 9, °, d.

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:92.

geminata (Fabricius). S. C. s. to Fla., w. to Tex. and s. to Peru; West Indies; tropical Asia and

Pacific Islands. Ecology: In the U. S., it is most common on or near the coast with its
incidence decreasing inland except in Florida where it is distributed over most of the
state. Nests are in the ground in open areas in dry to moist soil of various composition
surmounted by irregular piles of dirt; they may also be under cover of objects or in
rotting logs. Economic importance is similar to that of xyloni McCook, but its
importance has subsided since the introduction of the imported fire ants. A common
tramp species, native to the New World and spread by commerce to other tropical
regions of the world. Fire ant.

Atta geminata Fabricius, 1804. Systema Piezatorum, p. 423. °.

Atta Rufa Jerdon, 1851. Madras Jour. Lit. and Sci. 17:106. 8.

Solenopsis mandibularis Westwood, 1841. Ann. and Mag. Nat. Hist. 6:87. 9.

Myrmica virulens Smith, 1858. Cat. Hym. Brit. Mus. 6:132. 8.

Atta clypeata Smith, 1858. Cat. Hym. Brit. Mus. 6:169. 2, d.

Solenopsis cephalotes Smith, 1858. Linn. Soc. London, Jour., Zool. 3:149. 9.

Crematogaster laboriosus Smith, 1860. Linn. Soc. London, Jour., Zool. 4:109. ?.

Diplorhoptrum drewseni Mayr, 1861. Eur. Formicid., p. 71. 8.

Myrmica glaber Smith, 1862. Ent. Soc. London, Trans. (3) 1: 34. 9.

Myrmica polita Smith, 1862. Ent. Soc. London, Trans. (8) 1: 34. 8.

Myrmica (Monomarium (!)) saxicola Buckley, 1867. Ent. Soc. Phila, Proc. 6:341. %. Syn.
uncertain.

Atta Lincecumii Buckley, 1867. Ent. Soc. Phila., Proc. 6:344. %, 2{. Syn. uncertain.

Atta brazoensis Buckley, 1867. Ent. Soc. Phila., Proc. 6:345. 3, 2{. Syn. uncertain.

Atta coloradensis Buckley, 1867. Ent. Soc. Phila., Proc. 6:346. %, 2{. Syn. uncertain.

1386 Hymenoptera in America North of Mexico

Solenopsis geminata var. diabola Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:424. %, 2,
é.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:132-133 (larva). —Snelling,
1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3:7-9. —Wilson and Taylor, 1967.
Pacific Ins. Monog. 14:58-59.

Biology: Mann, 1920. Amer. Mus. Nat. Hist., Bul. 42:427. —Marlatt, 1928. U. S. Dept. Agr.,
Farmers’ Bul. 740:5. —Neig, 1930. Bombay Nat. Hist. Soc., Jour. 34:185. —Clark, 1931.
Tex. Agr. Expt. Sta. Bul. 435:1-12. —Cole, 1934. Ent. Soc. Amer., Ann. 27:395. —Phillips,
1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc., Bul. 15:12-17. —Smith, 1936.
Puerto Rico Univ., Jour. Agr. 20:838-839. —Plank and Smith, 1940. Puerto Rico Univ.,
Jour. Agr. 24:49-76. —Travis, 1941. Fla. Ent. 24:15-22. —Griffiths, 1942. Science 96:271-272.
—Lindquist, 1942. Jour. Econ. Ent. 35:850-851. —Kempf, 1961. Studia Ent. 4:507. —Smith,
1965. U. S. Dept. Agr., Tech. Bul. 1326:40-41 (economic importance). —Banks, Plumley, and
Hicks, 1973. Ent. Soc. Amer., Ann. 66:234-235 (polygyny in a colony). —Bass and Hays,
1976. Ga. Ent. Soc., Jour. 11: 34-36 (in S. C.).

globularia littoralis Creighton. N. C. s. to Fla., w. to La.; Mexico. Ecology: Commonly found
on open beaches; nests are constructed in or under rotten logs. S. globularia globularia
(Smith) occurs in Central and S. Amer.
Solenopsis (Euophthalma) globularia littoralis Creighton, 1930. Amer. Acad. Arts and
Sci., Proc. 66:110, 113. ¥, 2, do.
Solenopsis globularia mobilensis Smith, 1931. Ent. News 42:20. §. Nomen nudum.

Taxonomy: Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:22 (larva).

Biology: Smith, 1931. Ent. News 42:20. —Smith, 1933. Fla. Ent. 17:23.

huachucana Wheeler. Ariz. (Miller Canyon, Huachuca Mtns.).
Solenopsis huachucana Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:393. 8, @.

Taxonomy: Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66:118-120.

invicta Buren. N. C., S. C., Ga., Fla., Tenn., Ala., Miss., Ark., La., Tex.; Brazil. Ecology: The
most widespread of the two imported fire ants and the most economically important
being an annoying and sometimes destructful pest. The mound nests are unsightly and
sometimes interfere with agricultural operations; workers are agressive and can inflict a
painful sting; workers steal seeds from seedbeds and feed on germinating seeds of corn,
may gnaw holes in various fabrics, foster honeydew excreting insects, injure or kill
young rabbits, pigs, and other mammals, quail and other birds, gnaw into roots, stems,
buds, and fruits of various agricultural crops, and may girdle young plants. Introduced,
probably originating from Mato Grosso, Brazil; the earliest U.S. record is 1945 from
Daphne, Ala. Red imported fire ant. Because of the recent distinction of two introduced
species in the U. S., it is difficult to determine whether much of the literature pertains
to this species or S. richteri. Only selected references are given below; for a
bibliography through 1971 see U. S. Dept. Agr., Coop. Econ. Ins. Rpt., 1971, Sept., 21
(36): 639-652. Much of the literature is under the name S. saevissima richteri Forel.

Solenopsis invica (!) Buren, 1972. Ga. Ent. Soc., Jour. 7:9-15. 3, 2, d.

Taxonomy: Wilson, 1951. Evolution 5:68-79. —Wilson, 1952. Inst. Oswaldo Cruz, Mem.
50:49-68. —Wilson, 1953. Evolution 7:262-263. —Brown, 1957. Quart. Rev. Biol. 32:258-261.
—Wilson and Brown, 1958. Evolution 12:211-218 (morphological changes in introduced
population). —Snelling, 1963. Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3:10. —Buren,
1972. Ga. Ent. Soc., Jour. 7:9-15. —Buren, et. al., 1974. N. Y. Ent. Soc., Jour. 82: 113-124
(zoogeography of the imported fire ants).

Biology: Wilson, 1958. Sci. Amer. 198:36-41. —Bellinger, Dyer, King and Pratt, 1965. Ga.
Acad. Sci., Bul. 23:122 (review of fire ant problem). —Fincher and Lund, 1967. Ga. Ent.
Soc., Jour. 2:91-94 (biology and life cycle in Ga.). —Rhoades and Davis, 1967. Jour. Econ.
Ent. 60:544-558 (effects of meteorological factors on biology and control). —Anon. 1971
Sept., U. S. Dept. Agr., Coop. Econ. Ins. Rpt., 21 (36): 639-652 (bibliography through 1971).
—Collins and Markin, 1971. Ent. Soc. Amer., Ann. 64:1376-1380 (inquilines and other
arthropods).—Markin and Dillier, 1971. Ent. Soc. Amer., Ann. 64:562-565 (seasonal life

Superfamily FORMICOIDEA 1387

cycle in Miss.). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:41-43 (economic
importance). —Wilson and Oliver, 1969. Jour. Econ. Ent. 62:1268-1271 (food habits).
—Glancey, et al., 1970. Nature 226:863-864 (pheromone may induce brood tending).
—Wilson, Dillier, and Markin, 1971. Ent. Soc. Amer., Ann. 64:660-665 (foraging territories).
—Markin, et al., 1971. Ga. Ent. Soc., Jour. 6:145-156 (nuptial flight and flight ranges).
—Collins and Markin, 1971. Ent. Soc. Amer., Ann. 64:1376-1380 (inquilines and other
arthropods in nests). —Markin, Collins, and Dillier, 1972. Ent. Soc. Amer., Ann.
65:1053-1058 (colony founding by queens). —Harris and Burns, 1972. Environ. Ent.
1:362-365 (predation on lone star tick). —Bhatkar, et al., 1972. Environ. Ent. 1:274-279
(confrontation behavior between Lasius neoniger and the imported fire ant). —Stringer, et
al., 1972. Jour. Econ. Ent. 65:872-873 (air separation of different castes). —Glancey,
Stringer, and Bishop, 1973. Ga. Ent. Soc., Jour. 8:217-220 (trophic egg production).
—Glancey, et al., 1973. Ga. Ent Soc., Jour. 8:237-238 (multiple fertile queens in colonies).
—O’Neal and Markin, 1973. Ga. Ent. Soc., Jour. 8:294-303 (brood nutrition and parental
relationships). —Hubbard, 1974. Ga. Ent. Soc., Jour. 9: 127-132 (influence of nest material
and colony odor on digging). — Allen, et. al., 1974. Ent. Soc. Amer., Ann. 67: 43-46
(distribution and habits in Brazil). —Morrill, 1974. Environ. Ent. 3: 265-271 (production and
flight). —Morrill, 1975. Ga. Ent. Soc., Jour. 10: 162-164 (reduction of populations by tillage).
—Horton, et al., 1975. Ga. Ent. Soc., Jour. 10: 207-213 (food carrying ability and
recruitment time). —Lofgren, Banks, and Glancey, 1975. Ann. Rev. Ent. 20: 1-30 (biology
and control of imported fire ants). —Bass and Hays, 1976. Ga. Ent. Soc., Jour. 11: 34-36 (in
S. C.). —Adams, et al., 1976. Ga. Ent. Soc., Jour. 11: 165-169 (economic importance, impact
on soybean harvest).

Morphology: Blum, et al., 1958. Science 128:306-307 (chemical, insecticidal, and antibiotic
properties of venom). —Callahan, Blum, and Walker, 1959. Ent. Soc. Amer., Ann.
52:573-590 (histology of poison glands and sting). —Adrouny, Derbes, and Jung, 1959.
Science 130:479 (hemolytic component of fire ant venom). —Wilson, Durlach and Roth,
1959. Psyche 65:108-114 (chemical releases of necrophoric behavior). —Wilson, 1959.
Science 129:643-644 (nature of odor trail). —Wilson, 1962. Animal Behavior 19:134-147
(chemical communication among workers-mass foraging). —Blum and Callahan, 1960. XI
Int. Kong. Ent. Wien, Verh. B. 111:290-293 (chemical and biological properties of venom).
—Wilson, 1962. Animal Behavior 10:148-158 (chemical communication among workers).
—Wilson, 1962. Animal Behavior 10:159-164 (induction of social responses). —Walker and
Clower, 1961. Ent. Soc. Amer., Ann. 54:92-99 (alimentary canal of queen). —Caro, Derbes,
and Jung, 1957. Amer. Med. Assoc., Arch. Dermat. 75:475-488 (skin responses to sting).
—Thompson and Blum, 1967. Ent. Soc. Amer., Ann. 60:632-642 (spermatozoa). —Vinson,
1970. Ent. Soc. Amer., Ann. 63:930-935 (gustalory response to various electrolytes).
—MacConnell, Blum, and Fales, 1970. Science 168:840-841 (alkaloid from fire ant venom).
—Smith and Smith, 1971. Arch. Dermat. 103:438-441 (multiple fire ant stings - a
complication of alcoholism). —MacConnell, Blum, and Fales, 1971. Tetrahedron
26:1129-1139 (chemistry of fire ant venom). —Glancey, et al., 1976. Ga. Ent. Soc., Jour. 11:
83-88 (testes degeneration). —Robeau and Vinson, 1976. Ga. Ent. Soc., Jour. 11: 198-203
(effects of juvenile hormone analogues on caste differentiation).

krockowi Wheeler. N. Mex.; Mexico.
Solenopsis krockowi Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:428. 3, °.
molesta (Say). Ont. s. to Fla, w. to Wash., Calif. Ecology: Lestobiotic, usually nests in or near
nests of other ants from which they rob food and brood. A house-infesting ant and of
great annoyance because of their small size; sometimes nests in woodwork and masonry
of houses. Thief ant. The form occurring west of the Rockies is sometimes referred to
as the subspecies validiuscula.
Myrmica molesta Say, 1836. Boston Jour. Nat. Hist. 1:293. ?.
Myrmica minuta Say, 1836. Boston Jour. Nat. Hist. 1:294. 9.
Myrmica (Tetmamorium (!)) exigua Buckley, 1867. Ent. Soc. Phila., Proc. 6:342. 3, “2” =
é.
Solenopsis debilis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:461. %, 2, d.
Solenopsis molesta var. validiuscula Emery, 1895. Zool. Jahrb., Abt. f. System. 8:278. %.

1388 Hymenoptera in America North of Mexico

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:430. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bul. 22:584 (each caste). —Wheeler and Wheeler, 1955. Amer.
Midland Nat. 54:134-135 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:377-378. —McColloch and Hayes,
1916. Jour. Econ. Ent. 9:23-38. —Hayes, 1920. Kans. Agr. Expt. Sta., Tech. Bul. 7:1-54.
—Kckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342:21-22. —Metcealf and Flint, 1939.
Destructive and Useful Insects, p. 770. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:72.
—Macnamara, 1945. Canad. Ent. 77:40. —Grundmann and Peterson, 1953. Kans. Ent. Soc.,
Jour. 26:59. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 136-138. —Gregg, 1963.
Ants of Colo., pp. 372-375. —Ayre, 1963. Canad. Ent. 95:712-715 (feeding habits). —Smith,
1965. U. S. Dept. Agr., Tech. Bul. 1326:43-45 (economic importance). —Wheeler and
Wheeler, 1973. Ants of Deep Canyon, pp. 91-92.

pergandei Forel. Va. s. to Fla., w. to La. Ecology: Nests have been found in soil in rotting
stumps, and next to nests of other ants.
Solenopsis pergandei Forel, 1901. Soc. Ent. de Belg., Ann. 45:343. 8, 2, d.

Taxonomy: Smith, 1931. Ent. News 42:20. —Wheeler and Wheeler, 1960. Ent. Soc. Wash.,
Proc. 62:21-22 (larva).

Biology: Smith, 1931. Ent. News 42:20. —Smith, 1944. Fla. Ent. 27:15.
picta Emery. S. C. s. to Fla., w. to Tex. Ecology: Nests have been found in hollow twigs.
Solenopsis tenuis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:262. %. Preoce. by Mayr,
1877.
Solenopsis picta Emery, 1895. Zool. Jahrb., Abt. f. System. 8:278. &.
Solenopsis picta var. moerens Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:393. 8.

Taxonomy: Smith, 1942. Ent. Soc. Wash., Proc. 44:211. —Wheeler and Wheeler, 1960. Ent.
Soc. Wash., Proc. 62:22 (larva).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:10.
pilosula Wheeler. Tex. (Alice).
Solenopsis pilosula Wheeler, 1908. Amér. Mus. Nat. Hist., Bul. 24:426. 3, 6.
richteri Forel. Miss., Ala.; S. Brazil, Uruguay, Argentina. Ecology: A mound building ant with
similar habits and importance of invicta Buren. May have been more widespread in U.
S. until the introduction of invicta which may have replaced richteri in many localities.
Introduced, probably originating from southern S. Amer.; first recorded from Mobile,
Ala. in 1930 but may have been present there for 10 to 12 years. Black imported fire
ant. Until 1972, the imported fire ant had been known under one name, S. saevissima
richteri Forel; consequently, it is difficult to determine to which species, either invicta
or richteri, much of the earlier literature pertains. For a bibliography of the “imported
fire ant” through 1971, see U. S. Dept. Agr., Coop. Econ. Ins. Rpt., 1971, Sept., 21 (36):
639-652, as well as the list of references given under invicta.
Solenopsis Pylades var. Richteri Forel, 1909. Deut. Ent. Ztschr., p. 267. %, 2.

Taxonomy: Creighton, 1930. Amer. Acad. Arts and Sci., Proc. 66:87 (first U. S. record).
—Wilson, 1951. Evolution 5:68-79. —Wilson, 1952. Inst. Oswaldo Cruz, Mem. 50:49-68.
—Wilson, 1953. Evolution 7:262-263 (origin of variation). —Brown, 1957. Quart. Rev. Biol.
32:258-261. —Wilson and Brown, 1958. Evolution 12:211-218 (morphological changes in
introduced population). —Buren, 1972. Ga. Ent. Soc., Jour. 7:4-8 (two species of imported
fire ants).

Biology: Green, 1952. Jour. Econ. Ent. 45:593-597 (biology and control in Miss.). —Green,
1967. Miss. State Univ., Agr. Expt. Sta., Bul. 737, 23 pp.

salina Wheeler. W. Tex., Colo. w. to Calif.; Mexico. Ecology: Nests under rocks and wood;
lestobiotic.
Solenopsis salina Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:427. 9.

Biology: Gregg, 1963. Ants of Colo., pp. 376-378.

tennesseensis Smith. Fla., Tenn., Miss., Tex., Kans., Calif. (nr. Azusa).
Solenopsis (Diplorhoptrum) longiceps Smith, 1942. Ent. Soc. Wash., Proc. 44:210. 8.
Preoce. by Forel, 1907.

Superfamily FORMICOIDEA 1389

Solenopsis (Diplorhoptrum) tennesseensis Smith, 1951. In Muesebeck, et al., U. S. Dept.
Agr., Agr. Monog. 2:814. N. name.
texana catalinae Wheeler. Calif. (Catalina Is.).
Solenopsis texana catalinae Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:269. 3, °.

Taxonomy: Wheeler, 1905. South. Calif. Acad. Sci., Bul. 4:60.
texana texana Emery. Ont. s. to Fla., w. to Okla., Tex. Ecology: Nests have been found in logs,
stumps, and under bark.
Solenopsis pollux var. texana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:278. %.
Solenopsis rosella Kennedy, 1938. Canad. Ent. 70:232. %, 2, 3.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:430-431. —Wheeler and Wheeler,
1955. Amer. Midland Nat. 54:136 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14:70.
truncorum Forel. N. C. s. to Fla., w. to Colo., Ariz., Calif. Ecology: Nests have been found
under rocks and other objects; lestobiotic.
Solenopsis texana race truncorum Forel, 1901. Soc. Ent. de Belg., Ann. 45:346. 3, °.
Solenopsis molesta var. castanea Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:430. 8.

Biology: Gregg, 1963. Ants of Colo., pp. 378-380.

xyloni McCook. N. C. s. to Fla., w. to Colo., Calif.; Mexico. Ecology: Nests are in ground in

exposed soil or under cover of stones or other objects, sometimes in wood; they are
surmounted by irregular and variable-shaped mounds of loose soil. A serious pest in
some parts of its range: builds ugly mounds on lawns, inflicts painful stings, steals seeds
from seedbeds, kills young poultry and other birds, girdles nursery stock, gnaws into
buds, tubers, and fruits of various plants, bites holes in fabrics such as silk, nylon, and
linen, removes rubber insulation from telephone wires, and feeds on household foods.
Southern fire ant.

Solenopsis xyloni McCook, 1879. In Comstock, Rpt. Cotton Ins., p. 188. %, @.

Myrmica (Atta) sabeana Buckley, 1867. Ent. Soc. Phila., Proc. 6:343. %. Syn. questionable.

Solenopsis geminata maniosa Wheeler, 1915. Amer. Mus. Nat Hist., Bul. 34:396. %, 2, 2,
é.

Taxonomy: Wheeler and Wheeler, 1955. Amer. Midland Nat. 54:133 (larva). —Snelling, 1963.
Calif. Dept. Agr., Bur. Ent., Occas. Papers No. 3:9.

Biology: Severin, 1923. Jour. Econ. Ent. 16:96-97. —Smith, 1936. Jour. Econ. Ent. 29:120-122.
—Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342:10-21. —Mallis, 1938. Pan-Pacific
Ent. 14:87-91. —Metcalf and Flint, 1939. Destructive and Useful Insects, p. 771.
—Eagleson, 1940. Jour. Econ. Ent. 33:700. —Wray, 1962. Jour. Econ. Ent. 55:145 (apparent
introduction into N. C.). —Gregg, 1963. Ants of Colo., pp. 370-371, 373. —Smith, 1965. U. S.
Dept. Agr., Tech. Bul. 1326:38-40 (economic importance). —Wheeler and Wheeler, 1973.
Ants of Deep Canyon, pp. 92-94.

Morphology: Blum, Roberts, and Novak, 1961. Psyche 68:73-74 (chemical and biological
characters of venom).

Genus OLIGOMYRMEX Mayr

Oligomyrmex Mayr, 1867. Ent. Tidskr. 10:110.
Type-species: Oligomyrmex concinnus Mayr. Monotypic.
Aeromyrma Forel, 1891. Soc. Ent. de Belg., Ann. 35:307.
Type-species: Aeromyrma nosindambo Forel. Monotypic.
Pheidologeton subg. Aneleus Emery, 1900. Termes. Fuzetek 23:327.
Type-species: Solenopsis similis Mayr. Desig. by Wheeler, 1911.
Erebomyrma Wheeler, 1903. Biol. Bul. 4:138.
Type-species: Erebomyrma longii Wheeler. Monotypic.
Pheidologeton subg. Lecanomyrma Forel, 1913. Soc. Vaud. des Sci. Nat., Bul. 49:56.
Type-species: Pheidologeton (Lecanomyrma) butteli Forel. Monotypic.
Oligomyrmex subg. Octella Forel, 1915. Arkiv for Zool. 9:69.
Type-species: Oligomyrmex (Octella) pachycerus Forel. Monotypic.

1390 Hymenoptera in America North of Mexico

Spelaeomyrmex Wheeler, 1922. Amer. Mus. Novitates 45:9.
Type-species: Spelaeomyrmesx urichi Wheeler. Monotypic.

Oligomyrmex subg. Hendecatella Wheeler, 1927. Lab. Zool. Gen. e Agr. Portici, Bol. 20:93.
Type-species: Oligomyrmex (Hendecatella) capreolus Wheeler. Monotypic.
Solenopsis subg. Solenops Karawajew, 1930. Zool. Anz. 92:207. Preoce. by Dufour, 1820.

Type-species: Solenopsis (Solenops) weyeri Karawajew. Monotypic.
Sporocleptes Arnold, 1948. Natl. Mus. South. Rhodesia, Occas. Papers 2 (14): 219.
Type-species: Sporocleptes nicotiana Arnold. Monotypic.
Solenopsis subg. Crateropsis Patrizi, 1948. Ist. Ent. Univ. Bologna, Bol. 17:174.
Type-species: Solenopsis (Crateropsis) elmenteitae Patrizi. Monotypic.
Nimbamyrma Bernard, 1953. Mem. Inst. Franc. Afr. Noire 19, fase. 1, p. 240. Questionable
syn.
Type-species: Nimbamyrma villiersi Bernard. Monotypic.

A large genus in tropical Africa and Asia, with fewer species in the neotropics and only one
species reaching the United States. Colonies are small and collections are usually made from
under bark of logs, in rotten wood, or leaf litter. Most species may be lestobiotic in the nests of
other ants and termites.

Taxonomy: Wilson, 1962. Psyche 69:62-72. —Ettershank, 1966. Austral. Jour. Zool. 14:119-124
(generic syn. and list of world species). —Wheeler and Wheeler, 1953. Psyche 60:141
(larvae).

longii (Wheeler). Okla. (Ft. Sill), Tex. (Denton). Ecology: May be lestobiotic.
Erebomyrma longii Wheeler, 1903. Biol. Bul. 4:140. 3, 2, d.

Taxonomy: Mann, 1926. Psyche 33:104. —Wheeler, 1936. Amer. Acad. Arts and Sci., Proc.
71:197.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:423. —Wheeler, 1910. Ants, pp. 113,
140, 152, 158-159, 427-428.

TRIBE LEPTOTHORACINI

Genus MACROMISCHA Roger

Macromischa Roger, 1863. Berlin. Ent. Ztschr. 7:184.
Type-species: Macromischa purpurata Roger. Desig. by Wheeler, 1911.
Macromischa subg. Croesomyrmex Mann, 1920. Amer. Mus. Nat. Hist., Bul. 42:408.
Type-species: Macromischa (Croesomyrmex) wheeleri Mann. Orig. desig.
Macromischa subg. Antillaemyrmex Mann, 1920. Amer. Mus. Nat. Hist., Bul. 42:408.
Type-species: Macromischa (Antillaemyrmex) terricola Mann. Orig. desig.

A rather large genus in the neotropics with only three species reaching the United States.

Revision: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:138-142. —Mann, 1920. Amer. Mus.
Nat. Hist., Bul. 42:407-424. —Wheeler, 1931. Harvard Univ., Mus. Comp. Zool., Bul. 72:3-34.
—Wheeler, 1937. Harvard Univ., Mus. Comp. Zool., Bul. 81:441, 449-458, 463-465. —Smith,
1939. Ent. Soe. Amer., Ann. 32:502-509.

Taxonomy: Brown, 1973. In Meggers, et al., Tropical Forest Ecosystems in Afr. and S.
Amer., p. 181 (listed as a provisional syn. of Leptothorax).
floridana (Wheeler). Fla. (Paradise Key, Dade Co.). Ecology: Nest of the types was found in a
hollow branch of a small tree.
Antillaemyrmex floridanus Wheeler, 1931. Harvard Univ., Mus. Comp. Zool., Bul. 72:27.
3.

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:11.
polita Smith. Ariz. Ecology: One nest was found under the bark of a tree, but most specimens
have been taken on the ground.
Macromischa polita Smith, 1939. Ent. Soc. Amer., Ann. 32:503, 506. & .
subditiva Wheeler. La., Tex.; Mexico. Ecology: Nests have been found under willow bark, and
from dead hollow branches lying on the ground.
Macromischa subditiva Wheeler, 1903. Psyche 10:99. &.

Superfamily FORMICOIDEA 1391

Biology: Mitchell and Pierce, 1912. Ent. Soe. Wash., Proc. 14:73. —Smith, 1939. Ent. Soc.
Amer., Ann. 32:506. —Creighton, 1965. Psyche 72:282.

Genus ROGERIA Emery

Rogeria Emery, 1894. Soc. Ent. Ital., Bol. 26:188.
Type-species: Rogeria curvipubens Emery. Desig. by Wheeler, 1911.
Rogeria subg. Irogeria Emery, 1915. Soc. Ent. de France, Bul. p. 191.
Type-species: Rogeria procera Emery. Orig. desig.
A neotropical genus with about 24 species. Only two species reach the southwestern U. S.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:28 (larvae). —Kempf, 1961.
Rev. Brasil. Biol. 21:435-441. —Kempf, 1962. Studia Ent. 5:1-38. —Kempf, 1963. Rev.
Brasil. Biol. 23:189-196 (S. Amer. species). —Kempf, 1964. Studia Ent. 7:45-71. —Kempf,
1965. Rev. Brasil. Biol. 25:181-186. —Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci.
No. 236:1-6 (first record of genus in U. S.). —Wheeler and Wheeler, 1973. Psyche 80: 74
(larvae, revised description).

creightoni Snelling. Tex. (La Feria, Cameron Co.).
Rogeria creightoni Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. No. 236:2-4. 9.

huachucana Snelling. Ariz. (Cochise Co.). Ecology: Specimens found beneath stones.
Rogeria huachucana Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. No. 236:4-6. 9.

Genus LEPTOTHORAX Mayr

Other than those genera listed under Macromischa and the subgenera of Leptothorax, Brown
(1973) gives the following synonyms or possible synonyms: Chalepoxenus Menozzi, 1923?,
Doronomyrmex Kutter, 1945?, Epimyrma Emery, 1915?, Formicoxenus Mayr, 1855 ?, Gonepi-
myrma Bernard, 1948 ?, Icothorax Hamann and Klemm, 1967 ?, Leonomyrma Arnoldi, 1968 ?,
Myrmamnophilus Menozzi, 1924, Myrmetaerus Soudek, 1925 ?, Myrmoxenus Ruzksy, 1902 ?,
Symmyrmica Wheeler, 1904 ?, and Temnothorax Mayr, 1861. There is apparently considerably
more work to be done on this and related genera; consequently, I am using subgenera in Lep-
tothorax as has been done in past catalogs.

Taxonomy: Brown, 1973. In Meggers, et al., Tropical Forest Ecosystems in Afr. and S.

Amer., pp. 161-185.
Genus LEPTOTHORAX Subgenus NESOMYRMEX Wheeler

Leptothorax subg. Goniothorax Emery, 1896. Soc. Ent. Ital., Bol. 28:26, 58. Preoce. by

Milne-Edwards, 1879.
Type-species: Leptothorax vicinus Mayr. Desig. by Wheeler, 1911.
Nesomyrmex Wheeler, 1910. Amer. Mus. Nat. Hist., Bul. 28:259.
Type-species: Nesomyrmex clavipilis Wheeler. Monotypic.
Leptothorax subg. Caulomyrma Forel, 1914. Soc. Vaud. des Sci. Nat., Bul. 50:233.
Type-species: Leptothorax echinatinodis Forel. Orig. desig.
Limnomyrmex Arnold, 1948. Nat. Mus. So. Rhodesia, Occas. Papers 2 (14): 222.
Type-species: Limnomyrmex stramineus Arnold. Monotypic.

Most species of this subgenus are neotropical.

Revision: Kempf, 1959. Studia Ent. 2:391-432.
Taxonomy: Smith, 1950. Psyche 57:30. —Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann.
48:25 (larva). —Brown, 1971. Breviora 365:4-5 (generic syn.).
wilda Smith. S. Tex.; Mexico. Ecology: Arboreal, forms small colonies in plant cavities;

probably nocturnal.
Leptothorax (Goniothorax) wilda Smith, 1943. Ent. Soe. Wash., Proc. 45:155. §, 2.

Biology: Creighton, 1971. Ga. Ent. Soc., Jour. 6:207-210 (distribution and habits).
Genus LEPTOTHORAX Subgenus MYRAFANT Smith
Leptothorax subg. Myrafant Smith, 1950. Psyche 57:29.

1392 Hymenoptera in America North of Mexico

Type-species: Leptothorax curvispinosus Mayr. Orig. desig.

These small ants nest in small colonies, commonly in preformed cavities under bark, hollow
twigs, dried grass stems, old galls, or empty nut shells. A few may be found under rocks or in
the soil. Some species are lestobiotic, nesting near the nests of other ants, and others may be du-
lotic.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317-318, 320-323. —Wheeler, 1903.
Acad. Nat. Sci. Phila., Proc. 55:215, 224, 232-256.

Taxonomy: Smith, 1950. Psyche 57:29. —Smith, 1952. N. Y. Ent. Soc., Jour. 60:96-106
(tricarinatus - texanus complex). —Brown, 1955. Ent. News 66:43. —Wheeler and
Wheeler, 1955. Ent. Soe. Amer., Ann. 48:22-25 (larvae). —Cole, 1956. Tenn. Acad. Sci., Jour.
31:30-31 (tricarinatus - texanus complex). —Cole, 1958. Ent. Soc. Amer., Ann. 51:535-538
(nitens - carinatus complex). —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:23
(larvae).

ambiguus ambiguus Emery. Que. to Va., w. to N. Dak., S. Dak., Iowa, Nebr. Ecology: Nests in
soil in woodlands and grasslands or in hollow dead grass stems near the soil.
Leptothorax (Leptothorax) curvispinosus ambiguus Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:317, 320. 3.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22:588-589.
—Wesson and Wesson, 1940. Amer. Midland Nat. 24:97 (each caste). —Creighton, 1950.
Harvard Univ., Mus. Comp. Zool., Bul. 104:261-262. —Wheeler and Wheeler, 1955. Ent.
Soe. Amer., Ann. 48:22-23 (larva).

Biology: Sturtevant, 1925. Psyche 32:314. —Buren, 1944. Iowa State Col., Jour. Sci. 18:287.
—Kannowski, 1959. Insectes Sociaux 6:124, 150-151, 155 (parasitism; pleometrosis).
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 140-141.

ambiguus foveatus Smith. Ill. (Plainfield). Ecology: Found in nest of a species of
Aphaenogaster in a roadside ditch.
Leptothorax foveata Smith, 1934. Psyche 41:211. %.
ambiguus pinetorum Wesson and Wesson. Ohio (Jackson Co.).
Leptothorax ambiguus var. pinetorum Wesson and Wesson, 1940. Amer. Midland Nat.
24:97. 8, 2, d.
andrei Emery. N. Mex., Ariz., Nev., Calif. Ecology: Nests have been found under stones.
Leptothorax (Leptothorax) andrei Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 322. 9.

Taxonomy: Cole, 1958. Ent. Soc. Amer., Ann. 51:537-538 (each caste).

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:73. —Wheeler and Wheeler, 1973. Ants
of Deep Canyon, p. 95.
bradleyi Wheeler. Ga., Ala., Fla. Ecology: Colonies found in galleries in bark.
Leptothorax bradleyi Wheeler, 1913. Psyche 20:113. &.

Taxonomy: Wilson, 1952. Ent. News 63:68-71 (each caste).

carinatus Cole. W. Tex., Ariz. Ecology: Nests were found beneath stones.
Leptothorax (Leptothorax) carinatus Cole, 1957. Tenn. Acad. Sci., Jour. 32:213-215. 3, °.

Taxonomy: Cole, 1958. Ent. Soc. Amer., Ann. 51:537-538. —Wheeler and Wheeler, 1973.
Psyche 80: 70-71 (semipupa).
curvispinosus Mayr. Maine s. to Fla., w. to Iowa, Kans., Okla., Tex., Ariz. Ecology: Nests in
plant cavities such as hollow stems, twigs, and in acorns. Enslaved by Leptothorax
duloticus Wesson and Harpagoxenus americanus (Emery).
Leptothorax curvispinosus Mayr, 1866. Akad. der Wien, Math.-Nat. K1., Sitzber. 53:508. 8.
Stenamma gallarum Patton, 1879. Amer. Nat. 13:126. 8, °.

Taxonomy: Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36:451, 453 (worker, female). —Cole,
1940. Amer. Midland Nat. 24:56-57.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:385. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31:289, 305. —Headley, 1943. Ent. Soc. Amer., Ann. 36:743-753 (population studies).

Superfamily FORMICOIDEA 1393

—Talbot, 1957. Ecology 38:449-456 (populations). —Kannowski, 1959. Insectes Sociaux
6:124. —Wilson and Fagan, 1974. N. Y. Ent. Soc., Jour. 82: 106-112 (estimation of total
behavioral repertories).

furunculus Wheeler. Wyo., Colo.
Leptothorax furunculus Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:82. 9.

Biology: Gregg, 1963. Ants of Colo., pp. 381-382.
gallae Smith. Calif. Ecology: Commonly found on the canyon live oak, Quercus chrysolepis,
often in twig galls made by cynipids.
Leptothorax (Leptothorax) gallae Smith, 1949. Psyche 56:112. 8.
hispidus Cole. W. Tex.; Mexico. Ecology: Nests were found under stones at higher elevations.
Leptothorax (Leptothorax) hispidus Cole, 1957. Tenn. Acad. Sci., Jour. 32:42-45. 9, 9.

Taxonomy: Wheeler and Wheeler, 1973. Psyche 80: 71 (immatures).
longispinosus Roger. Que., Ont. s. to Ga., Ala. w. to Iowa. Ecology: Nests in plant cavities such

as hollow stems, twigs, and in acorns. Enslaved by Leptothorax duloticus Wesson and
Harpagoxenus americanus (Emery).

Leptothorax longispinosus Roger, 1863. Berlin. Ent. Ztschr. 7:180. &.

Leptothorax (Leptothorax) longispinosus laeviceps Buren, 1944. Iowa State Col., Jour. Sci.
18:286. &.

Leptothorax longispinosus iowensis Buren, 1945. Ent. Soc. Wash., Proc. 47:288. N. name
for laeviceps, thought to be preocce. by leviceps Emery, 1898.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22:588. —Cole, 1940.
Amer. Midland Nat. 24:56-57. —Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:24
(larva).

Biology: Wheeler, 1910. Ants, pp. 212, 222, 495, 504. —Dennis, 1938. Ent. Soc. Amer., Ann.
31:274, 289, 305. —Headley, 1943. Ent. Soc. Amer., Ann. 36:743-753 (population studies).
—Kannowski, 1959. Insectes Sociaux 6:125. —Lettendre and Pilon, 1972. Nat. Canad.
99:73-82 (ecology; in Que.).

mariposa Wheeler. Calif. (Yosemite Natl. Pk.). Ecology: Colonies were found under stones.
Leptothorax nitens var. mariposa Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:507.
S.

Taxonomy: Cole, 1958. Ent. Soc. Amer., Ann. 51:536.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:73-74.
minutissimus Smith. D. C. The type series of several females was associated with L.
curvispinosus workers, but the exact relationship is not known.
Leptothorax minutissimus Smith, 1942. Ent. Soc. Wash., Proc. 44:59. 2.
nevadensis eldoradensis Wheeler. Calif. (Coastal Range and lower elevations of the Sierras).
Leptothorax eldoradensis Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:414. &.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:73.
nevadensis melanderi Wheeler. W. Mont. to e. Wash.
Leptothorax melanderi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:81. %.
nevadensis nevadensis Wheeler. Wash., Oreg., Calif. (Eastern slopes of Sierras north to
Cascade Mtns.). Ecology: Nests in soil, usually under stones.
Leptothorax nevadensis Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 252. %, Oe

Taxonomy: Wheeler, 1909. N. Y. Ent. Soc., Jour. 17:81. —Wheeler and Wheeler, 1973. Psyche
80: 71 (larva).

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 95-96 (subspecies not
recognized).
nevadensis rudis Wheeler. Nev., Calif. (Sierras from Lake Tahoe to Sequoia Park).
Leptothorax nevadensis rudis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:508. %,
OF

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:73. —Cole, 1966. Brigham Young Univ.,
Sci. Bul., Biol. Ser. 7:17.

1394 Hymenoptera in America North of Mexico

nitens Emery. Wyo., Colo., Tex. w. to Wash., Oreg., Calif. Ecology: Found under rocks and in
duff.
Leptothorax (Leptothorax) nitens Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, Soe or
Leptothorax nitens occidentalis Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:233, 245. 3.
Leptothorax nitens var. heathii Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 245. &.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28:369-370. —Cole, 1958. Ent. Soc. Amer., Ann.
51:536. —Wheeler and Wheeler, 1973. Psyche 80: 71, 73 (larva).

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:73. —Gregg, 1963. Ants of Colo., pp.
382-384. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 96.
obliquicanthus Cole. Colo., N. Mex. Ecology: Found under stones in meadows and prairies.
Leptothorax (Myrafant) obliquicanthus Cole, 1953. Ent. Soc. Wash., Proc. 55:28-30. %.

Taxonomy: Gregg, 1953. Breviora 22:1-3.

Biology: Gregg, 1963. Ants of Colo., pp. 384-385, 387.

obturator Wheeler. Tex. Ecology: One colony was found in an oak gall.
Leptothorax obturator Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 249. 8, °, d.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:24 (larva).

Biology: Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19:663-664. —Wheeler, 1910. Ants, pp.
208-209.
rugatulus brunnescens Wheeler. N. Dak., Mont. s. to Colo., Utah. Ecology: Nests under rocks
or wood. Sometimes not distinguished from the typical subspecies in the literature.
Leptothorax rugatulus brunnescens Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.
52:510. 8.

Leptothorax rugatulus dakotensis Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11:247.
%.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:25 (larva).
Biology: Gregg, 1963. Ants of Colo., pp. 388-389.

rugatulus rugatulus Emery. N. Dak., S. Dak., Colo., N. Mex. w. to B. C., Calif. Ecology: Nests
under stones, wood, in decaying wood, in grasses.
Leptothorax (Leptothorax) rugatulus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317, 321.
3.
Leptothorax curvispinosus rugatulus var. Cockerelli Wheeler, 1903. Acad. Nat. Sci. Phila.,
Proc. 55:241. 8, 2.
Leptothorax curvispinosus annectens Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:242. 9.

Leptothorax rugatulus var. mediorufus Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.
52:510. 9, 2.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 241. —Cole, 1942. Amer.
Midland Nat. 28:369. —Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104:267-269.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:509. —Cole, 1934. Psyche 41:222.

—Cole, 1954. Tenn. Acad. Sci., Jour. 29:240. —Wheeler and Wheeler, 1963. Ants of N. Dak.,

pp. 141-143 (does not distinguish subspecies). —Gregg, 1963. Ants of Colo., pp. 386-388.
schaumi Roger. Maine to Ga, w. to Iowa., Kans., Tex. Ecology: Commonly nests in the bark of

trees.

Leptothorax schaumi Roger, 1863. Berlin. Ent. Ztschr. 7:180. %.

Leptothorax fortinodis Mayr, 1886 Zool.-Bot. Gesell. Wien, Verh. 36:451. 3, °.

Leptothorax fortinodis var. gilvus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:235. %, oF

Leptothorax fortinodis var. melanoticus Wheeler, 1903. Acad. Nat. Sci. Phila. Proc. 55:235.
UEeoe

Taxonomy: Wesson and Wesson, 1940. Amer. Midland Nat. 24:94-96. —Cole, 1940. Amer.
Midland Nat. 24:56. —Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104:269-271.
—Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:23 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:385. —Wheeler, 1916. Ind. Acad. Sci.,
Proc. 26:461.

Superfamily FORMICOIDEA 1395

schmittii Wheeler. Colo. (Canyon City).
Leptothorax Schmittii Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 242. 9.
Taxonomy: Gregg, 1963. Ants of Colo., pp. 388, 390-391.
silvestrii (Santschi). S. Ariz. Ecology: Nests in evergreen oaks at altitudes of 3500 ft. or more.
Tetramorium silvestrii Santschi, 1909. Soc. Ent. Ital., Bol. 41:6. &.

Taxonomy: Creighton, 1953. Amer. Mus. Novitates 1635:1-7 (each caste described; note on
habitat).
stenotyle Cole. Ariz. (nr. Rustler’s Park, Chiricahua Mtns.). Ecology: Nests were found under

stones.
Leptothorax (Leptothorax) angustinodus Cole, 1956. Tenn. Acad. Sci., Jour. 31:28-30. 9, 2.

Preoce. by Stitz, 1917.
Leptothorax (Leptothorax) stenotyle Cole, 1956. Tenn. Acad. Sci., Jour. 31:214. N. name.

terrigena Wheeler. Tex. (Austin).

Leptothorax terrigena Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 254. 9, 2.
texanus davisi Wheeler. N. Y., N. J., Fla.

Leptothorax texanus davisi Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:385. %, °.

Taxonomy: Smith, 1952. N. Y. Ent. Soc., Jour. 60:104-106.
texanus texanus Wheeler. Mich. (?), Ohio, s. to N. C., Ga. w. to Okla., Tex. Ecology: Nests have
been found in sandy soil.
Leptothorax texanus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223-245. 9, 2, d.

Taxonomy: Gregg, 1944. Ent. Soc. Amer., Ann. 37:446. —Smith, 1952. N. Y. Ent. Soc., Jour.
60:102-104. —Wheeler and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:24 (larva).

Biology: Smith, 1932. Ent. News 43:160. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:98.

tricarinatus neomexicanus Wheeler. W. Colo., Utah, N. Mex., Ariz. Ecology: Nests found in
soil, under stones, in open grassy areas.

Leptothorax neomexicanus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223-248. 9.

Taxonomy: Smith, 1952. N. Y. Ent. Soc., Jour. 60:100-102.

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22:341. —Cole, 1953. Ent. Soc. Wash.,
Proc. 55:28. —Gregg, 1963. Ants of Colo., pp. 393-394.
tricarinatus tricarinatus Emery. N. Dak., S. Dak., Iowa w. to Wyo., Colo., Utah. Ecology:

Nests under rocks and in soil.
Leptothorax (Leptothorax) tricarinatus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318,

321. 9:

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 247-248. —Buren, 1944. Iowa
State Col., Jour. Sci. 18:286, 288. —Smith, 1952. N. Y. Ent. Soc., Jour. 60:98-100 (worker,
male).

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., p. 143. —Gregg, 1963. Ants of Colo.,
pp. 391-393.

tuscaloosae Wilson. N. C., Ala. Ecology: Colony was found in small cavity in soil under a bed

of moss at base of an oak tree.
Leptothorax (Myrafant) tuscaloosae Wilson, 1950. Psyche 57:128-130. %, °.

wheeleri Smith. N. C., Ga, Fla., Ohio, Tenn., Ala., Miss. Ecology: Colonies found in cavities in

trees and under bark.
Leptothorax wheeleri Smith, 1929. Ent. Soc. Amer., Ann. 22:547. 8, 2.

Taxonomy: Wilson, 1952. Ent. News 60:67-68, 70 (worker, male).

Biology: Smith, 1931. Ent. News 42:18. —Wesson and Wesson, 1940. Amer. Midland Nat.
24:90, 96.

Genus LEPTOTHORAX Subgenus DICHOTHORAX Emery
Leptothorax subg. Dichothorax Emery, 1895. Zool. Jahrb., Abt. f. System. 8:323.

1396 Hymenoptera in America North of Mexico

Type-species: Leptothorax (Dichothorax) pergandei Emery. Desig. by Wheeler,
1911

The status of the two forms below needs clarification. Though regarded as subspecies, the
ranges overlap considerably.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 323-324. —Wheeler, 1903. Acad.
Nat. Sci. Phila., Proc. 55:224, 256-260.
pergandei floridanus Emery. N. J., N. C. to Fla., w. to Tex. Ecology: Has been found in
stumps, logs, and nut shells.
Leptothorax (Dichothorax) floridanus Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 324.
3.
Leptothorax (Dichothorax) pergandei flavus Smith, 1929. Ent. Soc. Amer., Ann. 22:549. 9,
2.
Leptothorax (Dichothorax) pergandei floridanus var. spinosus Smith, 1929. Ent. Soc.
Amer., Ann. 22:551. &.
Taxonomy: Smith, 1931. Ent. News 42:18-19.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:290, 305.
pergandei pergandei Emery. D. C. s. to Ga., Tenn., w. to Nebr., Tex. Ecology: Apparently a
ground nesting species.

Leptothorax (Dichothorax) pergandei Emery, 1895. Zool. Jahrb., Abt. f. System. 8:318, 323.
3

Leptothorax (Dichothorax) manni Wesson, 1935. Ent. News 46:208. %, 2, ¢. Preocc. by
Wheeler, 1914.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:224, 256-259 (worker, female).
—Smith, 1924. Ent. News 35:50. —Cole, 1940. Amer. Midland Nat. 24:56, 58. —Wheeler
and Wheeler, 1960. Ent. Soc. Wash., Proc. 62:23 (larva).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:290. —Buren, 1944. Iowa State Col., Jour.
Sci. 18:288.

Genus LEPTOTHORAX Subgenus LEPTOTHORAX Mayr

Leptothorax Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:431.
Type-species: Formica acervorum Fabricius. Desig. by Bingham, 1903.
Mychothorax Ruzsky, 1904. Sapiski Imp. Russ. Geog. Obshch. 41:288.
Type-species: Formica acervorum Fabricius. Orig. desig.

In North America, this subgenus is confined mostly to the northern and western United
States and Canada. The ants nest in living and dead trees, decaying wood, soil, or stumps. Some
are inquilines in nests of other ants and some are dulotic.

Taxonomy: Brown, 1955. Ent. News 66:43-50. —Wheeler and Wheeler, 1955. Ent. Soc. Amer.,
Ann. 48:21-22 (larvae).
crassipilis Wheeler. Wyo., Colo., N. Mex., Utah, Ariz., Nev. Ecology: Found under rocks, under
wood, in decaying logs.
Leptothorax (Mychothorax) acervorum crassipilis Wheeler, 1917. Amer. Acad. Arts and
Sci., Proc. 52:513. 3, 2, d.
Taxonomy: Cole, 1954. Tenn. Acad. Sci., Jour. 29:240-241.
Biology: Cole, 1953. Ent. Soc. Wash., Proc. 55:27. —Gregg, 1963. Ants of Colo., pp. 401-402,
405.

diversipilosus Smith. Wash. Ecology: An inquiline in nests of Formica obscuripes Forel.
Leptothorax (Mychothorax) diversipilosus Smith, 1939. Ent. Soc. Wash., Proc. 41:179. 8,
ergatoid °.

Taxonomy: Smith, 1956. Ent. Soc. Wash., Proc. 58:271-275 (female, ergatoid male).

Biology: Alpert and Akre, 1973. Ent. Soc. Amer., Ann. 66:753-760 (distribution, abundance,
behavior).

Superfamily FORMICOIDEA 1397

duloticus Wesson. Mich., Ohio, Ill. (?). Ecology: Dulotic, enslaves Leptothorax curvispinosus
Mayr and L. longispinosus Roger.
Leptothorax (Mychothorax) duloticus Wesson, 1937. Ent. News 48:125. 9, 2.

Taxonomy: Wesson, 1940. Brooklyn Ent. Soc., Bul. 35:81-83 (male, biology).

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24:94. —Talbot, 1957. Ecology
38:449-456 (populations). —Kannowski, 1959. Insectes Sociaux 6:124-215, 150-151
(parasitism).

hirticornis Emery. N. Dak., S. Dak., Colo., Utah., Calif. Ecology: An inquiline in nests of
Formica obscuripes Forel and possibly Formica integroides integroides Emery.
Leptothorax (Leptothorax) hirticornis Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317, 319.
om
Leptothorax (Leptothorax) hirticornis formidolosus Wheeler, 1915. Amer. Mus. Nat. Hist.,
Bul. 34:415. 8, ergatoid °.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223-224. —Smith, 1939. Ent. Soc.
Wash., Proc. 41:176-179 (worker, ergatoid female).

Biology: Weber, 1935. Ecol. Monog. 5:200. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart.
11:248. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 145-146. —Gregg, 1963. Ants of
Colo., pp. 403, 405. —Snelling, 1965. South. Calif. Acad. Sci., Bul. 64:16 (also description of
dealate female and ergatoid male).

muscorum (Nylander). Newfoundland (Labrador) w. to N. W. T., Alaska, s. to Conn., Wis., N.
Mex., Ariz., Calif.; Eurasia. Ecology: Has been found at 69° 22’ N., Kidluit Bay on
Richards Is., the northernmost record of any New World ant. Mostly found in
woodlands, in decaying stumps, logs, under bark of fallen trees, or under rocks. It is
best able to survive in extreme arctic conditions under stones.

Myrmica muscorum Nylander, 1846. Acta Soc. Sci. Fenn. 2:1054. 3, 2, d.

Leptothorax canadensis Provancher, 1887. Addit. Corr. Faune Ent. Canada, Hym. p. 245.
Perr

Leptothorax (Leptothorax) canadensis var. yankee Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:317, 319. 8, 2.

Leptothorax yankee var. kincaidi Pergande, 1900. Wash. Acad. Sci., Proc. 2:520. 3, 9.

Leptothorax muscorum var. sordidus Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223,
224. 9.

Leptothorax acervorum canadensis var. convivialis Wheeler, 1903. Acad. Nat. Sci. Phila.,
Proc. 55:228. ¥%, 9.

Leptothorax canadensis obscurus Viereck, 1903. Amer. Ent. Soc., Trans. 29:72. 9.

Leptothorax (Leptothorax) acervorum canadensis var. Calderoni Forel, 1914. Deut. Ent.
Ztschr., p. 617. 8, @.

Leptothorax (Mychothorax) muscorum var. septentrionalis Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52:511. %, 9, do.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 225-229. —Wheeler, 1917.
Conn. State Geol. and Nat. Hist. Survey Bul. 22:588. —Wheeler, 1917. Amer. Acad. Arts
and Sci., Proc. 52:512-513. —Brown, 1955. Ent. News 66:47-50 (also ecology). —Wheeler
and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:21 (larva). —Francoeur and Beique, 1966.
Canad. Ent. 98:142 (Provancher ant types).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:621. —Wheeler and Wheeler, 1944. N.
Dak. Hist. Quart. 11:247-248. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:73. —Cole,
1942. Amer. Midland Nat. 28:369-370. —Gregg, 1946. Amer. Midland Nat. 35:748.
—Kannowski, 1959. Insectes Sociaux 6:125. —Wheeler and Wheeler, 1963. Ants of N. Dak,
pp. 143-145. —Gregg, 1963. Ants of Colo., pp. 393-400. —Chapman, 1969. Ent. Soc. Amer.,
Ann. 62:1256-1259 (release and recovery of marked winged forms). —Moglich, Machwitz,
and Holldobler, 1974. Science 186: 1046-1047 (tandem calling).

1398 Hymenoptera in America North of Mexico

provancheri Emery. Que., Maine w. to Alta., N. Dak., Colo., N. Mex. Ecology: Apparently an
inquiline, found in nests of Myrmica incompleta incompleta Provancher and M.
lobicornis fracticornis Emery.
Leptothorax (Leptothorax) provancheri Emery, 1895. Zool. Jahrb., Abt. f. System. 8:317,
320. &.
Leptothorax emersoni Wheeler, 1901. Amer. Nat. 35:433. %, 2, d.
Leptothorax emersoni glacialis Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. (n. s.) 5:71. 8, 2,

3.
Leptothorax (Mychothorax) emersoni hirtipilis Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52:515. %.

Taxonomy: Wheeler, 1903. Acad. Nat. Sci. Phila., Proc. 55:223, 229. —Cole, 1954. Tenn. Acad.
Sci., Jour. 29:241. —Wheeler and Wheeler, 1973. Psyche 80: 73-74 (larva).

Biology: Wheeler, 1907. Wis. Nat. Hist. Soc., Bul. (n. s.) 5:78-83. —Wheeler, 1910. Ants, pp.
107, 393, 434-436. —Wheeler, 1901. Amer. Nat. 35:436-438. —Wheeler, 1903. Jour. Psychol.
and Neurol. 2:1-21. —Kannowski, 1957. Psyche 64:1-5. —Wheeler and Wheeler, 1963. Ants
of N. Dak., pp. 146-148. —Gregg, 1963. Ants of Colo., pp. 404-407.

Genus SYMMYRMICA Wheeler

Symmyrmica Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:1, 3.
Type-species: Symmyrmica chamberlini Wheeler. Monotypic.
Taxonomy: Brown, 1973. In Meggers et al., Tropical Forest Ecosystems in Afr. and S.
Amer., p. 185 (possible syn. of Leptothorazx).
chamberlini Wheeler. Utah, Oreg. Ecology: An inquiline in nest of Manica mutica (Emery).
Symmyrmica chamberlini Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:5. %, 2, apterous
ergatoid d.

Biology: Wheeler, 1910. Ants, pp. 432-434. —Wheeler, 1919. Amer. Phil. Soc., Proc. 58:22.
Genus HARPAGOXENUS Forel

Tomognathus Mayr, 1861. Die Europaischen Formiciden, pp. 29, 56. Preoce. by Dixon,
1850.
Type-species: Myrmica sublaevis Nylander. Monotypic.
Harpagoxenus Forel, 1893. Soc. Ent. de Belg., Ann. 37:167. N. name for Tomognathus.
Protomognathus Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:3.
Type-species: Tomognathus americanus Emery. Monotypic.

Ants of this genus are dulotic, enslaving and conducting raids on colonies of certain species of
Leptothorax. There is a single European species.

Revision: Smith, 1939. Ent. Soc. Wash., Proc. 41:165-172.

Taxonomy: Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:26-28 (larvae).

americanus (Emery). Mass., Ont. s. to N. C., w. to IIL, Mo. Ecology: Enslaves Leptothorax
curvispinosus Mayr and L. longispinosus Roger.
Tomognathus americanus Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 272 9.

Taxonomy: Creighton, 1927. Psyche 34:28 (male). —Smith, 1939. Ent. Soc. Wash., Proc.
41:166-168 (each caste). —Wheeler and Wheeler, 1955. Ent. Soc. Amer., Ann. 48:26-27
(larva).

Biology: Sturtevant, 1927. Psyche 34:1-9. —Creighton, 1927. Psyche 34:11-29. —Creighton,
1923. Psyche 36:48-50. —Wesson, 1939. Amer. Ent. Soc., Trans. 65:97-122.

canadensis Smith. N. S., Que., Maine w. to Mich., Minn. Ecology: Enslaves Leptothorax
muscorum (Nylander). Apparently closely related to the Palearctic H. sublaevis
(Nylander) and like that form has ergatoid females in addition to normal workers and

females.
Harpagoxenus canadensis Smith, 1939. Ent. Soc. Wash., Proc. 41:168. 2, ergatoid 9.

Taxonomy: Gregg, 1945. Canad. Ent. 77:74-76.

Superfamily FORMICOIDEA 1399

Biology: Gregg, 1946. Amer. Midland Nat. 35:748.
TRIBE MYRMECININI

Genus MYRMECINA Curtis

Myrmecina Curtis, 1829. Brit. Ent. 6:226, pl. 265.
Type-species: Myrmecina latreillii Curtis. Orig. desig.

Archaeomyrmex Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bul. 64:448, 451.
Type-species: Archaeomyrmex cacabau Mann. Orig. desig.

Apparently only one species occurs in North America, though several subspecies are some-
times recognized.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:271. —Smith, 1948. Ent. Soc. Wash.,
Proc. 50:238-240. —Snelling, 1965. South. Calif. Acad. Sci., Bul. 64:101-105. —Brown, 1967.
Ent. News 78:233-230.

Taxonomy: Brown, 1951. Brooklyn Ent. Soc., Bul. 46:103-106. —Wheeler and Wheeler, 1954.
Ent. Soe. Wash., Proc. 56:129-131 (larvae). —Brown, 1971. Breviora 365:1-2 (generic syn.).
americana Emery. Que. s. to Ga., w. to Iowa, Colo., N. Mex., Ariz., Calif. Ecology: Colonies are
small and nests are obscure, usually built in moist shady areas often under small stones.
Myrmecina latreillei americana Emery, 1895. Zool. Jahrb., Abt. f. System. 8:271. &.
Myrmecina latreillei americana var. brevispinosa Emery, 1895. Zool. Jahrb., Abt. f.
System. 8:271. 3, 9, d.
Myrmecina graminicola texana Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:422. 9.
Myrmecina graminicola quadrispina Enzmann, 1946. N. Y. Ent. Soc., Jour. 54:13. 8.
Myrmecina californica Smith, 1948. Ent. Soc. Wash., Proc. 50:239. §.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24:39. —Buren, 1944. Iowa State Col., Jour. Sci.
18:290. —Brown, 1949. Psyche 56:44-47. —Brown, 1951. Brooklyn Ent. Soc., Bul.
46:103-106. —Wheeler and Wheeler, 1954. Ent. Soc. Wash., Proc. 56:130 (larva).
—Francoeur, 1966. Nat. Canad. 93:439.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:373, 376. —Wheeler, 1906. Amer. Mus.
Nat. Hist., Bul. 22:332, 335-336. —Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:502.
—Talbot, 1934. Ecology 15:420, 427-428. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:274,
278-279, 304. —Gregg, 1963. Ants of Colo., pp. 364-365, 367.

TRIBE TETRAMORIINI
Revision: Bolton, 1976. Brit. Mus. (Nat. Hist.) Ent., Bul. 34: 283-379.
Genus TRIGLYPHOTHRIX Forel

Triglyphothrix Forel, 1890. Soc. Ent. de Belg., Ann. 34: CVI.
Type-species: Triglyphothrix walshi Forel. Monotypic.

A single introduced form is established in a number of localities in southeastern United
States. The genus is native to Africa and southern Asia.

Revision: Bingham, 1903 Fauna Brit. India, Hym. 2:172-175. —Bolton, 1976. Brit. Mus. (Nat.
Hist.) Ent., Bul. 34: 310-359 (world species).
lanuginosa (Mayr). S. C., Ga., Fla. Ala, Miss., La.; pantropical. Introduced, probably native to
India. Spread by commerce to most tropical regions of the world.
Tetramorium lanuginosa Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 972, 976. %.
Tetramorium obesum race striatidens Emery, 1889. Ann. Mus. Civ. Stor. Nat. Genova
7:501. &.

Taxonomy: Wheeler, 1912. N. Y. Ent. Soc., Jour. 20:46. —Wheeler, 1916. Jour. Econ. Ent.
9:568-569 (first notice of occurrence in U. S.). —Wilson and Taylor, 1967. Pacific Ins.
Monog. 14:70. —Wheeler and Wheeler, 1973. Psyche 80: 78 (larva). —Bolton, 1976. Brit.
Mus. (Nat. Hist.) Ent., Bul. 34: 350-352 (further synonymy).

Biology: Donisthorpe, 1927. British Ants, p. 393. —Smith, 1931. Ent. News 42:21.

1400 Hymenoptera in America North of Mexico

Genus TETRAMORIUM Mayr

Tetramorium Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5:423.
Type-species: Formica caespitum Linnaeus. Desig. by Girard, 1879.

Tetrogmus Roger, 1857. Berlin. Ent. Ztschr. 1:10.
Type-species: Tetrogmus caldarius Roger. Monotypic.

Tetramorium subg. Xiphomyrmex Forel, 1887. Schweiz. Ent. Gesell. Mitt. 7: 385.
Type-species: Tetramorium (Xiphomyrmex) kelleri Forel. Desig. by Wheeler, 1911.

This genus is better represented in Asia and Africa than in North America. Three of the four
species, guineense, pacificum, and simillimum, are, with little question, introduced; the other,
caespitum, which is also found in Europe, is generally believed to be native, though some wor-
kers also consider it as being introduced. This genus contains some important house pests.

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:972-977. —Bingham, 1903. Fauna
British India, Hym. 2:175-189. —Emery, 1916. Soc. Ent. Ital., Bol. 47:195. —Smith, 1938.
Wash. Acad. Sci., Jour. 28: 126-130 (Xiphomyrmex). —Smith, 1943. Ent. Soe. Wash., Proce.
45:1-5 (U. S. species).

Taxonomy: Wheeler and Wheeler, 1954. Amer. Midland Nat. 52: 445, 450 (larvae). —Cole,
1957. Tenn. Acad. Sci., Jour. 32: 209-219 (male, Xiphomyrmex). —Brown, 1957. Breviora
72:1-8 (native in N. Amer. ?). —Bolton, 1976. Brit. Mus. (Nat. Hist.) Ent., Bul. 34: 288,
359-365 (generic synonymy).

caespitum (Linnaeus). Ont. s. to Tenn., w. to Nebr., Mo., Wash., Nev., Calif.; Eurasia, Africa.

Ecology: One of the most common house-infesting ants in the large cities of the Atlantic
coast. They also steal seeds from seedbeds, gnaw into tubers, roots, and stalks of
various plants, attend honeydew excreting insects, and serve as an intermediate host for
poultry tapeworms. Colonies are populous and nests may be in exposed soil, under cover
of stones, pavement, or other objects, in rotting wood, or next to building foundations.
Possibly introduced by early colonists from Europe but believed by some to be a native
species. Parasite: Anergates atratulus (Schenck), Strongylognathus sp. Pavement ant.
Most common in the Atlantic seaboard states and more sparsely distributed inland.

Formica caespitum Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 581.

Myrmica (Myrmica) brevinodis var. transversinodis Enzmann, 1946. N. Y. Ent. Soc., Jour.
54:47. 3.

Taxonomy: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:972. —Donisthorpe, 1927. British
Ants, p. 189 (each caste). —Smith, 1943. Ent. Soc. Wash., Proc. 45:2. —Brown, 1949. Psyche
56:47. —Wheeler and Wheeler, 1954. Amer. Midland Nat. 52:445 (larva). —Wheeler and
Wheeler, 1960. Ent. Soc. Wash., Proc. 62:24-25 (larva). —Wheeler and Wheeler, 1973.
Psyche 80: 76, 78 (larva).

Biology: Smith, 1915. Va. Truck Expt. Sta., Bul. 16:1-15. —Wheeler, 1919. Amer. Phil. Soc.,
Proc. 58:23-26. —Wheeler, 1927. Psyche 34:164-165. —Donisthorpe, 1927. British Ants, pp.
193-198. —Metcealf and Flint, 1939. Destructive and Useful Insects, p. 771. —Mallis, 1941.
South. Calif. Acad. Sci., Bul. 40:74. —Brown, 1964. Ent. News 75:15. —Brown, 1957.
Breviora 72:1-4 (historical). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:45-47
(economic importance). —Lange, 1961. Jour. Econ. Ent. 54:1063. —Poldi, 1963. Pavia Univ.
Symposium Genet. et Biol. Ital. 12:182. —Weber, 1965. Ent. News 76: 137-139 (in
Philadelphia area). —Brian, Elmes, and Kelley, 1967. Jour. Anim. Ecol. 36:337-342
(populations). —Bruder and Gupta, 1972. Ent. Soc. Amer., Ann. 65:358-367. —Gurney, 1975.
Ins. World Digest 2 (5): 19-25 (stinging habits).

guineense (Fabricius). Ga., Fla. w. to Tex.; pantropical. Ecology: Nests in small to moderate
colonies in exposed soil, under stones or other objects, in rotting logs and stumps, in
stems of plants, and in branches and under bark of trees. Occasionally a house pest.
Introduced, spread throughout the tropical regions of the world by commerce.
Occasionally found in greenhouses farther north in N. Amer. Guinea ant.
Formica guineensis Fabricius, 1793. Ent. System. 2:357. 8.

Taxonomy: Emery, 1909. Deut. Ent. Ztschr., p. 695 (each caste). —Smith, 1936. Puerto Rico
Univ., Jour. Agr. 20:831, 852. —Smith, 1943. Ent. Soc. Wash., Proc. 45:3. —Wilson and

Superfamily FORMICOIDEA 1401

Taylor, 1967. Pacific Ins. Monog. 14:71-72 (Polynesia). —Wheeler and Wheeler, 1954. Amer.
Midland Nat. 52:449-450 (larva).

Biology: Marlatt, 1928. U.S. Dept. Agr., Farmers’ Bul. 740:6. —Phillips, 1934. (Hawaii Univ.)
Expt. Sta. Pineapple Prod. Coop. Assoc. Bul. 15:23-24. —Smith, 1943. Ent. Soc. Wash.,
Proc. 45:1-2. —Brown, 1958. Acta Hym. 1:28. —Taylor and Wilson, 1961. Psyche 68:138.
—Brown, 1964. Ent. News 75:14-15. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:47-48
(economic importance).

Morphology: Blum and Ross, 1965. Jour. Ins. Physiol. 11:857-868 (odor trail pheromone).
pacificum Mayr. Calif.; Oriental and Australian Regions. Introduced into a nursery in Calif;
may not be established.
Tetramorium pacificum Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20:972, 976. 3, °.

Taxonomy: Smith, 1943. Ent. Soc. Wash., Proce. 45:2-3. —Wilson and Taylor, 1967. Pacific Ins.

Monog. 14:72-73 (Polynesia).
simillimum (Smith). Ga., Fla.; Pantropical. Introduced, a tramp species probably native to
Africa. Occasionally found in greenhouses farther north in N. Amer.
Myrmica simillima Smith, 1851. List Hym. Brit. Mus. v. 6, p. 118. 8.

Taxonomy: Emery, 1909. Deut. Ent. Ztschr., pp. 695-696 (each caste). —Smith, 1936. Puerto
Rico Univ., Jour. Agr. 20:831, 853. —Smith, 1943. Ent. Soc. Wash., Proc. 45:2. —Wilson and
Taylor, 1967. Pacific Ins. Monog. 14:73 (Polynesia).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:101. —Marlatt, 1928. U. S. Dept. Agr.,
Farmers’ Bul. 740:6. —Smith, 1933. Fla. Ent. 17:24. —Phillips, 1934. (Hawaii Univ.) Expt.
Sta. Pineapple Prod. Coop. Assoc. Bul. 15:24. —Taylor and Wilson, 1961. Psyche 68:142-143.

spinosus hispidus (Wheeler). S. Ariz. Ecology: Found nesting in small craters in the desert. X.
spinosus spinosus Pergande occurs in Mexico.
Xiphomyrmex spinosus hispidus Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:415. 8.
spinosus insons (Wheeler). W. Tex., Ariz. Ecology: Nests found in small craters in dry, grassy

areas.
Xiphomyrmex spinosus insons Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:416. 3, 2, d.

spinosus wheeleri Forel. S. Ariz.; Mexico. Ecology: One colony was found beneath a stone in a
cactus desert.
Tetramorium (Xiphomyrmex) Wheeleri Forel, 1901. Soc. Ent. de Belg., Ann. 45:128. &.

Biology: Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:416.
Genus ANERGATES Forel

Anergates Forel, 1874. Schweiz. Naturf. Gesell. Denkschr. 26:93.
Type-species: Myrmica atratula Schenck. Monotypic.
The single species of this genus is a permanent parasite.
atratulus (Schenck). Conn., N. Y., N. J., Pa., Del., Md., D. C., Va; Europe. Ecology: A
parasitized nest of the host consists of a single fertile female of atratulus, a large
number of host workers, and a large number of pupoidal males and virgin females of
atratulus. Workers of the parasite are entirely lacking. Possibly introduced with its host
from Europe, though some workers are of the opinion that the host and atratulus are
native. Host: Tetramorium caespitum (L.).
Myrmica atratula Schenck, 1852. Nassau. Ver. f. Naturk. Jahrb. 8:91. °.
Anergates friedlandi Creighton, 1934. Psyche 41:193. °.
Taxonomy: Emery, 1922. In Wytsman, Gen. Ins., fasc. 174:205-206 (female, male).
—Donisthorpe, 1927. British Ants, pp. 96-97 (female, male). —Wheeler and Wheeler, 1955.
Amer. Midland Nat. 54:128-130 (larva). —Ettershank, 1966. Austral. Jour. Zool. 14:157-158.

Biology: Wheeler, 1910. Ants, pp. 498-504. —Donisthorpe, 1915. British Ants, p. 89.
—Wheeler, 1923. Social life Among the Insects, pp. 215-219. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bul. 104:241-243.

Morphology: Meyer, 1955. Insectes Sociaux 2:163-170.

1402 Hymenoptera in America North of Mexico
TRIBE OCHETOMYRMICINI

Genus OCHETOMYRMEX Mayr

Ochetomyrmex Mayr, 1877. Zool.-Bot. Gesell. Wien, Verh. 27:871.
Type-species: Ochetomyrmex semipolitus Mayr. Monotypic.
Wasmannia Forel, 1893. London Ent. Soc., Trans. 4:383.
Type-species: Tetramorium? auropunctatum Roger. Desig. by Wheeler, 1911.
Blepharidatta Wheeler, 1915. Harvard Univ., Mus. Comp. Zool., Bul. 59:484.
Type-species: Blepharidatta brasiliensis Wheeler. Monotypic.
Hercynia Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:43. Preoce. by Rafinesque, 1815.
Type-species: Hercynia panamana Enzmann. Monotypic.

A single species of this Neotropical genus has been introduced into the United States.

Taxonomy: Brown, 1948. Ent. News 59:102. —Brown, 1973. In Meggers, et al., Tropical
Forest Ecosystems in Afr. and S. Amer., pp. 178-185 (generic syn.).
auropunctata (Roger). Fla., Calif.; W. Indies, Mexico, Central and S. Amer. Ecology: Nests in

exposed soil, under cover of objects, or in wood. Attends honeydew excreting insects,
can sting severely, and infests houses. Introduced. Little fire ant.

Tetramorium ? auropunctatum Roger, 1863. Berlin. Ent. Ztschr. 7:182. %, 2, d.

Wasmannia ? glabra Santschi, 1931. Rev. Ent. Sao Paulo 1:272. 3.

Hercynia panamana Enzmann, 1947. N. Y. Ent. Soc., Jour. 55:44. 8, °.

Taxonomy: Smith, 1929. Jour. Econ. Ent. 22:243. —Smith, 1936. Puerto Rico Univ., Jour. Agr.
20:831, 854. —Brown, 1948. Ent. News 59:102. —Smith, 1954. Amer. Mus. Novitates
1671:7-8. —Wheeler and Wheeler, 1954. Amer. Midland Nat. 52:444 (larva). —Kempf, 1964.
Studia Ent. 7:66.

Biology: Wheeler, 1929. Psyche 36:89-90. —Spencer, 1941. Fla. Ent. 24:6-14. —Fernald, 1947.
Jour. Econ. Ent. 40:428. —Osburn, 1948. Fla. Ent. 31:11-15. —Smith, 1965. U. S. Dept.
Agr., Tech. Bul. 1326:48-49 (economic importance).

TRIBE CEPHALOTINI
Revision: Kempf, 1951. Rev. de Ent. 22:1-244.

Taxonomy: Kempf, 1958. Studia Ent. 1:1-168. —Kempf, 1973. Studia Ent. 16: 449-462 (generic
classification; key to genera).

Genus ZACRYPTOCERUS Wheeler

Zacryptocerus Ashmead, 1905. Canad. Ent. 37: 384. Nomen nudum. No description given;
the type-species is given as Cryptocerus multistrigus F. Smith, but no such species
exists.

Zacryptocerus Wheeler, 1911. N. Y. Acad. Sci., Ann. 21: 175.

Type-species: Cryptocerus clypeatus Fabricius. Monotypic.
Cryptocerus subg. Paracryptocerus Emery, 1915. Soc. Ent. de France, Bul., p. 192.
Type-species: Cryptocerus spinosus Mayr. Orig. desig.

Cryptocerus subg. Cyathocephalus Emery, 1915. Soc. Ent. de France, Bul., p. 192. Preocc.
by Kessler, 1868.

Type-species: Cryptocerus pallens Klug. Orig. desig.

Cryptocerus subg. Hypocryptocerus Wheeler, 1920. Psyche 27: 53.

Type-species: Formica haemorrhoidalis Latreille, Orig. desig.
Cyathomyrmex Creighton, 1933. Psyche 40:98. N. name for Cyathocephalus Emery.
Paracryptocerus subg. Harnedia Smith, 1949. Psyche 56:20.

Type-species: Cryptocerus umbraculatus Fabricius. Orig. desig.

Mainly Neotropical with slight extensions into the United States. The ants are arboreal, form-
ing small colonies in cavities of plants, especially twigs. Food consists largely of honeydew and
small arthropods. The soldier is believed to keep intruders from the nest by blocking the en-
trance hole with its head.

Revision: Smith, 1947. Ent. Soc. Wash., Proc. 49:29-40. —Kempf, 1958. Studia Ent. 1:65-158.

Superfamily FORMICOIDEA 1403

Taxonomy: Kempf, 1951. Rev. de Ent. 22:156-157, 233. —Wheeler and Wheeler, 1954. Wash.
Acad. Sci., Jour. 44:156-157 (larvae). —Kempf, 1973. Studia Ent. 16: 457-460 (syn. of
Paracryptocerus and Hypocryptocerus).

rohweri (Wheeler), n. comb. S. Ariz.; Mexico. Ecology: Arboreal.
Cryptocerus (Cyathocephalus) rohweri Wheeler, 1916. New England Zool. Club, Proc. 6:32.
mele

Taxonomy: Kempf, 1958. Studia Ent. 1:129-132. —Snelling, 1968. Los Angeles Co., Mus.,
Contrib. Sci. 132:5-9.

Biology: Smith, 1947. Ent. Soc. Wash., Proc. 49:34-37. —Creighton and Nutting, 1965. Psyche
72: 59-64 (habits and distribution).
texanus (Santschi), n. comb. S. Tex.; Mexico. Ecology: Arboreal.
Cryptocerus texanus Santschi, 1915. Soc. Ent. de France, Bul., p. 208. %, 2.

Taxonomy: Kempf, 1958. Studia Ent. 1:123-127.

Biology: Smith, 1947. Ent. Soc. Wash., Proc. 49:37-40. —Creighton, 1954. Psyche 61:41-57.
—Creighton, 1963. Psyche 70:133-143.
varians (Smith), n. comb. Fla.; Central Amer., W. Indies. Ecology: Arboreal.
Cryptocerus varians Smith, 1876. London Ent. Soce., Trans., p. 606. %.

Cryptocerus (Cyathocephalus) varians var. jamaicensis Forel, 1922. Rev. Suisse de Zool.
30:97. 8.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:102-104. —Wheeler and Wheeler,
1954. Wash. Acad. Sci., Jour. 44:156 (larva). —Kempf, 1958. Studia Ent. 1:155-158.

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:11. —Smith, 1947. Ent. Soc. Wash., Proc.
49:33.

TRIBE DACETINI

Taxonomy: Brown, 1948. Amer. Ent. Soc., Trans. 74:101-129 (preliminary generic revision of
higher Dacetini). —Brown, 1949. Amer. Ent. Soc., Trans. 75:43-51. —Brown, 1953. Amer.
Midland Nat. 50:1-137 (revisionary studies). —Wheeler and Wheeler, 1954. Psyche
61:111-145 (larvae). —Brown and Wilson, 1959. Quart. Rev. Biol. 34:278-294 (evolution of
dacetine ants).

Genus STRUMIGENYS Smith

Strumigenys Smith, 1860. Jour. Ent. (London) 1:72.

Type-species: Strumigenys mandibularis Smith. Monotypic.
Labidogenys Roger, 1862. Berlin. Ent. Ztschr. 6:249.

Type-species: Labidogenys lyroessa Roger. Monotypic.
Pyramica Roger, 1862. Berlin. Ent. Ztschr. 6:251.

Type-species: Pyramica gundlachi Roger. Monotypic.
Proscopomyrmex Patrizi, 1946. Bol. Inst. Ent. R. Univ. Bologna 15:294.

Type-species: Proscopomyrmex londianensis Patrizi. Monotypic.
Eneria Donisthorpe, 1947. Ann. and Mag. Nat. Hist. 14:598.

Type-species: Eneria excisa Donisthorpe. Monotypic.

This genus is best represented in the tropical regions of the world; over 50 species are known
from the New World, but only four of these have been recorded from the United States. Three
of these four species are believed to have been introduced. Colonies are small and nests are
usually found in leaf litter, plant cavities, rotting wood, or under objects.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325-326. —Wheeler, 1908. Amer. Mus.
Nat. Hist., Bul. 24:145-147. —Smith, 1931. Ent. Soc. Amer., Ann. 24:688-691 (N. Amer.).
—Brown, 1962. Psyche 69:238-267 (Neotropical species and key).

Taxonomy: Brown, 1948. Amer. Ent. Soc., Trans. 74:108. —Brown, 1949. Mushi 20:14.
—Wheeler and Wheeler, 1954. Psyche 61:135 (larvae). —Wheeler and Wheeler, 1960. Ent.
Soc. Wash., Proc. 62:25-26 (larvae). —Brown, 1959. Studia Ent. (n. s.) 2:25-30 (silvestriz
group). —Brown, 1960 (1959). Psyche 66:37-52 (gundlachi group).

1404 Hymenoptera in America North of Mexico

Biology: Brown, 1950. Brooklyn Ent. Soc., Bul. 45:87-89. —Brown and Wilson, 1960 (1959).
Quart. Rev. Biol. 34:281.

SPECIES GROUP SILVESTRII

silvestrii Emery. La. (Paradise); Brazil, Argentina, Cuba. Ecology: In La. found in a basal rot
hole in a live oak tree. Probably introduced.
Strumigenys silvestrii Emery, 1905. Soc. Ent. Ital., Bol. 37:168. 3, °.
Strumigenys (Strumigenys) caribbea Weber, 1934. Rev. de Ent. 4:43. 3, @.

Taxonomy: Brown, 1959. Studia Ent. (n. s.) 2:25-28. —Brown, 1962. Psyche 69:246, 257, 259.
SPECIES GROUP LOUISIANAE

louisianae Roger. N. C. to Fla., w. to Okla., Ariz.; Mexico s. to Bolivia, Argentina. Ecology:

The small colonies are beneath objects, in rotting wood, plant cavities; food consists of
small arthropods, especially Collembola.

Strumigenys lowisianae Roger, 1863. Berlin. Ent. Ztschr. 7:211. 8.

Strumigenys unidentata Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:570, 575. %.

Strumigenys unispinulosa Emery, 1890. Soc. Ent. Ital., Bol. 22:67. 8, 9.

Strumigenys unispinulosa var. longicornis Emery, 1894. Soe. Ent. Ital., Bol. 26:214. 3.

Strumigenys fusca Emery, 1894. Soc. Ent. Ital., Bol. 26:215. &.

Strumigenys louisianae var. obscuriventris Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.
24:145. §.

Strumigenys bruchi Forel, 1912. Soe. Ent. de Belg., Mem. 29:197-198. 9, 3.

Strumigenys infidelis Santschi, 1919. Soc. Cient. Argentina, An. 87:48. &.

Strumigenys eggersi var. cubaensis Mann, 1920. Amer. Mus. Nat. Hist., Bul. 42:430. 8.

Strumigenys (Strumigenys) louisianae laticephala Smith, 1931. Ent. Soc. Amer., Ann.
24:688, 690. 8.

Strumigenys (Strumigenys) louisianae soledadensis Weber, 1934. Rev. de Ent. 4:38-39. 8,
&

Strumigenys (Strumigenys) louisianae guatemalensis Weber, 1934. Rev. de Ent. 4:39. 8.
Strumigenys (Strumigenys) louisianae costaricensis Weber, 1934. Rev. de Ent. 4:39. 8.
Strumigenys clasmospongia Brown, 1953. Psyche 60:2. %.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325-326. —Wheeler, 1908. Amer.
Mus. Nat. Hist., Bul. 24:147. —Haug, 1932. Ent. Soc. Amer., Ann. 25:170-172. —Brown,
1953. Amer. Midland Nat. 50:28-31. —Brown, 1953. Psyche 60:1-2. —Wheeler and Wheeler,
1954. Psyche 61:136-137 (larva). —Brown, 1957. Quart. Rev. Biol. 32:271. —Brown, 1961.
Psyche 68: 64-67, 68 (variation and syn.). —Brown, 1962. Psyche 69:246-247, 257, 263.

Biology: Smith, 1931. Ent. Soc. Amer., Ann. 24:690-691. —Creighton, 1937. Psyche 44:97-109.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31:269, 291, 305. —Wilson, 1950. Brooklyn Ent. Soc.,
Bul. 45:85-86. —Wilson, 1953. Ent. Soc. Amer., Ann. 46:481-483. —Kempf, 1958. Studia Ent.
1:555.

SPECIES GROUP GUNDLACHI

eggersi Emery. Fla. (Miami, Fisher’s Is., Archbold Biol. Sta.); W. Indies, Mexico, S. Amer.
Probably introduced.
Strumigenys eggersi Emery, 1890. Soc. Ent. Ital., Bol. 22:68. 3, 2.

Taxonomy: Brown, 1960 (1959). Psyche 66:46-47 (worker, female). —Brown, 1962. Psyche
69:249, 257, 264.

Biology: Weber, 1952. Amer. Mus. Novitates 1554:5. —Kempf, 1958. Studia Ent. 1:554.
—Kempf, 1961. Studia Ent. 4:515.
gundlachi (Roger). Fla. (Royal Palm Ranger Sta., Everglades Natl. Pk.; Key Largo); W.
Indies, Mexico, Central Amer. Ecology: Usually found in leaf litter; feeds almost
exclusively on Collembola. Probably introduced.
Pyramica gundlachi Roger, 1862. Berlin. Ent. Ztschr. 6:253. “2”= &.
Strumigenys eggersi var. vincentensis Forel, 1893. London Ent. Soc., Trans. p. 378. &.

Superfamily FORMICOIDEA 1405

Strumigenys eggersi var. banillensis Santschi, 1930. Soe. Roy. Ent. d’Egypte, Bul. (n. s.), p.
80. 8.

Strumigenys bierigi Santschi, 1930. Soc. Roy. Ent. d’Egypte, Bul. (n. s.), p. 80. 3.

Strumigenys (Strumigenys) eggersi var. isthmica Santschi, 1931. Rev. de Ent. 1:276. %.

Strumigenys (Strumigenys) eggersi infuscata Weber, 1934. Rev. de Ent. 4:35. 3, 2.

Strumigenys (Strumigenys) eggersi var. berlesei Weber, 1934. Rev. de Ent. 4:36. %.

Taxonomy: Brown, 1960 (1959). Psyche 66:37-45 (worker, female). —Brown, 1962. Psyche
69:249, 257, 264.

Biology: Weber, 1952. Amer. Mus. Novitates 1554:4-5.

Genus SMITHISTRUMA Brown

Cephaloxys Smith, 1864. Linn. Soe. London, Jour. 8:77. Preoce. by Signoret, 1847.
Type-species: Cephaloxys capitata Smith. Monotypic.

Smithistruma Brown, 1948. Amer. Ent. Soc., Trans. 74:104-106.
Type-species: Strumigenys pulchella Emery. Orig. desig.

Smithistruma subg. Wessonistruma Brown, 1948. Amer. Ent. Soc., Trans. 74:106.
Type-species: Strumigenys pergandei Emery. Orig. desig.

Smithistruma subg. Weberistruma Brown, 1948. Amer. Ent. Soc., Trans. 74:106-107.
Type-species: Strumigenys (Cephaloxys) leptothrix Wheeler. Orig. desig.

Smithistruma subg. Platystrwma Brown, 1953. Amer. Midland Nat. 50:112.
Type-species: Strumigenys (Cephaloxys) depressiceps Wheeler. Orig. desig.

Ants of this genus nest in small colonies in the soil, various types of soil cover, rotting logs and

stumps, and beneath objects. They are most frequently collected in berlese funnels of samples of
leaf litter, duff, ete. Food consists almost exclusively of Collembola.

Revision: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325-329. —Wheeler, 1908. Amer. Mus.
Nat. Hist., Bul. 24:148. —Smith, 1931. Ent. Soc. Amer., Ann. 24:691-710. —Wesson and
Wesson, 1939. Psyche 46:91-112. —Brown, 1953. Amer. Midland Nat. 50:55-92 (key to spp.).

Taxonomy: Wheeler and Wheeler, 1954. Psyche 61:141-145 (larvae).

Biology: Brown, 1950. Brooklyn Ent. Soc., Bul. 45:87-89. —Wilson, 1953. Ent. Soc. Amer.,
Ann. 46:485-487. —Brown and Wilson, 1960 (1959). Quart. Rev. Biol. 34:282. —Brown, 1964.
Amer. Ent. Soc., Trans. 89:183-200.

abdita (Wesson and Wesson). Pa., Va., N. C., Ohio, Ill, Iowa. Ecology: Nests are found under
rocks in the soil.
Strumigenys (Cephaloxys) abdita Wesson and Wesson, 1939. Psyche 46:95, 109. %.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:89-90 (worker, female). —Brown, 1964.
Amer. Ent. Soc., Trans. 89:189-190, 199-200.

angulata (Smith). Ill., Ala., Miss. Ecology: Colonies found in a rotten pine stump; some
specimens found in nest of Strumigenys louisianae Roger.
Strumigenys (Cephaloxys) angulata Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 697. &.

Taxonomy: Smith, 1932. Ent. News 43:157-158 (also biology). —Wesson and Wesson, 1939.
Psyche 46:110. —Brown, 1953. Amer. Midland Nat. 50:54-55 (worker, female).
bimarginata (Wesson and Wesson). Ohio, II]. Ecology: Specimens found in soil cover.
Strumigenys (Cephaloxys) bimarginata Wesson and Wesson, 1939. Psyche 46:95-97. &.
Taxonomy: Brown, 1953. Amer. Midland Nat. 50:62-63 (worker, female).
bunki Brown. Ga., Fla., Miss., La. Ecology: Nests are found in the soil.
Smithistruma (Smithistruma) bunki Brown, 1950. Amer. Ent. Soe., Trans. 76:41-42. &.
Taxonomy: Brown, 1953. Amer. Midland Nat. 50:82-84 (worker, female). —Brown, 1964.
Amer. Ent. Soc., Trans. 89:190.

californica Brown. Calif.
Smithistruma (Smithistruma) californica Brown, 1950. Amer. Ent. Soc., Trans. 76:40-41.

2
Taxonomy: Brown, 1953. Amer. Midland Nat. 50:85-86.

1406 Hymenoptera in America North of Mexico

carolinensis Brown. N. C., S. C. Ecology: Specimens found in pine needle litter and oak leaf
mold.
Smithistruma carolinensis Brown, 1964. Amer. Ent. Soc., Trans. 89:185-186. &.

cloydi Pfitzer. Tenn. (Knoxville).
Smithistruma cloydi Pfitzer, 1951. Tenn. Acad. Sci., Jour. 16:198-200. 9.
Taxonomy: Brown, 1964. Amer. Ent. Soc., Trans. 89:191.
clypeata (Roger). N. J., Pa. s. to Fla., w. to Ill., Ark., La. Ecology: Usually found in forested
areas where it nests in soil cover, under stones, and in rotten logs.
Strumigenys clipeata(!) Roger, 1863. Berlin. Ent. Ztschr. 7:213. %.

Taxonomy: Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:571 (worker, female). —Emery,
1895. Zool. Jahrb., Abt. f. System. 8:326, 328 (each caste). —Wheeler, 1908. Amer. Mus.
Nat. Hist., Bul. 24:148. —Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 699-700. —Brown,
1953. Amer. Midland Nat. 50:58-59 (each caste). —Wheeler and Wheeler, 1954. Psyche
61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:191 (further distribution).

Biology: Smith, 1932. Ent. News 43:159. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:291, 305.
—Wesson and Wesson, 1939. Psyche 46:93-94. —Smith, 1955. Brooklyn Ent. Soc., Bul.
50:28. —Wilson, 1953. Ent. Soc. Amer., Ann. 46:486.

creightoni (Smith). D. C., N. C., Ga., Fla., Tenn., Ala. Ecology: Collections made from leaf
litter.
Strumigenys (Cephaloxys) creightoni Smith, 1931. Ent. Soc. Amer., Ann. 24:692, 705. &.

Taxonomy: Wesson and Wesson, 1939. Psyche 46:110. —Brown, 1953. Amer. Midland Nat.
50:80-81 (worker, female). —Brown, 1964. Amer. Ent. Soc., Trans. 89:191.

dietrichi (Smith). Md. s. to Fla., w. to Ohio, Ill, Ark., La. Ecology: Nests in or under bark of
rotting logs or stumps; a few are known from leaf litter. Sometimes occurs in or near
nests of other ants.
Strumigenys (Cephaloxys) dietrichi Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 696. 9.

Taxonomy: Smith, 1932. Ent. News 43:159. —Wesson and Wesson, 1939. Psyche 46:108.

—Brown, 1953. Amer. Midland Nat. 50:67-68 (worker, female). —Wheeler and Wheeler,
1954. Psyche 61:1438 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:191-192.

Biology: Kennedy and Schramm, 1933. Ent. Soc. Amer., Ann. 26:95-96. —Dennis, 1938. Ent.
Soc. Amer., Ann. 31:270, 272-273, 291, 305. —Wilson, 1953. Ent. Soc. Amer., Ann. 46:487.

filirrhina Brown. N. C., Mo. Ecology: Specimens taken in leaf mold.
Smithistruma (Smithistruma) filirrhina Brown, 1950. Amer. Ent. Soc., Trans. 76:37-38. &%.
Taxonomy: Brown, 1953. Amer. Midland Nat. 50:63-64. —Brown, 1964. Amer. Ent. Soc.,
Trans. 89:193.
filitalpa Brown. Ind., Ark.
Smithistruma (Smithistruma) filitalpa Brown, 1950. Amer. Ent. Soc., Trans. 76:39-40. %.
Taxonomy: Brown, 1953. Amer. Midland Nat. 50:79-80. —Brown, 1964. Amer. Ent. Soc.,
Trans. 89:193-194.
laevinasis (Smith). Va., N. C., Tenn., Ala., Miss., Ill. Ecology: Nests have been found in rotten

logs.
Strumigenys (Cephaloxys) clypeata var. laevinasis Smith, 1931. Ent. Soc. Amer., Ann.

24:691, 701. 8.

Taxonomy: Wesson and Wesson, 1939. Psyche 46:109. —Brown, 1953. Amer. Midland Nat.
50:61-62 (worker, female). —Brown, 1964. Amer. Ent. Soc., Trans. 89:194, 199.

Biology: Smith, 1932. Ent. News 43:159.
margaritae (Forel). Ga., Ala., Tex.; W. Indies, Mexico, Colombia. Ecology: Specimens have
been found in soil cover.
Strumigenys margaritae Forel, 1893. London Ent. Soc., Trans, p. 378. 8, 2, d.
Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:148. —Smith, 1931. Ent. Soc.
Amer., Ann. 24:691, 692-693. —Wesson and Wesson, 1939. Psyche 46:108. —Brown, 1953.
Amer. Midland Nat. 50:108-110. —Brown, 1964. Amer. Ent. Soc., Trans. 89:194-195.

Superfamily FORMICOIDEA 1407

missouriensis (Smith). N. Y., Va., N. C., Ohio, Miss., Il., lowa, Mo. Ecology: Primarily a soil
and soil cover inhabitant.
Strumigenys (Cephaloxys) missouriensis Smith, 1931. Ent. Soc. Amer., Ann. 24:691, 701.

om
Strumigenys (Cephaloxys) sculpturata Smith, 1931. Ent. Soc. Amer., Ann. 24:692, 706. %.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:72-73 (worker, female). —Wheeler and
Wheeler, 1954. Psyche 61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:195.

Biology: Wilson, 1953. Ent. Soc. Amer., Ann. 46:485-486.
ohioensis (Kennedy and Schramm). N. J. s. to Ga., w. to Ill, Ark., La. Ecology: Found in soil
cover and upper soil layers, often under objects; sometimes in rotting wood.
Strumigenys ohioensis Kennedy and Schramm, 1933. Ent. Soc. Amer., Ann. 26:98-99. %.
Strumigenys (Cephaloxys) manni Wesson and Wesson, 1939. Psyche 46:97. 3.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:87-89 (each caste). —Brown, 1964. Amer.
Ent. Soc., Trans. 89:196.
ornata (Mayr). Del., Md. s. to Fla., w. to Mich., Ill., Mo., Okla., Tex. Ecology: Most commonly
found in leaf litter or forest debris.
Strumigenys ornata Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:571. &.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325, 328. —Wheeler, 1908. Amer.
Mus. Nat. Hist., Bul. 24:148 (worker, female). —Smith, 1931. Ent. Soe. Amer., Ann. 24:691,
695-696. —Wesson and Wesson, 1939. Psyche 46:109. —Brown, 1953. Amer. Midland Nat.
50:65-67. —Brown, 1964. Amer. Ent. Soc., Trans. 89:197.

pergandei (Emery). Mass., N. Y., Ont. s. to N. C., Tenn., w. to Iowa, Mo., Kans. Ecology: Nests
in rotten logs, in soil, or under stones in soil. Often found in or near nests of other ants
where it may prey on myrmecophilous collembolans. Records from Mass. are the
northernmost for dacetine ants in the New World.
Strumigenys pergandei Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325. %, 2, d.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:148. —Smith, 1931. Ent. Soc.
Amer., Ann. 24:691, 698-699 (also biology). —Wesson and Wesson, 1939. Psyche 46:110.
—Brown, 1953. Amer. Midland Nat. 50:51-54. —Wheeler and Wheeler, 1954. Psyche
61:144-145 (larva). —Brown, 1964. Amer. Ent. Soc, Trans. 89:197.

Biology: Kennedy and Schramm, 1933. Ent. Soc. Amer., Ann. 26:95-98. —Wesson, 1936. Ent.
News 47:171-174. —Brown, 1952. Psyche 59:12. —Wilson, 1953. Ent. Soc. Amer., Ann.
46:487-488.

pilinasis (Forel). Pa. s. to N. C., Ala., w. to Ohio., Ill, Mo., Ark., La. Ecology: Nests under
stones, in soil cover, or in logs.
Strumigenys clypeata var. pilinasis Forel, 1901. Soc. Ent. de Belg., Ann. 45:339. &.
Strumigenys (Cephaloxys) clypeata var. brevisetosa Smith, 1935. Ent. Soc. Amer., Ann.
28:215. %.

Strumigenys (Cephaloxys) medialis Wesson and Wesson, 1939. Psyche 46:94, 110. &.
Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:148 (worker, female). —Smith,
1931. Ent. Soc. Amer., Ann. 24:691, 700. —Wesson and Wesson, 1939. Psyche 46:109.
—Brown, 1953. Amer. Midland Nat. 50:60-61 (worker, female). —Wheeler and Wheeler,

1954. Psyche 61:143 (larva). —Brown, 1964. Amer. Ent. Soc., Trans. 89:197-199.

Biology: Wilson, 1953. Ent. Soc. Amer., Ann. 46:485.
pulchella (Emery). N. Y., Pa. s. to Fla, w. to Ill, La. Ecology: Nests in rotting wood.

Strumigenys pulchella Emery, 1895. Zool. Jahrb., Abt. f. System. 8:325, 327. 9.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:148 (worker, female). —Smith,
1931. Ent. Soc. Amer., Ann. 24:691, 702. —Brown, 1953. Amer. Midland Nat. 50:70-72 (each
caste). —Brown, 1964. Amer. Ent. Soc., Trans. 89:198.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:291, 305. —Wesson and Wesson, 1939.
Psyche 46:111.

1408 Hymenoptera in America North of Mexico

reflexa (Wesson and Wesson). Md. s. to N. C., Tenn., Ala., w. to Ohio, Ill. Ecology: Nests in soil
or. under or in objects lying on soil.
Strumigenys (Cephaloxys) refleca Wesson and Wesson, 1939. Psyche 46:102, 111. &.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:73-75 (each caste).

rohweri (Smith). Miss.
Strumigenys (Cephaloxys) rohwert Smith, 1935.-Ent. Soc. Amer., Ann. 28:214. 8.

Taxonomy: Wesson and Wesson, 1939. Psyche 46:109. —Brown, 1953. Amer. Midland Nat.
50:57-58.
rostrata (Emery). N. J., Pa. s. to Fla., w. to Ill, Mo., La. Ecology: Nests in soil and rotting
wood.
Strumigenys rostrata Emery, 1895. Zool. Jahrb., Abt. f. System. 8:326, 329. 3, 2, o.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:148. —Smith, 1931. Ent. Soe.
Amer., Ann. 24:692, 704-705. —Wesson and Wesson, 1939. Psyche 46:110. —Brown, 1953.
Amer. Midland Nat. 50:84-85 (each caste). —Wheeler and Wheeler, 1954. Psyche 61:144
(larva).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31:273, 291-292, 305. —Wilson, 1953. Ent. Soc.
Amer., Ann. 46:485.
talpa (Weber). N. C., Fla., Ohio, Tenn., Ala., Ill., La. Ecology: Found in soil or humus.
Strumigenys (Cephaloxys) talpa Weber, 1934. Psyche 41:63. %.
Strumigenys (Cephaloxys) venatrix Wesson and Wesson, 1939. Psyche 46:108, 110. 8.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:76-78 (each caste). —Wheeler and Wheeler,
1954. Psyche 61:141 (larva).

Biology: Wilson, 1953. Ent. Soc. Amer., Ann. 46:486.

wrayi Brown. N. C. (Fayetteville). Ecology: Found in leaf mold.
Smithistruma (Smithistruma) wrayi Brown, 1950. Amer. Ent. Soc., Trans. 76:38-39. &.

Taxonomy: Brown, 1953. Amer. Midland Nat. 50:78-79.

Genus TRICHOSCAPA Emery

Strumigenys subg. Trichoscapa Emery, 1869. Accad. Degli Aspiranti Napoli, Ann. 2:24.
Type-species: Strumigenys (Trichoscapa) membranifera Emery. Monotypic.

The single species in this genus has been spread throughout the tropical regions of the world
by commerce.

Revision: Brown, 1948. Amer. Ent. Soc., Trans. 74:112-114.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:144-145, 148. —Smith, 1931. Ent.
Soc. Amer., Ann. 24:691, 693-695.

Biology: Brown and Wilson, 1960 (1959). Quart. Rev. Biol. 34:283.
membranifera (Emery). Ga., Fla., Ala., Miss., La., Calif. (nr. Stockton); Pantropical. Ecology:

Nests are in soil, wood, and plant cavities, and the ants are predaceous on a variety of
small soft-bodied arthropods. Introduced, probably of African origin.

Strumigenys (Trichoscapa) membranifera Emery, 1869. Accad. Degli Aspiranti Napoli,
Ann. 2:24. 3.

Strumigenys membranifera simillima Emery, 1890. Soc. Ent. Ital., Bol. 22:69. 8.

Strumigenys membranifera var. santschii Forel, 1904. Rev. Suisse de Zool. 12:6. 3.

Strumigenys (Cephaloxys) vitiensis Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bul.
64:461-462. &.

Strumigenys (Cephaloxys) silvestriana Wheeler, 1928. Bol. Lab. Zool. Sci. Agr. Portici
22:27-28. 9.

oe. (Cephaloxys) foochowensis Wheeler, 1928. Bol. Lab. Zool. Sci. Agr. Portici
22:28-29.

ae (Cephatozys) membranifera var. marioni Wheeler, 1933. Hawaii. Ent. Soc.,
Proc. 8:276.

Superfamily FORMICOIDEA 1409

Strumigenys (Cephaloxys) membranifera var. williamsi Wheeler, 1933. Hawaii. Ent. Soc.,
Proc. 8:275-277. %, °@.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:829, 856. —Brown, 1948. Amer.
Ent. Soc., Trans. 74:114. —Brown, 1949. Mushi 20:7, 22. —Wheeler and Wheeler, 1954.
Psyche 61:145 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:35 (Polynesia).

Biology: Weber, 1952. Amer. Mus. Novitates 1554:7. —Wilson, 1953. Ent. Soc. Amer., Ann.
46:483-485. —Brown, 1954. Harvard Univ., Mus. Comp. Zool., Bul. 112:6.

Genus QUADRISTRUMA Brown

Quadristruma Brown, 1949. Amer. Ent. Soc., Trans. 75:47.
Type-species: E'pitritus emmae Emery. Orig. desig.
The single species in the United States is an introduction.

emmae (Emery). Fla. (homestead); Pantropical. Ecology: Most commonly found under objects

on sea beaches or in debris from forests or agricultural areas near the sea. Introduced;
spread to many tropical regions of the world by commerce, possibly originating from
Africa.

Eiptritus emmae Emery, 1890. Soc. Ent. Ital., Bol. 22:70. 8.

Epitritus clypeatus Szabo, 1909. Arch. Zool. (Budapest) 1(7):1. %.

Epitritus clypeatus var. malesiana Forel, 1913. Zool. Jahrb., Abt. f. System. 36:83-84. %,
g

Epitritus wheeleri Donisthorpe, 1916. Ent. Rec. 28:121. 8.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:149. —Smith, 1936. Puerto Rico
Univ., Jour. Agr. 20:829, 858. —Brown, 1949. Amer. Ent. Soc., Trans. 75:48-50 (worker,
female). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14:42 (Polynesia).

Biology: Weber, 1934. Rev. de Ent. 4:51. —Brown, 1949. Mushi 20:21. —Brown, 1954.
Harvard Univ., Mus. Comp. Zool., Bul. 112:6. —Brown and Wilson, 1960 (1959). Quart. Rev.
Biol. 34:281. —Kempf, 1961. Studia Ent. 4:516.

TRIBE BASICEROTINI

Revision: Brown and Kempf, 1960. Studia Ent. 3:161-250 (world). —Taylor, 1968. Austral.
Jour. Zool. 16:333-348 (Indo-Australian).

Taxonomy: Wheeler and Wheeler, 1954. Psyche 61:111-145 (larvae).

Genus EURHOPALOTHRIX Brown and Kempf

Eurhopalothrix Brown and Kempf, 1960. Studia Ent. (n. s.) 3:202-203.
Type-species: Rhopalothrix Bolaui Mayr. Desig. by Brown and Kempf, 1961.

Taxonomy: Brown and Kempf, 1961 (1960). Psyche 67:44. —Wheeler and Wheeler, 1973.
Pan-Pacifie Ent. 49:207 (larvae).
floridana Brown and Kempf. Fla. (Highlands Hammock, Highlands Co.). Ecology: Collected in
leaf litter.
Eurhopalothrix floridana Brown and Kempf, 1960. Studia Ent. (n. s.) 3:207-208. 8.

TRIBE ATTINI

Taxonomy: Wheeler, 1948. Amer. Midland Nat. 40:664-689 (larvae). —Weber, 1970. Ent. Soc.
Wash., Proc. 72:414-415 (northern extent of tribe). —Wheeler and Wheeler, 1974. Ent. Soc.
Wash., Proc. 76:76-81 (larvae).

Biology: Weber, 1962. Amer. Mus. Nat. Hist., Bul. 71:45-49 (insect gardeners). —Weber,
1966. Science 153:587-604 (general). —Moser, 1967. Amer. Mus. Nat. Hist., Nat. Hist.
76:32-35 (trail of the leafcutters). —Weber, 1972. Amer. Scientist 60:448-456 (general).
—Weber, 1972. Amer. Phil. Soc., Mem. 92, 146 pp. (gardening ants, the attines).

1410 Hymenoptera in America North of Mexico

Morphology: Blum, Moser, and Cordero, 1964. Psyche 71:1-7 (source and specificity of odor
trail substances). —Moser, 1964. Science 143:1048-1049 (inquiline roach responds to trail
making substance). —Markl, 1970. Z. Vergl. Physiol. 69:2-37 (communication by
stridulatory signals). —Crewe and Blum, 1972. Jour. Ins. Physiol. 18:31-42 (alarm
pheromones, their phylogenetic significance).

Genus CYPHOMYRMEX Mayr

Cyphomyrmex Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12:690.
Type-species: Cyphomyrmex minutus Mayr. Monotypic.

Cyphomyrmex subg. Cyphomannia Weber, 1938. Rev. de Ent. 9:183.
Type-species: Cyphomyrmex (Cyphomannia) laevigatus Weber. Orig. desig.

A rather large Neotropical genus with extensions into the southern United States. These ants
form small colonies, usually in the soil, and live on fungi which they cultivate from a substratum
of vegetable matter of feces of certain insects.

Revision: Mayr, 1887. Zool.-Bot. Gesell. Wien, Verh. 37:555-562. —Wheeler, 1907. Amer. Mus.
Nat. Hist., Bul. 23:670, 719-728, 765-773. —Weber, 1940. Rev. de Ent. 11:406-427. —Kempf,
1964. Studia Ent. 7:1-44 (strigatus group). —Kempf, 1965. Studia Ent. 8:161-200 (rimosus
group).

Taxonomy: Wheeler, 1948. Amer. Midland Nat. 40:668-670 (larvae). —Weber, 1966. Ent.
News 77:166-168.

Biology: Weber, 1955. Science 121:109. —Weber, 1957. Ecology 38: 480-494 (cultivation of

fungus).

rimosus (Spinola). Fla., Ala., Miss., La., Tex., Ariz., Calif.; W. Indies and Mexico s. to

Argentina. Ecology: Nests are in soil, under bark of rotten wood, or in humus around
roots; cultures a fungus in form of a yeast. A puzzling and variable complex for which
many subspecies have been proposed; see Kempf (1965) for names to be considered.

Cryptocerus ? rimosus Spinola, 1853. Accad. Sci. Torino, Mem. (2) 13: 65. 8%, 3.

Meranoplus difformis Smith, 1858. Cat. Hym. Brit. Mus. v. 6, p. 195. %.

Cyphomyrmex minutus Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12:691. 8.

Cyphomyrmesx steinheili Forel, 1884. Soc. Vaud. des Sci. Nat., Bul. 20:368. 9.

Cyphomyrmex rimosus var. comalensis Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:719.
CROCE

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:719-712. —Weber, 1940. Rev. de
Ent. 11:411. —Weber, 1941. Rev. de Ent. 12:116-119. —Wheeler, 1948. Amer. Midland Nat.
40:668 (larva). —Weber, 1958. Ent. Soc. Wash., Proc. 60:259-260. —Wheeler and Wheeler,
1960. Ent. Soc. Wash., Proc. 62:28 (larva). —Kempf, 1965. studia Ent. 8:162, 198.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:106. —Weber, 1945. Rev. de Ent.
16:5-14. —Weber, 1947. Bol. de Ent. Venezolana 6:144. —Weber, 1955. Wash. Acad. Sci.,
Jour. 45:275-281. —Weber, 1972. Amer. Phil. Soc., Mem. 92, pp. 8, 26, 31-32, 34, 39-42, 51,
60, 71-72, 94-95, 98-99, 102, 105-108, 110, 117.

wheeleri Forel. Tex., Calif.; Mexico. Ecology: Nests in the soil beneath stones in very arid
habitats.
Cyphomyrmex wheeleri Forel, 1900. Schweiz. Ent. Gesell. Mitt. 10:282. 3, °.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:725-726 (each caste). —Weber,
1940. Rev. de Ent. 11:409. —Kempf, 1965. Studia Ent. 8:167-172.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40:74. —Weber, 1972. Amer. Phil. Soc.,
Mem. 92, pp. 26, 93. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 97-98.

Genus MYCETOSORITIS Wheeler

Atta subg. Mycetosoritis Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:714.
Type-species: Atta (Mycetosoritis) hartmanni Wheeler. Monotypic

A small genus found only in the United States and Brazil.

Superfamily FORMICOIDEA 1411

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104:317-318. —Kempf,
1964. Studia Ent. 7:1.

hartmanni (Wheeler). La., Tex. Ecology: Forms small colonies in soil; food consists of a fungus
grown upon a substratum composed of flower anthers.
Atta (Mycetosoritis) hartmanni Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:714. %, 2,
3.

Taxonomy: Wheeler and Wheeler, 1974. Ent. Soc. Wash., Proc. 76:77-79 (larva).

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:761-765. —Wheeler, 1910. Ants, pp.
334-335.

Genus TRACHYMYRMEX Forel

Atta subg. Trachymyrmex Forel, 1893. Soc. Ent. de Belg., Ann. 37:600.
Type-species: Atta septentrionalis McCook. Desig. by Wheeler, 1911.

A Neartcic and Neotropical genus found from New York south to Argentina, and the largest
genus of attine ants. In the United States, most forms are recorded from the more humid areas,
though a few occur in arid sections. Nests are in the soil. The fungi on which the ants feed is
grown on a substratum of plant particles and insect excrement.

Revision: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:706-714, 746-760. —Wheeler, 1911. N.
Y. Ent. Soc., Jour. 19:245-255.

Taxonomy: Weber, 1945. Rev. de Ent. 16:44. —Wheeler, 1948. Amer. Midland Nat. 40:673-674
(larvae).

Biology: Weber, 1955. Science 121:109. —Weber, 1960. Ent. News 71:1-6 (comparison of
worker and female behavior).
arizonensis (Wheeler). Ariz. (Huachuca Mtns.). Ecology: Found in arid, stony canyons, 5000 to
6000 ft.
Atta (Trachymyrmex) arizonensis Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:710. 9, 3.

Taxonomy: Wheeler, 1911. N. Y. Ent. Soc., Jour. 19:250 (worker).

Biology: Wheeler, 1911. Psyche 18:93-98. —Essig, 1926. Ins. of West. No. Amer., p. 862.
—Weber, 1972. Amer. Phil. Soc., Mem. 92, p. 93.
desertorum (Wheeler). Ariz. (nr. Tucson). Ecology: Nests were found in hard, pebbly, desert
soil.
Atta (Trachymyrmex) desertorum Wheeler, 1911. Psyche 18:98-100. &.
Taxonomy: Wheeler, 1911. N. Y. Ent. Soc., Jour. 19:249.

Biology: Wheeler, 1911. Psyche 18:100-101. —Essig, 1926. Ins. of West. No. Amer., p. 862.
jamaicensis (Andre). Fla. (Dania); W. Indies. Possibly introduced.
Atta (Acromyrmex) jamaicensis Andre, 1893. Rev. Ent. Caen 12:149. 8.
Trachymyrmex sharpi Forel, 1893. Ent. Soc. London, Trans., p. 372. %.
Atta (Trachymyrmex) maritima Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:107. &.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20:860. —Weber, 1967. Ent. News
78:107-109.

Biology: Weber, 1972. Amer. Phil. Soc., Mem. 92, pp. 102, 104 (fungi).
nogalensis Byars. Ariz. (Nogales).

Trachymyrmex nogalensis Byars, 1951. Ent. Soc. Wash., Proc. 53:109-111. 3, @.
septentrionalis (McCook). N. Y. (L. I., Staten Is.) s. to Fla., w. to Ill., La., Tex. Ecology: Nests
are in soil, usually with a small semicircular crater. Of some economic importance

because of the habit of cutting leaves from domesticated plants. The New York records
are the northernmost for an attine ant.

Oecodoma virginiana Buckley, 1867. Ent. Soc. Phila., Proc. 6:346. ¥. Placement of this
species is questionable.

Atta septentrionalis McCook, 1880. Acad. Nat. Sci. Phila., Proc. 32:359. %.

Atta (Trachymyrmex) septentrionalis var. obscurior Wheeler, 1907. Amer. Mus. Nat. Hist.,
Bul. 23:706. %, 2, 3

1412 Hymenoptera in America North of Mexico

Atta (Trachymyrmex) septentrionalis var. vertebrata Wheeler, 1911. N. Y. Ent. Soc., Jour.
19:246, 250. 8, 2.

Atta (Trachymyrmex) septentrionalis obscurior var. seminole Wheeler, 1911. N. Y. Ent.
Soce., Jour. 19:247, 250. 3, 9, do.

Atta (Trachymyrmex) septentrionalis obscurior var. crystallina Wheeler, 1911. N. Y. Ent.
Soe., Jour. 19:247, 250. %, °.

Atta (Trachymyrmex) septentrionalis obscurior var. irrorata Wheeler, 1911. N. Y. Ent.
Soc., Jour. 19:247, 250. &.

Taxonomy: Forel, 1884. Soc. Vaud. des Sci. Nat., Bul. 20:91. —Smith, 1924. Ent. News 35:52.
—Cole, 1940. Amer. Midland Nat. 24:59. —Wheeler, 1948. Amer. Midland Nat. 40:673
(larva). —Weber, 1958. Ent. News 69:52-53.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21:374, 386-387. —Wheeler, 1907. Amer.
Mus. Nat. Hist., Bul. 23:680-681, 746-753. —Wheeler, 1911. Psyche 18:95-97. —Wheeler,
1911. N. Y. Ent. Soc., Jour. 19:250-255. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:273, 292,
305. —Cole, 1939. Lloydia 2:153-160. —Cole, 1951 (1950). Ent. Soc. Amer., Ann. 43:499-500.
—Weber, 1956. Ecology 37:150-161. —Weber, 1966. Ent. News 77:241. —Moser, 1964.
Science 143:1048-1049 (inquiline roach responds to trail-making substance). —Weber, 1972.
Amer. Phil. Soc., Mem. 92, pp. 3, 13-15, 25-28, 32-33, 35-36, 40-44, 46, 51, 54-55, 62, 70, 99,
103-104, 106-107, 112, 117.

smithi neomexicanus Cole. N. Mex., Ariz. Ecology: Nests found in sand. T. smithi smithi
Buren occurs in Mexico.
Trachymyrmex smithi neomexicanus Cole, 1952. Tenn. Acad. Sci., Jour. 27:159-162. §.

Taxonomy: Cole, 1953. Tenn. Acad. Sci., Jour. 28:300-301 (female, male).
turrifex caroli (Wheeler). Tex. (Huntsville).
Atta (Trachymyrmex) turrifex caroli Wheeler, 1911. N. Y. Ent. Soc., Jour. 19:248. 9.
turrifex turrifex (Wheeler). La., Tex.; Mexico. Ecology: Nests in soil.
Atta (Trachymyrmex) turrifex Wheeler, 1903. Psyche 10:100. %, 2.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat., Hist., Bul. 23:709-710. —Wheeler, 1911. N. Y.
Ent. Soc., Jour. 19:249.

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:753-759. —Wheeler, 1911. Psyche
18:95-97.

Genus ACROMYRMEX Mayr

Atta subg. Acromyrmex Mayr, 1865. Reise d. Novara, Zool. v. 1, Hym., p. 83.
Type-species: Formica hystrix Latreille. Monotypic.
Atta subg. Moellerius Forel, 1893. Soc. Ent. de Belg., Ann. 37:589.
Type-species: Atta (Acromyrmex) landolti Forel. Desig. by Wheeler, 1911.
Pseudoatta Gallardo, 1916. Buenos Aires Mus. Nac. de Hist. Nat., An. 28:320. Syn.
uncertain.
Type-species: Pseudoatta argentina Gallardo. Monotypic.

A Neotropical genus with a slight extension into southwestern United States. These are
mostly grassland and semidesert fungus-growing ants which usually collect grass. They are com-
parable to Atta in biology and economic importance but colonies are apparently not so large and
the leaf-cutting habits not so extensive. The species listed below have previously been placed in
the subgenus Moellerius.

Revision: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:669-700, 703-706, 743-746. —Santschi,
1925. Rev. Suisse de Zool. 31:386-398.

Taxonomy: Weber, 1945. Rev. de Ent. 16:61-62. —Wheeler, 1948. Amer. Midland Nat.
40:674-676 (larvae). —Brown, 1973. In Meggers, et al., Tropical Forest Ecosystems in Afr.
and S. Amer., pp. 178-185 (generic syn.).

Biology: Weber, 1967. Ent. Soc. Amer., Ann. 78:107-109 (growth of young colonies in first
year).

Superfamily FORMICOIDEA 1413

versicolor chisosensis (Wheeler). Tex. (Mtns. of Big Bend area).
Atta (Moellerius) versicolor chisosensis Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:705.
3.
versicolor versicolor (Pergande). Ariz., Calif.; Mexico. Ecology: Nests are in soil in deserts;
collects pieces of leaves from bushes and herbs for a media on which to grow fungus.
Atta versicolor Pergande, 1893. Calif. Acad. Sci., Proc. 4:31. &.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:703-705 (each caste).

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:743-746. —Wheeler, 1911. Psyche
18:101. —Wheeler, 1917. Psyche 24:179-180. —Weber, 1972. Amer. Phil. Soc., Mem. 92, pp.
22, 26, 30, 32.

Genus ATTA Fabricius

Atta Fabricius, 1804. Systema Piezatorum, p. 421.
Type-species: Formica cephalotes Linnaeus. Desig. by Wheeler, 1911.
Oecodoma Latreille, 1818. Nouv. Dict. Hist. Nat. 23:223.
Type-species: Formica cephalotes Linnaeus. Desig. by Shuckard, 1840.
Archeatta Goncalves, 1942. Soc. Brasil. de Agron., Bol. 5:342.
Type-species: Oecodoma mexicana Smith. Orig. desig.
Atta subg. Neoatta Goncalves, 1942. Soc. Brasil. de Agron., Bol. 5:334.
Type-species: Formica sexdens Linnaeus. Orig. desig.
Atta subg. Palaeatta Borgmeier, 1950. Inst. Oswaldo Cruz, Mem. 48:244, 270.
Type-species: Atta bisphaerica Forel. Orig. desig.
Atta subg. E'piatta Borgmeier, 1950. Inst. Oswaldo Cruz, Mem. 48:246, 272.
Type-species: Atta laevigata Smith. Orig. desig.

A Neotropical genus with an extension into southern United States. The nests, which are deep
in the soil and usually have many lateral and vertical entrances, are often extremely large and
contain numerous individuals. Workers defoliate both wild and domesticated plants, including
trees, and they feed on a fungus grown from a substratum composed of macerated leaves and
other vegetable matter.

Revision: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:669-703, 729-742. —Goncalves, 1942.
Soc. Brasil. de Agron., Bol. 5:333-358. —Borgmeier, 1959. Studia Ent. 2:321-390.

Taxonomy: Wheeler, 1948. Amer. Midland Nat. 40:676-683 (larvae). —Borgmeier, 1950. Inst.
Oswaldo Cruz, Mem. 48:239-263. —Weber, 1958. Ent. News 69:7-13. —Smith, 1963. Ent.
Soe. Wash., Proc. 65:299-302 (key to spp. of U. S. and Mexico; biol. notes). —Weber, 1968.
Ent. Soc. Wash., Proc. 70:348-350 (Panama species).

Biology: Weber, 1962. Amer. Mus. Nat. Hist., Nat. Hist. 61:45-51.

Morphology: Blum, et al., 1968. Comp. Biochem. Physiol. 26: 291-299 (alkanones and terpenes
in the mandibular glands of Atta species).
mexicana (Smith). Ariz. (Organ Pipe Cactus Natl. Mon.); Mexico s. to Honduras. Ecology:
Nests are in a variety of ecological situations and their fungus gardens are grown on a
wide variety of vegetable substances.
Oecodoma mexicana Smith, 1858. Cat. Hym. Brit. Mus. 6:185. 2, d.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:702-703. —Goncalves, 1942. Soc.
Brasil. de Agron., Bol. 5:335, 337, 343. —Byars, 1949. Jour. Econ. Ent. 42:545 (Ariz. record;
soldier). —Borgmeier, 1950. Inst. Oswaldo Cruz, Mem. 48:260-261. —Borgmeier, 1959.
Studia Ent. 2:351-353 (each caste). —Smith, 1963. Ent. Soc. Wash., Proc. 65:299-302.

texana (Buckley). La., Tex.; e. Mexico. Ecology: Nests are in well-drained sand or loamy soils.
The interior of the nest may reach a depth of 15 to 20 ft. and contains innumerable
interconnected chambers. It is connected to the exterior by numerous holes which are
surrounded by crater-shaped piles of dirt. In a large nest there may be 1,000 entrance
holes occupying an area of 4,500 square feet. The ants grow a fungus primarily on
macerated leaves. An economically important pest; invades houses; cuts leaves from
domesticated plants; steals seeds; builds unsightly nests; damages roads, walks, stock, or

1414 Hymenoptera in America North of Mexico

equipment by cave-ins of the nest; and can inflict painful bites. Texas leafcutting ant;
also known locally as the town ant, night ant, cut ant, parasol ant, pack ant, or fungus
ant.

Myrmica (Atta) texana Buckley, 1860. Acad. Nat. Sci. Phila., Proc. 12:233. ¥, 2, d.

Taxonomy: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:700-703. —Goncalves, 1942. Soc.
Brasil. de Agron., Bol. 5:335, 337, 343. —Byars, 1949. Jour. Econ. Ent. 42:545. —Borgmeier,
1950. Inst. Oswaldo Cruz, Mem. 48:260-261. —Borgmeier, 1959. Studia Ent. 2:356-358.
—Smith, 1963. Ent. Soc. Wash., Proc. 65:299-301. —Wheeler and Wheeler, 1974. Ent. Soc.
Wash., Proc. 76:79-81 (larva).

Biology: Wheeler, 1907. Amer. Mus. Nat. Hist., Bul. 23:729-742. —Hunter, 1912. U. S. Dept.
Agr., Bur. Ent. Cir. 148:1-4. —Jones, 1917. Jour. Econ. Ent. 10:561. —Snyder, 1937. La.
Conserv. Rev., pp. 14-17. —Walter, Seaton, and Mathewson, 1938. U. S. Dept. Agr. Cir.
494:1-18. —Metcalf and Flint, 1939. Destructive and Useful Insects, p. 771. —Smith, 1939.
South. Forest Expt. Sta., Occas. Papers 84:2-6. —Johnston, 1944. Jour. Forestry 42:130-132.
—Bennett, 1958. U. S. Dept. Agr., Forest Pest Leaflet 23:1-4 (rev. 1967). —Warter, Moser,
and Blum, 1962. La. Acad. Sci., Proc. 25:42-46 (correlation between night hawk feeding and
nuptial flights). —Moser, 1962. Forests and People 12:40-41 (nesting habits). —Moser,
1963. Ent. Soc. Amer., Ann. 56:286-291 (contents and structure of nest in summer).
—Moser, 1964. Science 143:1048-1049 (inquiline roach responds to trail-making substance).
—Moser, 1967. Insectes Sociaux 14:295-312 (mating activities). —Moser, 1967. Amer. Mus.
Nat. Hist., Nat. Hist. 76:33-35. —Weber, 1972. Amer. Phil. Soc., Mem. 92, pp. 3, 24, 30-33,
35, 37-38, 65, 81, 84.

Morphology: Moser and Blum, 1963. Science 140:1228 (trail marking substance, source and
potency). —Moser, Brownlee, and Silverstein, 1968. Jour. Ins. Physiol. 14:529-535 (alarm
pheromones). —Hermann, Moser, and Hunt, 1970. Ent. Soc. Amer., Ann. 63:1152-1158
(poison apparatus, morphological and behavioral changes). —Moser and Silverstein, 1967.
Nature 215:206-207 (volatility of trail marking substance). —Tumlinson, e¢ al., 1971.
Nature 234: 348-349 (identification of trail pheromone). —Tumlinson, et al., 1972. Jour.
Insect. Physiol. 18: 809-814 (a volatile trail pheromone). —Riley, et al., 1974. Science 183:
760-762 (biological responses to its alarm pheromone).

UNPLACED TAXA OF MYRMICINAE

Pheidole buckleyi Smith, 1951. In Muesebeck, et al, U. S. Dept. Agr., Agr. Monog. 2:806. N.
name for Atta pennsylvanica Buckley.

Crematogaster (Acrocoelia) sanguinea coachellai Enzmann, 1946. N. Y. Ent. Soc., Jour. 54:95.
%. In same paper, fig. 3 of the petiole is given as that of Crematogaster (Acrocoelia)
lineolata var. coachellai.

Myrmica corrugata Say, 1836. Boston Jour. Nat. Hist. 1:291. 2, d.

Myrmica dimidiata Say, 1836. Boston Jour. Nat. Hist. 1:293. 9 (?). N. Amer.

Myrmica (Monomarium (!)) diversa Buckley, 1867. Ent. Soc. Phila., Proc. 6:337. 8, 2. Tex.

Myrmica inflecta Say, 1836. Boston Jour. Nat. Hist. 1:292. 3.

Myrmica (Monomarium (!)) lineolata Buckley, 1867. Ent. Soc. Phila., Proc. 6:340. §, 2. Preoce.
by Say, 1836. No locality.

Solenopsis madara Roger, 1863. Berlin. Ent. Ztschr. 7:200. 9, 9.

Myrmica (Monomarium (!)) montana Buckley, 1867. Ent. Soc. Phila., Proc. 6:339. %. Tex.

Myrmica novaeboracensis Buckley, 1867. Ent. Soc. Phila., Proc. 6:337. 2. N. Y.

Myrmica opposita Say, 1836. Boston Jour. Nat. Hist. 1:292. 3, d. N. Amer.

Atta pennsylvanica Buckley, 1867. Ent. Soc. Phila., Proc. 6:345. %, 2{. Pa. Previously placed in
Pheidole where it would be preoce. by Roger, 1863; see Pheidole buckleyi Smith.

Atta picea Buckley, 1867. Ent. Soc. Phila., Proc. 6:344. 3, 2{. Tex.

Oecodoma pilosa Buckley, 1867. Ent. Soc. Phila., Proc. 6:348. 3. Tex.

Myrmica (Diplorhoptrum) scabrata Buckley, 1867. Ent. Soc. Phila., Proc. 6:343. %. Conn.

Myrmica (Atta) sublanuginosa Buckley, 1867. Ent. Soc. Phila., Proc. 6:343. %. Tex.

Oecodoma tardigrada Buckley, 1867. Ent. Soc. Phila., Proc. 6:349. 8, 2, d. Tex.

Pheidole pubiventris race timmi Forel, 1901. Mitt. Naturh. Mus. Hamburg 18:62. %, 2.
“Probably of Mexican origin - Dr. R. Timm.”

Superfamily FORMICOIDEA 1415
SUBFAMILY DOLICHODERINAE

A small subfamily with most Nearctic representatives found in the southern portion of the
United States. Most species are small, drab-colored insects and are recognized by the single seg-
mented pedicel, lack of a constriction between the first and second gastric segments, and the
slitlike cloacal orifice.

Taxonomy: Starcke, 1933. Tijdschr. v. Ent. 76: XX VI-XXXII (larvae). —Wheeler and
Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 169-210 (larvae). —Wheeler and Wheeler, 1966.
Ent. Soc. Amer., Ann. 59: 726-732 (larvae). —Wheeler and Wheeler, 1974 (1973).
Pan-Pacifie Ent. 49: 396-401 (larvae).

Morphology: Eisner, 1957. Harvard Univ., Mus. Comp. Zool., Bul. 116: 453-464
(proventriculus; use as a generic character).

TRIBE DOLICHODERINI

Genus DOLICHODERUS Lund

Brown (1973) listed Monacis Roger, 1862, Diabolus Karavaiev, 1926, Diceratoclinea Wheeler,
1935, and Karawajewella Donisthorpe, 1944 as synonyms of Dolichoderus and Hypoclinea Mayr,
1855, and Acanthoclinea Wheeler, 1935 as provisional synonyms. Because of the doubt as to the
exact status of some of the genera, subgeneric groupings are used as in some previous North
American literature until the genus has been adequately studied.

Taxonomy: Brown, 1973. In Meggers, et al., Tropical Forest Ecosystems in Afr. and S.
Amer., pp. 178-185 (generic syn.).

Genus DOLICHODERUS Subgenus DOLICHODERUS Lund

Dolichoderus Lund, 1831. Ann. des Sci. Nat., Zool. 23: 130.
Type-species: Formica attelaboides Fabricius. Monotypic.

Not known to occur in the Nearctic Region.

Genus DOLICHODERUS Subgenus HYPOCLINEA Mayr

Hypoclinea Mayr, 1855. Zool.-Bot. Gesell. Wien, Verh. 5: 377.
Type-species: Formica quadripunctata Linnaeus. Desig. by Wheeler, 1911.

In North America, this subgenus is found only in the eastern half of the continent. Colonies
are small to moderately large and nests are constructed in the soil, in curled leaves and hollow
stems of plants, and in cartons attached to plants. Workers attend honeydew-excreting insects
and feed on small arthropods. Some forms emit a fluid which has a peculiar smoky or pungent
odor.

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 953-960. —Mayr, 1886. Zool.-Bot.
Gesell. Wien, Verh. 36: 434-437. —Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 305-319.

Taxonomy: Brown, 1950. Wasmann Jour. Biol. 8: 249. —Wheeler and Wheeler, 1951. Ent. Soc.
Wash., Proc. 53: 172 (larvae). —Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59:
726-730 (larvae).

Biology: Kannowski, 1959. Insectes Sociaux 6: 129-134 (flight comparisons of some spp.).
—Torossian, 1960. Insectes Sociaux 7: 383-393 (biology of D. quadripunctatus (L.)).

Morphology: Torossian, 1959. Insectes Sociaux 6: 369-374 (tropholactic and proctodeal
exchange in D. quadripunctatus (L.)).
mariae Forel. Mass. to Ga., w. to Minn., Ill., Okla., La. Ecology: Nests of the rather large

colonies are found in the soil, preferably sand, beneath tufts of grass or small bushes.

Dolichoderus Mariae Forel, 1884. Soc. Vaud. des Sci. Nat., Bul. 20: 349. &.

Dolichoderus mariae davisi Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306, 308. &.

Dolichoderus (Hypoclinea) mariae var. blatchleyi Wheeler, 1917. Ind. Acad. Sci., Proc. 26:
462. 8.

1416 Hymenoptera in America North of Mexico

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bul. 22: 589 (worker, female). —Cole, 1940. Amer. Midland
Nat. 24: 60. —Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 726 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 316-319, 387-388. —Smith, 1924. Ent.
News 35: 81. —Talbot, 1956. Psyche 63: 134-139 (flight activities). —Kannowski, 1959.
Insectes Sociaux 6: 129-133.

plagiatus (Mayr). N. B., Ont. s. to Ga., Tenn., w. to N. Dak., Ill. Ecology: Colonies are small and
nests are found in inconspicuous places such as under forest debris in the soil, in hollow
stems, and in curled-up leaves.
Hypoclinea plagiata Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 957, 960. &.
Dolichoderus borealis Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 408. &.
Dolichoderus plagiatus var. inornatus Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306,
313. &.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 310 (each caste). —Wheeler, 1917.
Conn. State Geol. and Nat. Hist. Survey Bul. 22: 590. —Cole, 1940. Amer. Midland Nat. 24:
60. —Gregg, 1944. Ent. Soc. Amer., Ann. 37: 467. —Wheeler, and Wheeler, 1966. Ent. Soc.
Amer., Ann. 59: 727 (larva). —Francoeur and Beique, 1966. Canad. Ent. 98: 142
(Provancher types).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306, 315-316, 388. —Wheeler, 1908.
Amer. Mus. Nat. Hist., Bul. 24: 621-622. —Wesson and Wesson, 1940. Amer. Midland Nat.
24: 90, 99. —Kannowski, 1959. Insectes Sociaux 6: 133. —Wheeler and Wheeler, 1963. Ants
of N. Dak., pp. 149-151. —Kannowski, 1967. Ent. Soc. Amer., Ann. 60: 1246-1252
(population studies).

pustulatus Mayr. N.S. s. to Fla., w. to IIl., Okla., Tex. Ecology: Colonies are small and nests
are found under piles of detritus or in a hard, thin carton shell above ground about
blades of a tuft of grass.
Dolichoderus pustulatus Mayr, 1886. Zool.-Bot. Gesell. Wien., Verh. 36: 435-436. %, 9.
Dolichoderus plagiatus var. beutenmuelleri Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20:
304. 3.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306, 313, 388. —Smith, 1924. Ent.
News 35: 82. —Smith, 1931. Ent. News 42: 22. —Wheeler and Wheeler, 1966. Ent. Soc.
Amer., Ann. 59: 727-728 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 315-316, 388. —Wheeler, 1917. Ind.
Acad. Sci., Proc. 26: 462. —Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 12. —Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 89, 99. —Kannowski, 1959. Insectes Sociaux 6:
133-134, 155. —Kannowski, 1967. Ent. Soc. Amer., Ann. 60: 1246-1252 (population studies).

taschenbergi (Mayr). N. S., Ont. s. to S. C., w. to Man., N. Dak., S. Dak., La. Ecology: Nests of

rather large colonies are constructed in the soil in wooded areas or the edge of woods
and usually have a low mound of thatch (grass, twigs, needles) over the entrance hole.
Workers commonly attend honeydew-excreting insects.

Hypoclinea Taschenbergi Mayr, 1866. Akad. der Wiss. Wien, Math.-Nat. Kl. Sitzber. 53:
498. 3.

Dolichoderus taschenbergi var. gagates Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306,
310. %. Preoce. by Emery, 1890.

Dolichoderus (Hypoclinea) taschenbergi var. aterrimus Wheeler, 1915. Amer. Mus. Nat.
Hist., Bul. 34: 417. N. name for gagates Wheeler.

Dolichoderus tachenbergi var. wheeleriella Forel, 1916. Rev. Suisse de Zool. 24: 458. N.
name for gagates Wheeler.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 306, 309-310. —Smith, 1924. Ent.
News 35: 82. —Cole, 1940. Amer. Midland Nat. 24: 60. —Wheeler and Wheeler, 1966. Ent.
Soc. Amer., Ann. 59: 728 (larva).

Biology: Wheeler, 1915. Psyche 22: 306. —Smith, 1924. Ent. News 35: 81-82. —Logier, 1923.
Canad. Ent. 55: 247. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 292-293. —Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 99. —Kannowski, 1959. Ent. Soc. Amer., Ann. 52:

Superfamily FORMICOIDEA 1417

755-760 (flight activities). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 151-154.
—Bradley and Hinks, 1968. Canad. Ent. 100: 40-50 (attending aphids on jack pine).
—Bradley, 1972. Canad. Ent. 104: 245-249 (transplanting colonies in jack pine stands in
Man.). —Burns, 1973. Canad. Ent. 105: 97-104 (foraging and tending behavior).

TRIBE TAPINOMINI

Genus LIOMETOPUM Mayr

Liometopum Mayr, 1861. Die Europaischen Formiciden, pp. 25, 38.
Type-species: Formica microcephala Panzer. Monotypic.

A Holarctic genus found only in the western United States in the Nearctic region. The Neare-
tic forms nest in the soil beneath cover or under bark or in crevices in trees and the nest cham-
bers are usually subdivided by a network of paperlike material. Colonies are often populous.
Workers forage in files sometimes several hundred feet long and commonly attend coccids and
aphids. They are pugnacious and eject a repellent secretion with a strong and disagreeable odor
like that of butyric acid.

Revision: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 321-333.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 172, 181 (larvae).
apiculatum Mayr. W. Tex., Colo., N. Mex., Ariz.; Mexico. Ecology: Usually in foothill areas at
elevations of 4000 to 7000 feet.
Liometopum apiculatum Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 961. 9.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 331 (female). —Wheeler, 1905.
Amer. Mus. Nat. Hist., Bul. 21: 322-324 (each caste). —Wheeler and Wheeler, 1951. Ent.
Soe. Wash., Proc. 53: 181 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 322-324. —Wheeler, 1917. Psyche 24:
177-178. —Gregg, 1963. Univ. Colo. Studies, Biol. Ser. 11, pp. 1-6. —Gregg, 1963. Ants of
Colo., pp. 488-441. —Van Pelt, 1971. Ent. Soc. Amer., Ann. 64: 1186 (trophobiosis and
feeding habits).

occidentale luctuosum Wheeler. Wyo., Colo., w. Tex., w. to Ariz., Calif. Ecology: At higher
elevations, 4000 to 8000 feet.
Liometopum apiculatum luctuwosum Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 325. %.

Taxonomy: Forel, 1914. Deut. Ent. Ztschr. 6: 619 (male). —Cole, 1942. Amer. Midland Nat.
28: 371.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 332. —Mallis, 1941. South Calif. Acad.
Sci., Bul. 40: 75. —Gregg, 1963. Ants of Colo., pp. 441-443. —Cole, 1966. Brigham Young
Univ., Sci. Bul., Biol. Ser. 7: 18 (Nevada Test Site). —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 100-101.

occidentale occidentale Emery. Oreg., Calif.; Mexico. Ecology: Found at lower elevations, sea
~ level to 4000 feet.
Liometopum microcephalum var. occidentale Emery, 1895. Zool. Jahrb., Abt. f. System. 8:
330. 8, 2.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 324-325. —Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 8, 17.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 522. —Mallis, 1941. South Calif.
Acad. Sci., Bul. 40: 75. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 100.

Genus IRIDOMYRMEX Mayr

Iridomyrmex Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 702.
Type-species: Formica detecta Smith. Desig. by Bingham, 1903.

Tapinoma subg. Doleromyrma Forel, 1907. Mus. Nat. Hungarici, Ann. 5: 28.
Type-species: Tapinoma (Doleromyrma) darwinianum Forel. Monotypic.
Anonychomyrma Donisthorpe, 1946. Ann. and Mag. Nat. Hist. 13: 588. Syn. uncertain.

Type-species: Anonychomyrma myrmex Donisthorpe. Monotypic.

1418 Hymenoptera in America North of Mexico

Three of the four species of this cosmopolitan genus found in North America have been in-
troduced. Most species nest in the soil and some are found in buildings. Workers collect
honeydew but are also entomophagous.

Revision: Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 953-960. —Smith, 1929. Jour. Econ.
Ent. 22: 241-243.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 172, 184 (larvae).
—Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 729 (larvae).
glaber (Mayr). Fla. (Winter Park); Australian and Oriental regions. Ecology: Florida specimens
collected from a stump of queen palm, Arecastrum romanzoffianum. Probably
introduced. These specimens are tentatively referred to glaber since the taxonomy of
glaber and related species in the Old World are unstudied.
Hypoclinea glabra Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 705. %, 3.

Taxonomy: Brown, 1958. Acta Hym. 1:40-41 (also biological notes; New Zealand; Australia).
—Wheeler and Wheeler, 1974 (1973). Pan-Pacifie Ent. 49: 398 (larva, from Australian
specimens).

humilis (Mayr). Md. to Fla. w. to IIl., Tex., Ariz., Calif.; Mexico, S. Amer.; Europe; S. Africa;
Hawaii; Australia. Ecology: Nests in soil, rotting wood or debris and lives in large
colonies that contain many females. Workers forage in files and tend
honeydew-excreting insects though they also feed on other sweet substances. A most
persistent and troublesome house-infesting ant; also known to steal seeds from
seedbeds, drive poultry from their nests, kill newly hatched chickens, foster
honeydew-excreting insects, disrupt bee hives, and gnaw into ripened fruits. Introduced,
native to S. Amer. and spread to the U. S. and other parts of the world by commerce;
probably introduced first into New Orleans on coffee ships from Brazil sometime prior
to 1891. Argentine ant.

Hypoclinea (Iridomyrmex) humilis Mayr, 1868. Soc. Nat. Modena, Ann. 3: 164. 2.

Taxonomy: Wheeler, 1913. In Newell and Barber, U. S. Dept. Agr. Bur. Ent. Bul. 122: 27-30
(each caste). —Smith, 1929. Jour. Econ. Ent. 22: 241. —Wheeler and Wheeler, 1951. Ent.
Soc. Wash., Proc. 53: 186 (larva). —Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59:
729 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 79-80 (Polynesia).

Biology: Newell, 1909. Jour. Econ. Ent. 2: 174-192 (life history). —Newell and Barber, 1913.
U.S. Dept. Agr. Bur. Ent., Bul. 122: 1-98. —Gallardo, 1915. Buenos Aires Mus. Nac. de
Hist. Nat., An. 27: 23-25. —Barber, 1920. U. S. Dept. Agr. Farmers’ Bul. 1101: 1-11.
—Woglum and Borden, 1921. U. S. Dept. Agr. Bul. 965: 1-43. —Harned and Smith, 1922.
Jour. Econ. Ent. 15: 261-264. —Essig, 1926. Ins. of West. N. Amer., pp. 865-866. —Smith,
1936. U. S. Dept. Agr. Cir. 387: 1-39. —Mallis, 1942. Sci. Monthly 55: 536-545 (half a
century with the Argentine ant). —Flanders, 1943. Calif. Citrograph 28: 117, 128, 137
(Argentine ant versus parasites of black scale). —McCluskey, 1958. Science 128: 536-537
(daily rhythm in male Argentine ant). —Skaife, 1962. The Study of Ants, p. 12. —Pavan,
1963. Pavia Univ., Sympos. Genet. et Biol. Ital. 12: 122-131. —Haskins and Haskins, 1965.
Ecology 46: 737 (extension of range in Bermuda at expense of Pheidole megacephala (F.)).
—Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326, pp. 52-54 (economic importance).
—Markin, 1968. Kans. Ent. Soc., Jour. 41: 511-516 (study in citrus groves in Calif.).
—Markin and McCoy, 1968. Ent. Soc. Amer., Ann. 61: 505-509 (a nematode in pharyngeal
glands). —Markin, 1969. Insectes Sociaux 16: 159-190 (food distribution in laboratory
colonies). —Dechene, 1970. Wasmann Jour. Biol. 28: 175-184 (behavioral patterns).
—Fluker and Beardsley, 1970. Ent. Soc. Amer., Ann. 63: 1290-1296 (ecological notes, in
Hawaii). —Markin, 1970. Jour. Econ. Ent. 63: 740-744 (foraging behavior in Calif. citrus
groves). —Markin, 1970. Insectes Sociaux 17: 127-158. —Markin, 1970. Ent. Soc. Amer.,
Ann. 63: 1238-1242 (seasonal life cycle in S. Calif.). —Erickson, 1971. Psyche 78: 257-266
(displacement of native ant spp. by Argentine ant).

Morphology: Pavan, 1950 (1948). 8th Internat. Cong. Ent., Proc., Stockholm, pp. 863-865
(physiology). —Pavan, 1955. Soc. Ital. di Sci. Nat. Atti 94: 379-477. —Weber, 1961. Ent.
Soc. Wash., Proc. 63: 218 (use of poison): —Torossian, 1961. Insectes Sociaux 8: 189-191.
—Nachtwey, 1963. Insectes Sociaux 10: 50-53.

Superfamily FORMICOIDEA 1419

iniquus nigellus Emery. Mass., IIl., other scattered localities mostly in northeastern U. S.;
Central Amer. Ecology: Found in greenhouses in U. S. Introduced. /. iniquus iniquus
Mayr occurs in Central and S. Amer. and W. Indies.
Tridomyrmex iniquus var. nigellus Emery, 1890. Soc. Ent. Ital., Bol. 22: 56. 8.
Biology: Wheeler, 1929. Psyche 36: 89-90. —Smith, 1929. Jour. Econ. Ent. 22: 241-242.
pruinosus analis (Andre). N. Dak., Kans., Okla., Tex. w. to s. Idaho, Calif.; Mexico, Guatemala.
Ecology: Nests are in soil either under objects or in exposed situations surmounted by a

craterlike mound.
Tapinoma anale Andre, 1893. Rev. Ent. de France 12: 148. §.
Iridomyrmex pruinosus var. testaceus Cole, 1936. Ent. News 47: 121. %, @.

Taxonomy: Smith, 1929. Jour. Econ. Ent. 22: 241. —Eckert and Mallis, 1937. Calif. Agr. Expt.
Sta. Cir. 342: 8. —Cole, 1942. Amer. Midland Nat. 28: 372-373 (also biological notes).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 342. —Smith, 1924. Ent. News 35: 83.

—Dennis, 1938. Ent. Soc. Amer., Ann. 31: 294, 305. —Wheeler and Wheeler, 1963. Ants of

N. Dak., pp. 154-155. —Gregg, 1963. Ants of Colo., pp. 4836-438. —Cole, 1966. Brigham

Young Univ., Sci. Bul., Biol. Ser. 7: 18 (Nevada Test Site). —Wheeler and Wheeler, 1973.

Ants of Deep Canyon, pp. 101-102.

pruinosus pruinosus (Roger). N. Y. to Fla., w. to Wis., Ohio, Tex., N. Mex.; W. Indies.

Ecology: Nests are in exposed soil with craterlike mounds or under the cover of objects.
A house pest in the Gulf Coast states.

Tapinoma pruinosum Roger, 1863. Berlin Ent. Ztschr. 7: 165. §.

Taxonomy: Smith, 1929. Jour. Econ. Ent. 22: 241. —Cole, 1940. Amer. Midland Nat. 24: 64-65.
—Cole, 1942. Amer. Midland Nat. 28: 372. —Santschi, 1930. Soc. Roy. Ent. d’Egypte, Bul.
23: 81. —Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 185. (larva). —Wheeler
and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 729 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 389. —Smith, 1930. Fla. Ent. 14: 5.
—Wheeler, 1911. Harvard Univ., Mus. Comp. Zool., Bul. 54: 497. —Warter, Moser, and
Blum, 1962. La. Acad. Sci., Proc. 25: 42-46 (correlation between night hawk feeding and
nuptial flights of pruinosus pruinosus). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:
54-55 (economic importance).

Morphology: Blum, Warter, Monroe, and Chidchester, 1963. Jour. Ins. Physiol. 9: 881-885
(chemical releasers of social behavior - methyl-n-amy] ketone).

Genus FORELIUS Emery

Forelius Emery, 1888. Ztschr. f. Wiss. Zool. 46: 389.
Type-species: Iridomyrmex mccooki Forel. Monotypic.
A small New World genus.
Taxonomy: Wheeler and Wheeler, 1966. Ent. Soc. Amer., Ann. 59: 729 (larvae).

foetidus (Buckley). Ark., Kans., Okla., Tex., Colo. w. to Calif.; Mexico. Ecology: Nests are
under objects or in exposed soil with small craters of soil. Incorrectly reported from D.

C.
Formica foetida Buckley, 1866. Ent. Soc. Phila., Proc. 6: 167. 3, 2.

Iridomyrmex McCooki Forel, 1878. Soc. Vaud. des Sci. Nat., Bul. 15: 382.
Forelius MacCooki race Andrei Forel, 1912. Soc. Ent. de Belg., Mem. 20: 44. &.
Taxonomy: Forel, 1886. Soc. Ent. de Belg., Bul. (C. R.) 30: 39. —Wheeler and Wheeler, 1974
(1973). Pan-Pacifiec Ent. 49: 401 (larva).
Biology: Cole, 1937. Ent. News 48: 137. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 48: 137.
—Lindquist, 1942. Jour. Econ. Ent. 35: 850-852. —Gregg, 1963. Ants of Colo., pp. 434-436.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 102-103.

Genus CONOMYRMA Forel

Conomyrma Forel, 1913. Rev. Zool. Africaine 2: 350.
Type-species: Prenolepis pyramica Roger. Desig. by Santschi, 1922.

1420 Hymenoptera in America North of Mexico

Conomyrma subg. Biconomyrma Kusnezov, 1952. Acta Zool. Lilloana (Tucuman) 10:
429-430.

Type-species: Dorymyrex pyramicus var. brunnea Forel. Desig. by Kusnezov, 1959.

A New World genus. The Nearctic forms construct nests in the soil which are surmounted by

irregular or craterlike mounds of soil. They are usually found in open sunny areas in situations

where other ant species find conditions intolerable. The workers are predaceous, active and

agressive and emit a fluid with a disagreeable odor. The species below have previously been as-

signed to the genus Dorymyrmex Mayr, a genus now restricted to South America. The taxa of
this genus occurring in the Gulf coast states are not yet clearly understood.

Taxonomy: Gallardo, 1916. Buenos Aires Mus. Nac. de Hist. Nat., An. 28: 1-130. —Santschi,
1922. Soe. Vaud. des Sci. Nat., Bul. 54: 365. —Kusnezov, 1952. Acta Zool. Lilloana
(Tucuman) 10: 427, 429, 433-488. —Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53:
183. —Kusnezov, 1959. Zool. Anz. 162: 38-51. —Snelling, 1973. Los Angeles Co. Mus.,
Contrib. Sci. 238: 1-6 (in the U. S.).

bicolor (Wheeler). Tex., N. Mex., Utah, Ariz., Nev., Calif.; Mexico. Ecology: Nests are in
exposed situations in the soil and are usually surmounted by irregular or craterlike
mounds.
Dorymyrmex pyramicus var. bicolor Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 342.
Sao

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 372. —Cole, 1957. N. Y. Ent. Soc., Jour. 65:

130. —Wilson, 1957. Psyche 64: 76.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 76. —Eckert and Mallis, 1937. Calif.
Agr. Expt. Sta. Cir. 342: 19. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 18
(Nevada Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 103-104.

flavopecta (Smith). Fla. Occurrence in other Gulf Coast states is not certain.
Dorymyrmex pyramicus flavopectus Smith, 1944. Fla. Ent. 27: 15. %.
insana (Buckley). N. C. to Fla., w. to Ill, N. Dak., Oreg., Calif.; Mexico; W. Indies. Ecology:
Nests are in soil in open sunny places and are surmounted by an irregular or craterlike
mound. A pest species mainly in the South where it commonly enters houses and builds
ugly mounds on lawns. Pyramid ant. Records in the literature for Dorymyrmex
pyramicus (Roger) in N. Amer. should refer to this species; Conomyrma pyramica
(Roger) is a valid species but is found only in S. Amer.
Formica insana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 165. %, °.
Dorymyrmex flavus McCook, 1879. In Comstock, Rpt. on Cotton Insects, p. 186. &.
Dorymyrmex pyramicus var. nigra Pergande, 1895. Calif. Acad. Sci., Proc. 5: 871. &.
Dorymyrmex pyramica brunnea var. antillana Forel, 1911. Deut. Ent. Ztschr., p. 306.
Dorymyrmex pyramicus var. smithi Cole, 1936. Ent. News 47: 120. 8.
Conomyrma (?Biconomyrma) wheeleri Kusnezov, 1952. Acta Zool. Lilloana (Tucuman) 10:
438. 8.

Taxonomy: Emery, 1895. Zool. Jahrb., Abt. f. System 8: 331-332 (male). —Cole, 1940. Amer.
Midland Nat. 24: 61. —Cole, 1942. Amer. Midland Nat. 28: 371. —Smith, 1936. Puerto Rico
Univ., Jour. Agr. 20: 861, 864. —Buren, 1944. Iowa State Col., Jour. Sci. 18: 291. —Wheeler
and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 183 (larva). —Wilson, 1967. Psyche 64: 76
(sympatry of bicolor and pyramica). —Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci.
238: 5 (syn.).

Biology: Wheeler, 1910. Ants, pp. 146, 210, 205, 426. —Smith, 1924. Ent. News 35: 82.
—Dennis, 1938. Ent. Soc. Amer., Ann. 31: 293, 305. —Eckert and Mallis, 1937. Calif. Agr.
Expt. Sta. Cir. 342: 19. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 155-157.
—Gregg, 1963. Ants of Colo., pp. 432-434. —Wilson, 1967. Psyche 64: 76. —Smith, 1965. U.
S. Dept. Agr., Tech. Bul. 1326: 56-57 (economic importance). —Cole, 1966. Brigham Young
Univ., Sci. Bul., Biol. Ser. 7: 18 (Nevada Test Site). —Wheeler and Wheeler, 1973. Ants of
Deep Canyon, pp. 105-106.

Morphology: Metcalf and Metcalf, 1970. Ent. Soc. Amer., Ann. 63: 34-35 (effects of isomeres

of 2-heptanone on alarm behavior). —Blum and Warter, 1966. Ent. Soc. Amer., Ann. 59:
774-779 (alarm and digging behavior; isolation of 2-heptanone).

Superfamily FORMICOIDEA ‘1421

Genus TAPINOMA Foerster

Tapinoma Foerster, 1850. Hym. Stud. 1: 43.
Type-species: Tapinoma collina Foerster. Monotypic.
Micromyrma Dufour, 1857. Soc. Ent. de France, Ann. 5: 60.
Type-species: Micromyrma pygmaea Dufour. Monotypic.
Tapinoma subg. Tapinoptera Santschi, 1925. Eos 1: 348.
Type-species: Tapinoma vexatum Santschi. Monotypic.

At least one of the three North American species is introduced. Nests are found in a variety
of situations and workers commonly attend honeydew-excreting insects. All of the species emit a
substance with an odor similar to that of butyric acid.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 172 (larvae).

Morphology: Torossian, 1960. Insectes Sociaux 7: 171-174 (physiology of the Palearctic
Tapinoma erraticum (Latreille)). —Weber, 1961. Ent. Soc. Wash., Proc. 63: 217-218 (use of
poison).

litorale Wheeler. S. Fla.; W. Indies. Ecology: Apparently arboreal, nesting in twigs of trees
and bushes, hollow culms, or between leaves.
Tapinoma litorale Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 109. %, 2, 6.

Taxonomy: Smith, 1936. Puerto Rico Univ., Jour. Agr. 20: 861-862.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 150. —Wheeler, 1932. N. Y. Ent. Soc.,
Jour. 40: 13.

melanocephalum (Fabricius). Fla., where it is established; occasionally found in greenhouses

or heated buildings in other parts of the continent such as Ont., Iowa., N. Mex., Calif.;
tropicopolitan. Ecology: Highly adaptable in nesting habits; found in soil, rotten wood,
decayed parts of trees, under bark, in plant cavities, houses, and greenhouses. Workers
are fond of honeydew but also feed on live and dead insects. A house-infesting ant.
Introduced; widely distributed in the tropical regions of the world by commerce.

Formica melanocephala Fabricius, 1793. Ent. System. 2: 353. &.

Formica nana Jerdon, 1851. Madras Jour. Lit. Sci. 17: 125. 8.

Myrmica pellucida Smith, 1857. Linn. Soc. London, Jour., Zool. 2: 71. %.

Formica familiaris Smith, 1860. Linn. Soc. London, Jour., Zool., Sup. 4: 96. 8.

Tapinoma (Micromyrma) melanocephalum var. australis Santschi, 1928. Insects of Samoa
5: 53. 8.

Taxonomy: Smith, 1928. Ent. Soc. Amer., Ann. 21: 311. —Smith, 1936. Puerto Rico Univ.,
Jour. Agr. 20: 861-862. —Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 197
(larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 79-82 (Polynesia).

Biology: Wheeler, 1910. Ants, pp. 154, 156. —Marlatt, 1928. U. S. Dept. Agr. Farmers’ Bul.
740: 6. —Phillips, 1934. Hawaii Univ., Expt. Sta. Pineapple Prod. Coop. Assoc., Bul. 15:
20-21. —King, 1948. Iowa Acad. Sci., Proc. 55: 395. —Kempf, 1961. Studia Ent. 4: 520.
—Brown, 1964. Ent. News 75: 14-15. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326:
59-60 (economic importance). —Tamsitt and Fox, 1966. Ent. Soc. Wash., Proc. 68: 268
(attacking lab. flea colony). —Gomez-Nunez, 1971. Jour. Med. Ent. 8: 735-737 (an inhibitor
of Rhodinus prolixus populations).

Morphology: Weber, 1961. Ent. Soc. Wash., Proc. 63: 218 (employment of venom).
sessile (Say). N. S., Que. s. to Fla., w. to Wash., Calif.; Mexico. Ecology: A very adaptable ant

found in a wide variety of habitats. Most nests are in the soil beneath objects but also
under bark, in stumps, plant cavities, insect galls, refuse piles, and bird and mammal
nests. Colonies may contain thousands of individuals and numerous reproductive
females. A common and troublesome house-infesting ant. Food consists mainly of
honeydew and workers attend honeydew-excreting insects. Odorous house ant.

Formica sessilis Say, 1836. Boston Jour. Nat. Hist. 1: 287. 3, 2.

Tapinoma boreale Roger, 1863. Berlin. Ent. Ztschr. 7: 165. 8, °.

Formica gracilis Buckley, 1866. Ent. Soc. Phila., Proc. 6: 158. 8, °.

Formica parva Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. &.

1422 Hymenoptera in America North of Mexico

Tapinoma boreale Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 238. 9, ¢.
Preocc. by Roger, 1863.

Taxonomy: Wheeler and Wheeler, 1951. Ent. Soc. Wash., Proc. 53: 196 (larva). —Francoeur
and Beique, 1966. Canad. Ent. 98: 143 (Provancher types).

Biology: Smith, 1928. Ent. Soc. Amer., Ann. 21: 307-329. —Metcalf and Flint, 1939.
Destructive and Useful Insects, Ed. 2: 770. —Weber, 1941. Canad. Ent. 73: 140-141.
—Kannowski, 1959. Insectes Sociaux 6: 126-129. —Wheeler and Wheeler, 1963. Ants of N.
Dak., pp. 157-160. —Gregg, 1963. Ants of Colo., pp. 443-447. —Judd, 1964. Canad. Ent. 96:
990 (in galls on goldenrod). —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 13826: 57-59
(economic importance). —Bobb, 1965. Jour. Econ. Ent. 58: 925 (as predator of Neodiprion
pratti pratti (Dyar)). —Kulman, 1965. Jour. Econ. Ent. 58: 70 (as predator of Malacosoma
americanum (F.)). —Wang and Brook, 1970. Jour. Econ. Ent. 63: 1971-1973 (toxicological
and biological studies). —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 106-107.

Morphology: Crozier, 1970. Canad. Jour. Genet. and Cytol. 12: 541-546 (pericentric
rearrangement polymorphism).

UNPLACED TAXA OF DOLICHODERINAE

Bothriomyrmex dimmocki Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 417. 3, 2. Mass. (Mt.
Tom, Springfield).

Formica masonia Buckley, 1866. Ent. Soc. Phila., Proc. 6: 165. §. Tex. (Fort Mason).

Formica (Hypochira (!)) subspinosa Buckley, 1866. Ent. Soc. Phila., Proc. 6: 169. %. Tex.

SUBFAMILY FORMICINAE

This subfamily is the predominant group of ants in North America, most common in the north
and in the mountains with their numbers decreasing rapidly toward the southern part of the
United States. They are recognized by the single segmented pedicel, lack of a constriction
between the first and second gastric segments, and the round acidopore, usually with a circle of
hairs.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 126-171, 175-217 (larvae).

—Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 205-222 (larvae). —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 648-656 (larvae).

Biology: Kannowski, 1963. Symp. Genet. et Biol. Ital. 12: 74-102 (flight activities). —Wilson
and Regnier, 1971. Amer. Nat. 105: 279-289 (evolution of alarm-defense system).

Morphology: Blum and Wilson, 1964. Psyche 71: 28-31 (anatomical source of trail substances
in formicine ants). —Hung, 1969. Ent. Soc. Amer., Ann. 62: 455-456 (chromosome numbers
of various spp.).

TRIBE PLAGIOLEPIDINI

Genus PLAGIOLEPIS Mayr

Plagiolepis Mayr, 1861. Eurp. Formicid., p. 42.
Type-species: Formica pygmaea Latreille. Monotypic.

Plagiolepis subg. Anacantholepis Santschi, 1914. Meddel. Goteborgs Mus. Zool. 3: 36.
Type-species: Plagiolepis (Anacantholepis) decora Santschi. Monotypic.

Aporomyrmex Faber, 1969. Pflanzenschutzber. 39: 52.
Type-species: Aporomyrmex ampeloni Faber. Orig. desig.

Plagiolepis subg. Paraplagiolepis Faber, 1969. Pflanzenschutzber. 39: 65.
Type-species: Plagiolepis xene Starcke. Monotypic.

An Old World genus with a single introduced species found in the United States.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 129, 132 (larvae). —Brown.
1973. In Meggers, et al., Tropical forest ecosystems in Afr. and S. Amer., pp. 178-185.

Superfamily FORMICOIDEA 1423

alluaudi Emery. Calif. (Catalina Is.); W. Indies, Pacific Islands, apparently pantropical.

Introduced, probably native to Africa from where it has been widely distributed by
commerce.

Plagiolepis alluaudi Emery, 1894. Soc. Ent. de France, Ann. 63: 71. %.

Plagiolepis mactavishi Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 166. 9.

Plagiolepis foreli Santschi, 1920. Soc. Vaud. des Sci. Nat., Bul. 53: 165. &.

Plagiolepis foreli var. ornata Santschi, 1920. Soc. Vaud. des Sci. Nat., Bul. 53: 166. 9.

Plagiolepis foreli Mann, 1921. Harvard Univ., Mus. Comp. Zool., Bul. 64: 473. 3. Preoce. by
Santschi, 1920.

Plagiolepis augusti Emery, 1921. Soc. Ent. de Belg., Ann. 61: 317. N. name for foreli
Mann.

Taxonomy: Smith, 1957. N. Y. Ent. Soc., Jour. 65: 196 (also biol. notes). —Brown, 1958. Acta
Hym. 1: 49. —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 85-86 (Polynesia).

Biology: Phillips, 1934. (Hawaii Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc., Bul. 15: 17-18
(in Hawaii). —Taylor and Wilson, 1961. Psyche 68: 143.

Genus ACROPYGA Roger

Acropyga Roger, 1862. Berlin. Ent. Ztschr. 6: 242.
Type-species: Acropyga acutiventris Roger. Monotypic.
Acropyga subg. Rhizomyrma Forel, 1893. Ent. Soc. London, Trans., p. 347.
Type-species: Acropyga (Rhizomyrma) goldii Forel. Desig. by Wheeler, 1911.
Acropyga subg. Atopodon Forel, 1912. Rev. Suisse de Zool. 20: 771.
Type-species: Acropyga (Atopodon) inerzae Forel. Desig. by Wheeler, 1913.
Acropyga subg. Malacomyrma Emery, 1922. Mus. Nat. Hung., Ann. 19: 109.
Type-species: Acropyga silvestrii Emery. Monotypic.

The small yellow ants of this genus are subterranean in habit and are fond of honeydew. Ac-
cording to Weber (1944) all the New World species may be obligate coccidophiles. Some species
may be of economic importance because they tend and disperse coccids on the roots of coffee
plants.

Revision: Weber, 1944. Ent. Soc. Amer., Ann. 37: 89-122 (New World spp.; also biol. and
association with coccids).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 136 (larvae). —Snelling,
1973. Los Angeles Co. Mus., Contrib. Sci. 236: 6-7 (first record of genus in U. S.).
epedana Snelling. Ariz. (Montezuma Pass, 5800 ft., Huachuca Mts., Cochise Co.) Ecology:
Specimens were found under a stone.
Acropyga epedana Snelling, 1973. Los Angeles Co. Mus., Contrib. Sci. 236: 7-8. %.

TRIBE MYRMELACHISTINI

Genus MYRMELACHISTA Roger

Myrmelachista Roger, 1863. Berlin. Ent. Ztschr. 7: 162.
Type-species: Myrmelachista kraatzii Roger. Monotypic.
Myrmelachista subg. Decamera Roger, 1863. Berlin. Ent. Ztschr. 7: 166. Preoce. by
Mulsant, 1842.
Type-species: Myrmelachista (Decamera) nigella Roger. Monotypic.
Aphomomyrmex subg. Neaphomus Menozzi, 1935. Zool. Jahrb., Abt. f. System 67: 324.
Synonymy questionable.
Type-species: Aphomomyrmex (Neaphomus) goetschi Menozzi. Monotypic.
Hincksidris Donisthorpe, 1944. Ent. Monthly Mag. 80: 59. N. name for Decamera Roger.

A Neotropical genus represented in the United States by a single species probably introduced
into Florida.
Taxonomy: Wheeler, 1934. Harvard Univ., Mus. Comp. Zool., Bul. 77: 187-206. —Wheeler and
Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 129, 137 (larvae). —Kusnezov, 1959. Acta Zool.
Lilloana (Tucuman) 17: 379.

1424 Hymenoptera in America North of Mexico

ramulorum Wheeler. Fla. (Highland City, Polk Co.); W. Indies. Ecology: Specimens in Florida
collected on sweet orange. Probably introduced.
Myrmelachista ambigua ramulorum Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24:

1553925 do.
Taxonomy: Wheeler, 1934. Harvard Univ., Mus. Comp. Zool., Bul. 77: 189. —Smith, 1936.

Puerto Rico Univ., Jour. Agr. 20: 872-878 (also biol notes). —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 47: 138 (larva).
Biology: Anonymous, 1968. FAO Plant Protect. Bul. 16: 53 (Fla. record; biol).

TRIBE BRACHYMYRMECINI
Genus BRACHYMYRMEX Mayr

Brachymyrmex Mayr, 1868. Soc. Nat. Modena, Ann. 3: 163.
Type-species: Brachymyrmex patagonicus Mayr. Monotypic.
Brachymyrmex subg. Bryscha Santschi, 1925. Buenos Aires Mus. Nac. de Hist. Nat., An.

31: 652.
Type-species: Brachymyrmex pilipes Mayr. Orig. desig.

Most species of this New World genus are found south of the United States. They usually
form small colonies in the soil under various objects and are easily transported by commerce in
soil or plants. They are fond of honeydew and attend honeydew excreting insects.

Revision: Wheeler, 1903. Psyche 10: 102-103. —Santschi, 1923. Buenos Aires Mus. Nac. de

Hist. Nat., An. 31: 650-674.
Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 129, 139 (larvae).

—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 652 (larvae, Brachymyrmecini).

Biology: Arnaud and Quate, 1951. Pan-Pacifie Ent. 27: 171 (note on swarming).
depilis Emery. N.S. s. to Fla. w. to B. C., Calif. Ecology: Nests are small and inconspicuous

and are in the soil under stones, rotten wood, or other objects.
Brachymyrmex heerii depilis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 635. §.
Brachymyrmex nanellus Wheeler, 1903. Psyche 10: 102. %, d.
Brachymyrmex depilis flavescens Grundmann, 1952. Kans. Ent. Soc., Jour. 25: 117. 8.

Taxonomy: Santschi, 1923. Buenos Aires Mus. Nac. de Hist. Nat., An. 31: 664. —Cole, 1940.
. Amer. Midland Nat. 24: 65. —Gregg, 1944. Ent. Soc. Amer., Ann. 37: 469. —Cole, 1953.
Ent. News 64: 266. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 47: 139 (larva).

—Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 210 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 389. —Smith, 1927. Ent. News 38:
313. —Dennis, 1938. Ent. Soc. Amer., Ann. 37: 469. —Headley, 1952. Ent. Soc. Amer., Ann.

45: 436, 439. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 161-163. —Gregg, 1963.
Ants of Colo., pp. 447-449. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp.

108-109.
obscurior Forel. Fla., s. Tex.; Mexico, Central Amer., W. Indies. Possibly introduced into U. S.
Brachymyrmez heeri var. obscurior Forel, 1893. Ent. Soc. London, Trans., p. 345. %, 2, d.

Taxonomy: Santschi, 1923. Buenos Aires Mus. Nac. de Hist. Nat., An. 31: 654, 666. —Smith,
1936. Puerto Rico Univ., Jour. Agr. 20: 866-867 (also biol. notes). —Brown, 1957. Harvard

Univ., Mus. Comp. Zool., Bul. 116: 237.
Biology: Plank and Smith, 1940. Puerto Rico Univ., Jour. Agr. 24: 60.

TRIBE CAMPONOTINI

Genus CAMPONOTUS Mayr

This large genus of ants is found throughout North America and most other parts of the

world. They are commonly referred to as carpenter ants, especially members of the subgenus
Camponotus, because of their habit of nesting in wood. However, many species nest in the soil,

Superfamily FORMICOIDEA 1425

under stones, or in hollow twigs or branches.

Many subgeneric groupings have been proposed for Camponotus on a worldwide basis, many
of which may or may not be valid or may eventually represent distinct genera. Brown (1973)
gave 54 genus-group names to be considered with Camponotus. Seven subgenera have been
recognized in North America, and these are separated here as in the past literature.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 667-682. —Wheeler, 1910. N. Y. Acad.
Scei., Ann. 20: 295-354.

Taxonomy: Emery, 1920. Rev. Zool. Afr. 8: 16-19 (subgenera). —Wheeler, 1921. Psyche 28:
17. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 181 (larvae). —Wheeler and
Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 216 (larvae). —Wheeler and Wheeler 1970. Ent.
Soc. Amer., Ann. 63: 649 (larvae). —Brown, 1973. In Meggers, et al., Tropical forest
ecosystems in Afr. and S. Amer., pp. 178-185 (genus-group names).

Biology: Smith, 1942. Ent. News 53: 133-135 (effect of reduced food supply on stature).
—Smith, 1942. Tenn. Acad. Sci., Jour. 17: 367-373 (polymorphism). —Riordan, 1960.
Insectes Sociaux 7: 353-355 (location of nests by a radioactive isotope). —Sanders, 1964.
Canad. Ent. 96: 894-909 (biol. of carpenter ants in N. B.). —Patrick, 1969. Amer. Midland
Nat. 82: 605-610 (changes in carpenter ants harboring dicrocoeliid metacercariae).
—Sanders and Baldwin, 1969. Canad. Ent. 101: 416-418 (Iridium-192 as a tag for carpenter
ants). —Sanders, 1970. Ecology 51: 865-873 (distribution of carpenter ant colonies in
spruce fir forests in Ont.). —Ayre and Blum, 1971. Physiological Zool. 44: 77-83 (attraction
and alarm). —Sanders, 1972. Canad. Ent. 104: 1681-1687 (seasonal and daily activity
patterns in n.w. Ont.). —Sanders, 1973. Ent. Soc. Ontario, Proc. 102: 13-16 (aggregation of
alate carpenter ants in Ont.).

Genus CAMPONOTUS Subgenus CAMPONOTUS Mayr

Camponotus Mayr, 1861. Die Europaischen Formiciden, pp. 25, 35.
Type-species: Formica ligniperda Latreille. Desig. by Bingham, 1903.

Most species of this subgenus build their nests'in decaying wood. Some are common house in-
festing ants that make their nest in the timber and woodwork of buildings and tunnel nest
passages into adjacent wood. Dry, sound wood is rarely attacked, and the presence of nests of
carpenter ants in buildings is usually evidence that the wood was damp and partially decayed.

americanus Mayr. Ont. s. to Fla. w. to Mich., Iowa, Mo., Okla., Tex. Ecology: Prefers to nest in
the soil, usually under stones or rotten logs.
Camponotus americanus Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 661. 9, °.
Camponotus (Camponotus) castaneus stirps rufinasis Santschi, 1936. Rev. d. Ent. 6: 204.
3.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 323-325 (each caste). —Cole, 1940.
Amer. Midland Nat. 24: 82, 84 (also biol. notes). —Gregg, 1944. Ent. Soc. Amer., Ann. 37:
477. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 185 (larva). —Wheeler and
Wheeler, 1958. Ent. Soc. Amer., Ann. 61: 216 (larva).

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 273, 275, 300-301.

Morphology: Ayre and Blum, 1971. Physiological Zool. 44: 77-83 (attraction and alarm by
pheromones).
ferrugineus (Fabricius). Mass., N. Y. s. to Ga. w. to Mich., IIL, Nebr., Kans. Ecology: Nests are
located in and beneath well-rotted logs and stumps with galleries often extending into
the soil. They have also been found in dead standing trees and occasionally in moist or
faulty wood in buildings. Red carpenter ant.
Formica ferruginea Fabricius, 1798. Sup. Ent. System., p. 279. ¥, °-

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 668. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 299, 338-339 (each caste; biol. notes). —Wheeler, 1917. Conn. State Geol. and
Nat. Hist. Survey Bul. 22: 600-601. —Cole, 1940. Amer. Midland Nat. 24: 86. —Brown,
1950. Ent. News 61: 158-160. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 188
(larva).

1426 Hymenoptera in America North of Mexico

Biology: Pricer, 1908. Biol. Bul. 14: 177-218. —Dennis, 1938. Ent. Soc. Amer., Ann. 31:
301-302. —Williams, 1961. Ohio Jour. Sci. 61: 279 (habits; nest). —Smith, 1965. U. S. Dept.
Agr., Tech. Bul. 1326, pp. 67-69 (economic importance).

Morphology: Tanquary, 1913. Ill. State Lab. Nat. Hist., Bul. 9: 454-475 (embryology).
herculeanus (Linnaeus). Newfoundland (Labrador and insular) w. to Alaska s. to N. Y., Pa.,
Wis., Minn., N. Dak., Colo., N. Mex., Utah, Oreg.; Eurasia. Ecology: Probably the
dominant ant in the forests of boreal and alpine N. Amer. Large colonies are found in
rotting logs and stumps, especially conifers.
Formica herculeana Linnaeus, 1758. Syst. Nat., Ed. 10, p. 579. §.
Camponotus herculeanus var. Whymperi Forel, 1902. Ent. Soc. London, Trans., p. 699. &,
Qe

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 330-333 (each caste; biol. notes).
—Cole, 1942. Amer. Midland Nat. 28: 387. —Creighton, 1950. Harvard Univ., Mus. Comp.
Zool., Bul. 104: 367. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 185-187
(larva). —Yasumatsu and Brown, 1957. Kyushu Univ., Fac. Agr., Jour. 11: 45. —Arnoldi,
1967. Zool. Zhur. 46: 1817-1818. —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63:
651 (larva).

Biology: Jones, 1929. Colo. Agr. Expt. Sta. Bul. 341, 96 pp. (tends 19 spp. in 6 gen. of aphids
on 12 spp. of host plants). —Gregg, 1946. Amer. Midland Nat. 35: 753. —Brown, 1949. Ent.
News 60: 99. —Holldobler, 1950. Ztschr. f. Angew. Ent. 31: 583 (biol. and habits in
Europe). —Weber, 1950. Amer. Ent. Soc., Trans. 76: 188. —Perttunen, 1955. Ann. Ent.
Fenn. 21: 38 (reactions to air humidity). —Brown, 1955. Ent. News 66: 47-48. —Holldobler,
1961. Insectes Sociaux 8: 14 (rhythmical behavior). —Holldobler, 1962. Waldhygiene 4: 228
(nest types, ecomomic importance in Germany). —Holldobler, 1962. Ztschr. f. Angew. Ent.
49: 338. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 164-165. —Gregg, 1963. Ants of
Colo., pp. 656-660. —Sanders, 1964. Canad. Ent. 96: 894 (biol. and habits, N. B.). —Sanders,
1972. Canad. Ent. 101: 1618-1687 (seasonal and daily activity patterns, Ont.).

Morphology: Ayre, 1963. Ent. Expt. and Appl. 6: 165-170 (feeding behavior and digestion).
—Ayre and Blum, 1971. Physiological Zool. 44: 77-83 (attraction and alarm by
pheromones).

laevigatus (Smith). Mont., Colo., N. Mex. w. to B. C., Oreg., Calif.; Mexico. Ecology: Found in
wooded and forested areas where it nests in rotting logs and stumps. Occasionally found
in buildings.
Formica laevigata Smith, 1858. Cat. Hym. Brit. Mus., v. 6, p. 55. ¥, ?.
Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 327-330 (each caste; biol. notes).

—Cole, 1942. Amer. Midland Nat. 28: 387-3888. —Wheeler and Wheeler, 1968. Ent. Soc.
Amer., Ann. 61: 216 (larva).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 556. —Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 29. —Furniss, 1944. Oreg. Agr. Expt. Sta. Cir. 158: 1-12.
—Gregg, 1963. Ants of Colo., pp. 660-663.

modoc Wheeler. S. Dak., Colo. w. to B. C., Oreg., Calif. Ecology: Found in forested areas
where it makes its nests in rotting logs and stumps. Has been considered a subspecies
of herculeanus or pennsylvanicus by various authors.

Camponotus (Camponotus) herculeanus var. modoc Wheeler, 1910. N. Y. Acad. Sci., Ann.
20: 299, 333. 3, 2, do.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 387. —Brown, 1950. Ent. News 61: 158.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 557. —Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 29. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 90.
—Furniss, 1944. Oreg. Agr. Expt. Sta. Cir. 158: 1-12. —Gregg, 1963. Ants of Colo., pp.
665-669.

novaeboracensis (Fitch). N. S., Que. s. to Va. w. to B. C., Oreg., Utah, Colo. Ecology: Prefers
wooded areas where it normally nests in rotting logs and stumps. Sometimes a house
pest.
Formica Novaeboracensis Fitch, 1855. N. Y. State Agr. Soc., Trans. 14: 766. %.

Superfamily FORMICOIDEA 1427

Camponotus herculeanus ligniperdus var. pictus Forel, 1879. Soc. Vaud. des Sci. Nat., Bul.
163595 OS O.

Camponotus herculeanus ligniperdus var. noveboracensis Forel, 1899. Soc. Ent. de Belg.,
Ann. 43: 447. Emend.

Camponotus herculeanus ligniperda var. rubens Wheeler, 1906. Psyche 13: 41. 9, d.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 340 (each caste). —Buren, 1944.
Iowa State Col., Jour. Sci. 18: 298. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46:
182-185 (larva). —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larva).

Biology: Jones, 1929. Colo. Agr. Expt. Sta. Bul. 341: 96 pp. (attends aphids). —Gregg, 1944.
Ent. Soc. Amer., Ann. 37: 457-458. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11:
250-251. —Kannowski, 1959. Insectes Sociaux 6: 134-135. —Wheeler and Wheeler, 1963.
Ants of N. Dak., pp. 165-168. —Gregg, 1963. Ants of Colo., pp. 663-665. —Sanders, 1964.
Canad. Ent. 96: 896 (biology and habits, N. B.). —Gotwald, 1968. N. Y. Ent. Soc., Jour. 76:
278-296 (food gathering behavior). —Sanders, 1972. Canad. Ent. 104: 1681-1687 (seasonal
and daily activity pattern, Ont.).

Morphology: Smith, 1942. Tenn. Acad. Sci., Jour. 17: 368.
pennsylvanicus (DeGeer). N. B., Que. s. to Fla. w. to N. Dak., Tex. Ecology: Nests are found in

live and dead trees, rotting logs and stumps, and in wood products such as fences,
telegraph poles, and buildings. A common and important house infesting ant because of
its adaptability to nest in woodwork of buildings. It commonly feeds on household foods,
and it is annoying. Nests are started in buildings usually in moist or faulty wood. The
first native North American ant to be described. Black carpenter ant.

Formica pensylvanica (!) DeGeer, 1773. Mem. Serv. Hist. Ins., v. 3, p. 603. 3, 2, d.

Camponotus herculeanus herculeanus herculeano-pennsylvanicus Forel, 1879. Soc. Vaud.
des Sci. Nat., Bul. 16: 57. % (?).

Camponotus herculeanus pennsylvanicus var. mahican Wheeler, 1910. N. Y. Acad. Sci.,
Ann. 20: 299, 338. N. name for herculeano-pennsylvanicus Forel.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 335-336 (each caste). —Wheeler,
1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 600. —Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 293. —Townsend, 1945. Ky. Agr. Expt. Sta. Cir. 59: 1-27 (references to 1945).
—Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 187-188 (larva). —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larva).

Biology: McCook, 1877. Amer. Ent. Soc., Trans. 6: 253-296. —Pricer, 1908. Biol. Bul. 14:
177-218. —Herrick, 1914. Insects Injurious to the Household and Annoying to Man, pp.
177-178. —Gibson, 1916. Canad. Ent. 48: 365-366. —Graham, 1918. Minn. State Ent. Rpt.
17: 32-40 (as destroyers of sound wood). —Back, 1937. U. S. Dept. Agr. Leaflet 147. —Van
Pelt, 1958. Tenn. Acad. Sci., Jour. 33: 120-122 (parasitism by a fungus). —Wheeler and
Wheeler, 1963. Ants of N. Dak., pp. 168-170. —Sanders, 1964. Canad. Ent. 96: 899 (biology
and habits, N. B.). —Smith, 1965. U. S. Dept. Agr. Tech. Bul. 1326, pp. 63-67 (economic
importance). —Sanders, 1972. Canad. Ent. 104: 1681-1687 (seasonal and daily activity
patterns, Ont.).

Morphology: McCook, 1878. Phila. Acad. Nat. Sci., Proc. 30: 15-19. —Fielde, 1903. Biol. Bul. 5:
320-325. —Fielde and Parker, 1904. Phila. Acad. Nat. Sci., Proc. 56: 642-649. —Forbes,
1938. Ent. Soc. Amer., Ann. 31: 181-195 (anatomy and histology of worker). —Smith, 1942.
Tenn. Acad. Sci., Jour. 17: 367-373. —Forbes, 1952. N. Y. Ent. Soc., Jour. 60: 157-171 (male
genitalia). —Forbes, 1954. Jour. Morph. 95: 523-548 (male reproductive system). —Forbes,
1956. Insectes Sociaux 3: 505-511 (male digestive tract). —Forbes and McFarlane, 1961. N.
Y. Ent. Soc., Jour. 69: 92-103 (comparative anatomy of digestive glands of female and
male). —Keister, 1963. Ent. Soc. Amer., Ann. 56: 336-340 (trachael system). —Hermann
and Blum, 1968. Psyche 75: 216-227 (poison apparatus). —Ayre and Blum, 1971.
Physiological Zool. 44: 77-83 (attraction and alarm by pheromones).

quercicola Smith. Calif. (Los Angeles Co. to San Diego Co.). Ecology: Nests in dead limbs of
Quercus agrifolia; nocturnal.
Camponotus (Camponotus) quercicola Smith, 1953. N. Y. Ent. Soc., Jour. 61: 211-214. 3,
ne

1428 Hymenoptera in America North of Mexico

schaefferi Wheeler. S. Ariz. Ecology: Nests in dead oak limbs at elevations of about 5000 to
8000 feet.

Camponotus schaefferi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 88. 8, 2.
Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 344-345 (also biol. notes).

Biology: Wheeler, 1917. Amer. Acad. Arts. and Sci., Proc. 52: 557.
texanus Wheeler. Tex. Ecology: Nests have been found in oak logs.
Camponotus texanus Wheeler, 1903. Psyche 10: 108. §, 2, d.

Taxonomy: Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 90. —Wheeler, 1910. N. Y. Acad. Sci.,
Ann. 20: 300, 344 (also biol. note).

Genus CAMPONOTUS Subgenus TANAEMYRMEX Ashmead

Tanaemyrmex Ashmead, 1905. Canad. Ent. 37: 384.

Type-species: Formica longipes Gerstaecker. Orig. desig.
Myrmoturba Forel, 1912. Soc. Ent. de Belg., Mem. 20: 91.

Type-species: Formica maculata Fabricius. Desig. by Wheeler, 1913.

Most species of this subgenus nest in the soil under stones or other objects; occasionally nests

may be surmounted by a small crater. The ants rarely nest in wood, but, if so, the wood is
usually buried in the soil.

acutirostris Wheeler. Tex., N. Mex., Ariz. Ecology: Nests have been found in the ground
under stones.

Camponotus (Camponotus) acutirostris Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 298, 317.
COE On

Camponotus acutirostris var. clarigaster Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34:
420. 8.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 561-562.

castaneus (Latreille). N. Y. s. to Fla. w. to Iowa, Okla., Tex. Ecology: Nests in rotting logs and

stumps, exposed soil, or in soil under objects. Occasionally enters buildings usually in
search of food.

Formica castanea Latreille, 1802. Hist. Nat. Fourmis, p. 118. 8, 2, d.

Formica mellea Say, 1831. Deser. N. Spp. N. Amer. Ins. Found in La. by Jos. Barabino, p.
14. 3.

Camponotus clarus Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 660. &.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 321-323. —Cole, 1940. Amer.
Midland Nat. 24: 82, 84.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 402. —Gregg, 1944. Ent. Soc. Amer.,
Ann. 37: 457, 477. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326, pp. 60-62.
dumetorum Wheeler. S. Calif. Ecology: Nests are common in the chaparral and are in the soil
usually surmounted by flat craters.

Camponotus (Camponotus) maculatus dumetorum Wheeler, 1910. N. Y. Acad. Sci., Ann.
20: 354. 8, d.

Taxonomy: Snelling, 1970. Ent. Soc. Wash., Proc. 72: 390-397.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 560. —Mallis, 1941. South. Calif.
Acad. Sci., Bul. 40: 90-91.
festinatus (Buckley). Tex., Colo., N. Mex., Ariz., Calif.; Mexico. Ecology: Nests in the ground
under stones, logs, or dried cow dung.
Formica festinata Buckley, 1866. Ent. Soc. Phila., Proc. 6: 165. 8, °.

Camponotus (Camponotus) fumidus var. pubicornis Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 668, 670. %.

Camponotus fragilis Pergande, 1893. Calif. Acad. Sci., Proc. 4: 26. °.

Camponotus (Camponotus) fumidus var. spurcus Wheeler, 1910. N. Y. Acad. Sci., Ann. 20:
299, 315. 3, 2.

Superfamily FORMICOIDEA 1429

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 312 (also biol. notes). —Snelling, 1968.
Ent. Soc. Wash., Proc. 70: 350-355. —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63:
651 (larva).
Biology: Wheeler, 1901. Amer. Nat. 35: 518, 520, 533.
incensus Wheeler. Fla. (Pigeon Key near Miami).
Camponots (Tanaemyrmex) incensus Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 14. 8.

ocreatus Emery. N. Mex., Ariz., s. Nev., s. Calif.; Mexico. Ecology: Nests are found under
stones.

Camponotus (Camponotus) maculatus ocreatus Emery, 1893. Zool. Jahrb., Abt. f. System.
7: 668, 673. &.

Camponotus (Camponotus) acutirostris primipilaris Wheeler, 1910. N. Y. Acad. Sci., Ann.
20: 298, 319. ¥, 2.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 561-562. —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larva).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 272. —Cole, 1966. Brigham Young Univ., Sci.
Bul., Biol. Ser. 7: 20 (Nev. Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon,
pale

sansabeanus bulimosus Wheeler. N. Mex., Ariz. Ecology: Nests in the ground under stones.

Camponotus (Camponotus) maculatus bulimosus Wheeler, 1910. N. Y. Acad. Sci., Ann. 20:
B08 9519S:

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 560-561. —Cole, 1954. Tenn.
Acad. Sci., Jour. 29: 272.

sansabeanus sansabeanus (Buckley). Ark., La. w. to Colo., Utah, Ariz., s. Calif. Ecology: Nests
under rocks and stones in dry woods.
Formica San Sabeana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 167. 8, 2, d.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 672-673. —Wheeler, 1910. N. Y.
Acad. Sci., Ann. 20: 298, 307-308 (also biol. notes). —Cole, 1942. Amer. Midland Nat. 28:
387. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 196 (larva).

Biology: Wheeler, 1901. Amer. Nat. 35: 518, 520, 533. —Wheeler, 1910. Ants, pp. 349, 393.
—Gregg, 1963. Ants of Colo., pp. 669-671. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, p. 111 (half of collections belong to subsp. torrefactus).

Morphology: Wheeler, 1910. Ants, pp. 24, 49.

sansabeanus torrefactus Wheeler. Colo., Ariz., Utah, Nev. Ecology: Nests are found under
stones.

Camponotus (Camponotus) maculatus sansabeanus var. torrefactus Wheeler, 1910. N. Y.
Acad. Sci., Ann. 20: 298, 308. 8, 3.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 387, 388 (also biol. note).
Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 559.

semitestaceus Emery. Okla., Tex. w. to Wash., Oreg., Calif.; Mexico. Ecology: Nests are found
under stones or in soil surmounted by low craters. Most U. S. records and references in
the literature to maccooki Forel refer to semitestaceus; maccooki is apparently
restricted to Guadalupe Is.

Camponotus (Camponotus) maculatus vicinus var. semitestaceus Emery, 1893. Zool.
Jahrb., Abt. f. System. 7: 668, 672. °.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 298, 304, 306. —Snelling, 1970. Ent. Soc.
Wash., Proc. 72: 390-397 (separation of maccooki and semitestaceus).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 345. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 559. —Essig, 1926. Ins. West. N. Amer., p. 868. —Cole, 1934. Ent.
Soc. Amer., Ann. 27: 403. —Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Cir. 342: 28.
—Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 91-92. —Cole, 1966. Brigham Young Univ.,

Sci. Bul., Biol. Ser. 7: 19-20. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp.
112-113.

1430 Hymenoptera in America North of Mexico

socius Roger. N. C. s. to Fla. w. to La.; Brazil. Ecology: Nests are built in branches and rotten
logs that are covered by sand. Introduced into the U. S.
Camponotus socius Roger, 1863. Berlin. Ent. Ztschr. 7: 140. 2.
Camponotus (Tanaemyrmex) socius var. osceola Wheeler, 1932. N. Y. Ent. Soc., Jour. 40:
15. 8.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 298, 319-321 (each caste).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 14-15. —Creighton, 1950. Harvard Univ.,
Mus. Comp. Zool., Bul. 104: 374. —Holldobler, 1971. Z. Vergl. Physiol. 75: 123-142
(recruitment behavior).

tortuganus Emery. S. Fla. Ecology: May nest in soil under stones or under rotting wood. A
frequent house pest and reported to have been found nesting in siding and roofing of
buildings.

Camponotus maculatus tortuganus Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 336. &.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 310-312 (each caste).

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 13-14. —Smith, 1965. U.S. Dept. Agr.,
Tech. Bul. 1326, pp. 62-63.
vafer Wheeler. Ariz. (Huachuca Mts., 5000 to 6000 ft.). Ecology: Nests were found under

stones.

Camponotus (Camponotus) vafer Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 315. %, 2.

vicinus Mayr. Man., N. Dak., Colo., Okla., w. to B. C., Oreg., Calif.; Mexico. Ecology: Nests in

soil under stones or in rotting wood buried in the soil.

Formica Tejonia Buckley, 1866. Ent. Soc. Phila., Proc. 6: 161. ¢. Questionably the same as
vicinus.

Camponotus vicinus Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 940. &.

Camponotus (Camponotus) maculatus vicinus var. nitidiventris Emery, 1893. Zool.
Jahrb., Abt. f. System. 7: 668, 672. 8.

Camponotus (Camponotus) maculatus vicinus var. infernalis Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 298, 305. 8.

Camponotus (Camponotus) maculatus vicinus var. luteangulus Wheeler, 1910. N. Y.
Acad. Sci., Ann. 20: 298, 304. %, do.

Camponotus (Camponotus) maculatus vicinus var. maritimus Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 298, 305. 3, 9, d.

Camponotus (Camponotus) maculatus vicinus var. plorabilis Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 298, 303. 3, 2, d.

Camponotus (Myrmoturba) maculatus vicinus var. subrostrata Forel, 1914. Deut. Ent.
Ztschr., p. 620. %.

Camponotus (Myrmoturba) maculatus Maccooki berkeleyensis Forel, 1914. Deut. Ent.
Ztschr., p. 619. %.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 387. —Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bul. 104: 381. —Snelling, 1970. Ent. Soc. Wash., Proc. 72: 390-397. —Wheeler
and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 220 (larva).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 345. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 599-560. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 92.
—McClure, 1943. Ecol. Monog. 13: 19. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
170-171. —Gregg, 1963. Ants of Colo., pp. 671-675. —Cole, 1966. Brigham Young Univ., Sci.
Bul., Biol. Ser. 7: 20 (Nev. Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon,
pp. 113-114.

Genus CAMPONOTUS Subgenus MYRMOTHRIX Forel

Myrmothrix Forel, 1912. Soc. Ent. de Belg., Mem. 20: 91.
Type-species: Formica abdominalis Fabricius. Desig. by Wheeler, 1913.

Most members of this subgenus are Neotropical with two forms extending into southern
United States.

Revision: Santschi, 1936. Rev. de Ent. 6: 207-218.

Superfamily FORMICOIDEA 1431

abdominalis floridanus (Buckley). N. C. s. to Fla. w. to s. Miss. Ecology: Nests are in and
under rotten logs and stumps, usually in damp situations. A house infesting ant,
especially in Florida, where it is known to nest in the woodwork of buildings and feed
on household foods. May also damage beehives. C. (M.) abdominalis abdominalis (F.)
occurs in Central and South America. Florida carpenter ant.

Formica Floridana Buckley, 1866. Ent. Soe. Phila., Proc. 6: 161. 8.
Camponotus atriceps stirps Yankee Forel, 1884. Soc. Vaud. des Sci. Nat., Bul. 20: 340. &.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 668. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 299, 325-326. —Santschi, 1936. Rev. de Ent. 6: 213-214.

Biology: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 326. —Smith, 1965. U. S. Dept. Agr., Tech.
Bul. 1326, pp. 69-70 (economic importance).
abdominalis transvectus Wheeler. S. Tex.; Mexico.
Camponotus (Camponotus) abdominalis transvectus Wheeler, 1910. N. Y. Acad. Sci., Ann.
20: 299, 326. %, 2, d.

Taxonomy: Santschi, 1936. Rev. de Ent. 6: 218.

Genus CAMPONOTUS Subgenus MYRMENTOMA Forel

Myrmentoma Forel, 1912. Soc. Ent. de Belg., Mem. 20: 92.
Type-species: Formica lateralis Olivier. Desig. by Wheeler, 1913.

The Nearctic forms of this subgenus nest in insect galls, in branches and stems of plants,
under bark of trees, in wood and buildings, and sometimes in the soil. Colonies are small, consist-
ing of a few dozen to several hundred individuals. The listing below essentially follows that of
Creighton (1950) though there will undoubtedly be some changes in the near future.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 216-232. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bul. 104: 382-390.
anthrax Wheeler. S. Calif. Ecology: Nests were found in soil under large stones.
Camponotus anthrax Wheeler, 1911. N. Y. Ent. Soc., Jour. 19: 96. 3, 2, od.

Taxonomy: Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 650 (larva).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 558. —Mallis, 1941. South. Calif.
Acad. Sci., Bul. 40: 89.
caryae caryae (Fitch). N. Y. s. to D. C. w. to Mich., Ohio. Ecology: Apparently associated with
trees, especially hickory.
Formica caryae Fitch, 1855. N. Y. State Agr. Soc., Trans. 14: 855. ¥, 9, d.

Camponotus marginatus discolor var. cnemidatus Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 669, 678. %.

Taxonomy: Smith, 1940. Ent. Soc. Wash., Proc. 42: 137-141 (each caste; biol. note).
caryae clarithorax Emery. S. Calif.

Camponotus marginatus discolor var. clarithorax Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 670, 678. %, 2, do.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 231-232 (each caste). —Wheeler,
1910. N. Y. Acad. Sci., Ann. 20: 301.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 89. 5
caryae discolor (Buckley). Ohio, S. C., Fla. w. to N. Dak., Iowa, Kans., Tex. Ecology: Nests are
in plant cavities in twigs, branches, under bark, in logs and stumps, or in insect galls. A
house infesting ant that may nest in woodwork in houses, especially in preformed
cavities or in rotting or faulty wood.
Formica discolor Buckley, 1866. Ent. Soc. Phila., Proc. 6: 166. 3, °.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 230-231 (each caste). —Buren, 1944.
Iowa State Col., Jour. Sci. 18: 293.

Biology: Wheeler, 1902. Tex. Acad. Sci., Trans. 4: 23. —Wesson and Wesson, 1940. Amer.
Midland Nat. 24: 103. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 172-173. —Smith,
1965. U.S. Dept. Agr., Tech. Bul. 1326, pp. 70-71.

1432 Hymenoptera in America North of Mexico

essigi Smith. Idaho, Oreg., Nev., Calif.
Camponotus caryae var. essigi Smith, 1923. Ent. News 24: 306. 9.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 89.

hyatti bakeri Wheeler. Calif. (Channel Islands). Ecology: Nests in soil.
Camponotus hyatti var. bakeri Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 271. %, °@.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 346.
hyatti hyatti Emery. Nev., Calif.; Mexico (Baja Calif.). Ecology: Colonies have been found in
the stem of Yucca and in soil under a dead juniper limb.
Camponotus (Camponotus) hyatti Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 669, 680.
3.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300, 345-346.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 19 (Nev. Test Site).
nearcticus Emery. Ont. s. to Fla. w. to N. Dak., Colo., Tex.; B. C. s. to Idaho, Calif. Ecology:
Forms small colonies in dead twigs and branches, under bark of live and dead trees, in
insect galls, pine cones, and rotting logs and stumps; also in wood products such as fene:
posts and in woodwork of houses, especially the roofing.
Formica americana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 154. 3, 2. Questionably the
same as nearcticus.
Camponotus (Camponotus) marginatus var. nearcticus Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 669 675. 8, 9.
Camponotus (Camponotus) marginatus var. minutus Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 669, 676. %, @.
Camponotus (Camponotus) marginatus var. decipiens Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 669, 676. %, °.
Camponotus fallax fallax var. pardus Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 225. &,

een Or

Camponotus fallax fallax var. tanquaryi Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 226.
Ce Lenn

Camponotus fallax rasilis var. pavidus Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 228.
oe?

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300. —Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 293. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 193 (larva).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 21: 402-403. —Wheeler, 1910. N. Y. Ent.
Soe., Jour. 18: 220-221. —Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17: 24.
—Kannoswki, 1959. Insectes Sociaux 6: 134. —Gregg, 1963. Ants of Colo., pp. 675, 677.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 173-174. —Smith, 1965. U. S. Dept.
Agr., Tech. Bul. 1326, pp. 72-73 (economic importance).

nevadensis Gregg. Nev. (Fuller Lake, 3 mi. S. Verdi, 6000 ft.).
Camponotus (Myrmentoma) nevadensis Gregg, 1973. Southwest. Nat. 18: 39-43. %, 2, 6.
sayi Emery. N. C. s. to Fla. w. to Nebr., Kans., Colo., Ariz., Calif. Ecology: The small colonies
are found in insect galleries in wood, in twigs and branches, insect galls, under bark, in
stalks of plants, and in rotting logs and stumps. A house infesting ant which may nest in
woodwork and feed on human foods. In most literature as rasilis Wheeler.
Camponotus (Camponotus) sayi Emery, 1894. Zool. Jahrb., Abt. f. System. 7: 679. §.
Camponotus sayi var. bicolor Pergande, 1894. Calif. Acad. Sci., Proc. 4: 161. 3, 2, d.
Preoce. in Camponotus by Latreille, 1798.
Camponotus fallax rasilis Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 227. 3, 2, d.
Camponotus sayi californica Emery, 1925. In Wytsman, Gen. Ins. 183: 118. N. name for
bicolor Pergande.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300. —Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 293-294. —Snelling, 1968. Ent. Soc. Wash., Proc. 70: 355-358.

Biology: Wheeler, 1910. N. Y. Ent. Soe., Jour. 18: 228. —Dennis, 1938. Ent. Soc. Amer., Ann.
31: 302-303. —Gregg, 1963. Ants of Colo., pp. 675, 677. —Smith, 1965. U. S. Dept. Agr.,
Tech. Bul. 1326, pp. 73-74 (economic importance).

Superfamily FORMICOIDEA 1433

subbarbatus Emery. N. Y.s. to N. C. w. to Mich., Iowa, Kans. Ecology: Nests are in plant
cavities, in twigs and branches, and under bark.
Camponotus (Camponotus) marginatus subbarbatus Emery, 1893. Zool.
Jahrb., Abt. f. System. 7: 669, 676. %, 2, d.
Camponotus (Camponotus) marginatus subbarbatus var. paucipilis Emery, 1893. Zool.
Jahrb., Abt. f. System. 7: 669, 677. %, do.

Taxonomy: Wheeler, 1910. N. Y. Ent. Soc., Jour. 18: 222, 229. —Wheeler, 1910. N. Y. Acad.
Sci., Ann. 20: 300. —Buren, 1944. Iowa State Col., Jour. Sci. 18: 293.

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24: 90, 103.

Genus CAMPONOTUS Subgenus COLOBOPSIS Mayr

Colobopsis Mayr, 1861. Die Europaischen Formiciden, pp. 25, 38.
Type-species: Formica truncata Spinola. Desig. by Bingham, 1903.

Ants of this subgenus are most abundant in the southern portions of the United States. They
make their nests in hollow twigs or branches of trees and shrubs, in insect galls and nuts. The

soldiers and females have a peculiar cylindrical, truncated head which the soldiers use for
blocking the single entrance hole to the nest.

Revision: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 139-158.

Taxonomy: Wheeler and Wheeler 1953. Ent. Soc. Amer., Ann. 46: 188 (larvae).

etiolatus Wheeler. Tex.; Mexico. Ecology: Nests have been found in insect galls and in twigs
of trees.
Camponotus (Colobopsis) abditus var. etiolatus Wheeler, 1904. Amer. Mus. Nat. Hist., Bul.
201 5ON OR 2 OG

Taxonomy: Wheeler, 1934. Harvard Univ., Mus. Comp. Zool., Bul. 77: 216. —Wheeler and
Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 650 (larva).
Biology: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 352-353.
hunteri Wheeler. Tex. (Victoria). Ecology: Type series taken from twig of a pecan.
Camponotus (Colobopsis) pylartes var. hunteri Wheeler, 1910. N. Y. Acad. Sci., Ann. 20:
301, 353. 8, 2f.
Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 562.

impressus (Roger). Md. s. to Fla. w. to cent. Tex. Ecology: Colonies have been found in culms
of sedges.
Colobopsis impressa Roger, 1863. Berlin. Ent. Ztschr. 7: 160. &.

Taxonomy: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 144-146 (each caste). —Wheeler,
1910. N. Y. Acad. Sci., Ann. 20: 302. —Smith, 1955. Brooklyn Ent. Soc., Bul. 50: 98.
Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 16.
mississippiensis Smith. Md. s. to Fla. w. to Il, Okla., La.
Camponotus (Colobopsis) mississippiensis Smith, 1923. Psyche 30: 83. 9, 2.
Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 188-190 (larva).

Biology: Smith, 1923. Ent. News 35: 127. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 303, 306.
obliquus Smith. Ala., Miss. Ecology: Found nesting in a hickory nut.
Camponotus (Colobopsis) obliquus Smith, 1930. Ent. Soc. Amer., Ann. 23: 256. Y.
Biology: Smith, 1931. Ent. News 42: 23.
papago Creighton. Ariz.; Mexico.
Camponotus (Colobopsis) papago Creighton, 1952. Psyche 59: 153-162. 9, f, °, d.
pylartes fraxinicola Smith. N. C. to Ala. w. to Ark.
Camponotus (Colobopsis) pylartes fraxinicola Smith, 1923. Psyche 30: 86. %, 2.

Biology: Smith, 1924. Ent. News 35: 127.

1434 Hymenoptera in America North of Mexico

pylartes pylartes Wheeler. La., Tex. Ecology: Colonies found in twigs and spines of trees and
shrubs.
Camponotus (Colobopsis) pylartes Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 147. 8,
DOr

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 120. —Wheeler, 1910. N. Y. Acad.
Scei., Ann. 20: 301.

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 153-158.

Genus CAMPONOTUS Subgenus MYRMAPHAENUS Emery

Myrmaphaenus Emery, 1920. Rev. Zool. Bot. Africaines 8: 237.
Type-species: Camponotus leydigi Forel. Orig. desig.

Paracolobopsis Emery, 1920. Rev. Zool. Bot. Africaines 8: 249.
Type-species: Camponotus salvini Forel. Orig. desig.

Neomyrmamblys Wheeler, 1921. Psyche 28: 19.
Type-species: Camponotus fastigatus Roger. Desig. by Santschi, 1921.

Only two species of this Neotropical subgenus reach the United States.

ulcerosus Wheeler. Tex., Ariz.; Mexico. Ecology: Nests are constructed in the soil beneath
stones. A carton shield is constructed at the nest entrance with an opening the same
size as the head of the major worker. Thus, the major worker functions as a door in this
opening.
Camponotus (Camponotus) ulcerosus Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 351. 8.
Camponotus (Camponotus) bruesi Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 299, 349. 3.

Taxonomy: Creighton, 1951. Psyche 58: 47-64 (also biology).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 562. —Creighton, 1953. Psyche
60: 82-84. —Samuelson, 1961. Pan-Pacific Ent. 37: 189 (association with larvae of Strymon
melinus).

yogi Wheeler. S. Calif. Ecology: Most colonies have been found in living stems of Haplopappus
pinifolius where they live in burrows made by buprestids.
Camponotus yogi Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 420. 8.

Taxonomy: Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 650 (larva). —Creighton
and Snelling, 1966. Psyche 73: 187-195 (female, male; biological notes).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 562. —Mallis, 1941. South. Calif.
Acad. Sci., Bul. 40: 92.

Genus CAMPONOTUS Subgenus MYRMOBRACHYS Forel

Myrmobrachys Forel, 1912. Soc. Ent. de Belg., Mem. 20: 91.
Type-species: Formica senex Smith. Desig. by Wheeler, 1913.

This Neotropical subgenus extends into Florida, Texas, and Arizona. The species found in the
United States usually nest under bark of trees, in branches of trees and shrubs, and in logs and
stumps.

mina Forel. S. Ariz.; Mexico. Ecology: Most colonies have been found nesting in mesquite.
Camponotus Mina Forel, 1879. Soc. Vaud. des Sci. Nat., Bul. 16: 83. %.
Camponotus erythropus Pergande, 1894. Calif. Acad. Sci., Proc. 4: 28. 8.
Camponotus (Camponotus) mina zuni Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 300. &.

Taxonomy: Creighton, 1965. Amer. Mus. Novitates 2239: 1-7 (also biological notes).

planatus Roger. S. Fla., S. Tex.; W. Indies, Mexico. Ecology: Arboreal, nesting in branches and
under bark.
Camponotus planatus Roger, 1863. Berlin. Ent. Ztschr. 7: 148. 8, 2, d.

Taxonomy: Wheeler, 1910. N. Y. Acad. Sci., Ann. 20: 301, 348-349. —Wheeler and Wheeler,
1953. Ent. Soc. Amer., Ann. 46: 194 (larva).

Superfamily FORMICOIDEA 1435

Biology: Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 15. —Wheeler, 1942. Harvard Univ., Mus.
Comp. Zool., Bul. 90: 258.
trepidulus Creighton. S. Ariz. (Baboquivari Mtns.). Ecology: Colonies were found in dead limbs
of Quercus oblongifolia.
Camponotus (Myrmobrachys) trepidulus Creighton, 1965. Amer. Mus. Novitates 2239: 7-9.
Cee SeuGr

TRIBE LASIINI
Genus LASIUS Fabricius

Ants of this holarctic genus nest in exposed soil, under objects or in rotting wood. Colonies are
small to moderate in size. Workers attend and may foster honeydew-excreting insects. The
treatment here follows the revision by Wilson (1955).

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637-639. —Wheeler, 1916. Psyche 23:
168-173. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 1-199.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 147-150 (larvae).

Biology: Kannowski, 1959. Insectes Sociaux 6: 145-146 (flight comparisons). —Pontin, 1961.
Jour. Animal Ecology 30: 47-54 (population stabilization and competition between L. flavus
and L. niger). —Pontin, 1963. Jour. Animal Ecology 32: 565-567 (competition and ecology
of L. flavus and L. niger).

Morphology: Bernardi, et. al., 1967. Tetrahedron Letters 40: 3893-3896 (components of
secretion of mandibular glands).

Genus LASIUS Subgenus LASIUS Fabricius

Lasius Fabricius, 1805. Systema Piezatorum, p. 415.
Type-species: Formica nigra Linnaeus. Desig. by Bingham, 1903.
Donisthorpea Morice and Durrant, 1914. Ent. Soc. London, Trans., pp. 421-423.
Type-species: Formica nigra Linnaeus. Orig. desig.
alienus (Foerster). N. S., N. B. s. to Fla. w. to Man. N. Dak., S. Dak., Neb., Kans., Ark., Miss.;

B. C., Mont., Idaho, Wash., Oreg., Calif., S. Ariz.; Mexico; Eurasia. Ecology: In N. Amer.,
this ant shows a preference for well-shaded woodlands where it nests in rotting logs
and stumps or under stones. Only occasionally has it been found in open areas. In the
early American literature, this species has been recorded as americanus Emery, and
alienus has commonly been confused with neoniger. A frequent house pest. Cornfield
ant is the approved common name for alienus.

Formica aliena Foerster, 1850. Hym. Studien 1: 36-38. 8, d.

Prenolepis lasioides Emery, 1869. Accad. Natur. Napoli, Ann. 2: 6-7. 8, °, d.

Prenolepis fuscula Emery, 1869. Accad. Natur. Napoli, Ann. 2: 8. 9.

Lasius fumatus Emery, 1870. Soc. Ent. Ital., Bol. 2: 194.

Lasius niger var. americanus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: Gade GEORG.

Lasius niger var. grandis Forel, 1909. Soc. Ent. de Belg., Ann. 53: 104-105. %.

Lasius niger alienus var. alieno-americanus Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 525-526. ?.

Lasius niger turcicus Santschi, 1921. Soc. Esp. Hist. Nat., Bol. 21: 115-116. % (2 misdet.).

Lasius niger lasioides var. barbara Santschi, 1921. Soc. Esp. Hist. Nat., Bol. 21: 170. 9.

Acanthomyops niger alienus var. flavidus Kuznetzov-Ugamskij, 1927. Rev. Russ. d’Ent.
21: 189. %.

Acanthomyops niger alienus var. turkmenus Kuznetzov-Ugamskij, 1927. Rev. Russ. d’Ent.
21: 189. &.

Lasius brunneus var. obscurata Stitz, 1930. Mitt. Zool. Mus. Berlin 16: 239-240. , °.

Lasius alienus illyricus Zimmermann, 1934. Zool.-Bot. Gesell. Wien, Verh. 84: 50-52. ¥, °,
3.

Lasius alienus var. pannonica Roszler, 1942. Siebenburgischer Verh. Naturw.,
Hermannstadt, Verh. und Mitt. 91-92 :40. 9, °.

Lasius alienus var. pontica Starcke, 1944. Ent. Ber. 11: 156-157. 9.

1436 Hymenoptera in America North of Mexico

Taxonomy: Gregg, 1945. Ent. Soc. Amer., Ann. 38: 529-546. —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 46: 147-148 (larva).

Biology: Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 77-89. —Benjamin, 1958.
Canad. Ent. 90: 419. —Kannowski, 1959. Insectes Sociaux 6: 135-1386. —Wheeler and
Wheeler, 1963. Ants of N. Dak., pp. 178-179. —Gregg, 1963. Ants of Colo., pp. 454-457.
—Burns, 1964. Ent. Soc. Amer., Ann. 57: 138 (association with tuliptree scale). —Smith,
1965. U. S. Dept. Agr., Tech. Bul. 1326, pp. 80-81 (economic importance). —Wheeler and
Wheeler, 1973. Ants of Deep Canyon, pp. 115-116.

Morphology: Regnier and Wilson, 1969. Jour. Ins. Physiology 15: 893-898 (alarm-defense
system).
crypticus Wilson. N. Dak. s. to N. Mex. w. to Alta., Idaho, Oreg., Utah; S. Calif. Ecology:
Abundant in prairie regions where it nests under stones or occasionally in crater nests
in soil.
Lasius (Lasius) crypticus Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113:
104-118. 3, 2, do.

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 179-182.
neoniger Emery. Que., Maine s. to Fla. w. to Idaho, Wyo., Colo., N. Mex.; Calif. (Sierras);
Alaska (?). Ecology: Nests almost exclusively in open areas, either under stones or in
crater nests. One of the dominant ants found in lawns, cultivated fields, grassy road
strips, and prairies. A common house and lawn pest and also fosters honeydew-excreting
insects. In American literature, neoniger has often been confused with alienus
(=americanus Emery), and many early references applying these names are here
referred to neoniger.
Lasius niger var. neoniger Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637. §.

Taxonomy: Gregg, 1945. Ent. Soc. Amer., Ann. 38: 534.

Biology: Forbes, 1894. 18th Rpt. State Ent. Ill, 171 pp. —Forbes, 1908. Ill. Agr. Expt. Sta.
Bul. 131: 31-44. —Tanquary, 1913. Ill. State Lab., Nat. Hist. Bul. 9: 417-443. —Metcalf and
Flint, 1939. Destructive and Useful Insects, pp. 371-374, 770. —Severin, 1920. S. Dak. State
Ent. Cir. 20: 3. —Talbot, 1945. Amer. Midland Nat. 34: 504-506. —Talbot, 1946. Ecology 27:
65-70. —Schread and Chapman, 1948. Conn. (State) Agr. Expt. Sta. Bul. 515: 4-11.
—Talbot, 1953. Mich. Univ., Lab. Vertebrate Biol., Contrib. No. 63, pp. 3-12. —Wilson,
1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 100-104. —Ayre, 1962. Canad. Jour. Zool.
40: 157-164 (parasitism). —Wheeler and Wheeler, 1963. Ants. of N. Dak., pp. 182-185.
—Gregg, 1963. Ants of Colo., pp. 461-465. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326,
pp. 81-84 (economic importance). :

niger (Linnaeus). Colo., N. Mex., Ariz., Utah, Mont., Idaho, Wash., Oreg., Calif.; Mexico;

Eurasia. Ecology: Found in forests or in open situations where it usually nests under
stones, though occasionally in rotting wood.

Formica nigra Linnaeus, 1758. Syst. Nat., Ed. 10, 1: 580. &.

Lasius niger var. alieno-niger Forel, 1874. Les Fourmis de la Suisse, pp. 47, 49. 8, ?.

Lasius niger var. alienoides Emery, 1891. Explor. Sci. Tunisie, Paris, p. 16. 8.

Lasius niger flavescens Forel, 1903. Mus. Zool. Acad. Imp. Sci. St. Petersburg, Ann. 8:
386-387. &.

Lasius niger emeryi Ruzsky, 1905. Schrift. Naturforsch.-Ges. Univ. Kasan 38: 313-314. 8.

Acanthomyops niger nitidus Kuznetzov-Ugamskij, 1927. Rev. Russ. d’Ent. 21: 188. &.

Acanthomyops niger alienus var. pilicornis Kuznetzov-Ugamskij, 1927. Rev. Russe d’Ent.
21: 189. &.

Acanthomyops niger var. minimus Kuznetzov-Ugamskij, 1928. Ants of South Ussuri
Region, U.S. S. R. Natl. Geog. Soc. Publ., p. 20. &.

Lasius emarginatus var. nigrescens Stitz, 1930. Mitt. Zool. Mus. Berlin 16: 240. 2.

Lasius niger coloratus Santschi, 1937 Soc. Ent. de Belg., Bul. 68: 387. %, 9.

Lasius emarginatus var. japonicus Santschi, 1941. Mitt. Schweiz. Ent. Ges., 18: 277-278. 9,
oe

Lasius transylvanica Roszler, 1943. Zool. Anz. 144: 44-46. 3, 6.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 148-150 (larva).

Superfamily FORMICOIDEA 1437

Biology: Goswald, 1932. Ztschr. Wiss. Zool. 142: 1-156 (in Germany). —Wilson, 1955. Harvard
Univ., Mus. Comp. Zool., Bul. 113: 72-75. —Wheeler and Wheeler, 1973. Ants of Deep
Canyon, pp. 116-117.

pallitarsis (Provancher). N. S., Que. w. to B. C., Alaska s. to N. Y., N. C., Mich., Wis., Minn., S.
Dak., N. Mex., Ariz., Nev., Calif.; Siberia. Ecology: Found mostly in forested areas
where it nests in rotting logs and stumps or under stones. Occasionally a house infesting
ant.

Formica pallitarsis Provancher, 1881. Nat. Canad. 12: 355. 3, d.
Lasius niger sitkaensis Pergande, 1900. Wash. Acad. Sci., Proc. 2: 519. %.

Taxonomy: Wheeler, 1917. Harvard Univ., Mus. Comp. Zool., Bul. 61: 18, 21. —Cole, 1942.
Amer. Midland Nat. 28: 374. —Francoeur and Beique, 1966. Canad. Ent. 98: 144
(Provancher types).

Biology: Wheeler, 1915. Psyche 22: 206. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool.,
Bul. 113: 45-47. —Medler, 1958. Ent. Soc. Wash., Proc. 60: 258 (flights). —Wheeler and
Wheeler, 1963. Ants of N. Dak., pp. 185-188. —Gregg, 1963. Ants of Colo., pp. 461-465.
—Corbet and Ayre, 1968. Canad. Field-Nat. 82: 230-231 (swarming and mating).
—Kannowski, 1969. Canad. Field-Nat. 83: 283-285 (nuptial flights). —Corbet and Ayre,
1969. Canad. Field-Nat. 83: 285-286 (swarming and mating). —Akre and Hill, 1973. Kans.
Ent. Soc., Jour. 46: 526-536 (behavior of a myrmecophilous beetle).

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome numbers).
sitiens Wilson. Colo., N. Mex., Ariz., Nev.; Mexico. Ecology: Found in the lower altitudinal
forest belts in mountains, 7000 to 8000 feet where it nests under stones in dry open
situations.
Lasius (Lasius) sitiens Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 108-111.
SES iGr

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 20 (Nev. Test Site).
Genus LASIUS Subgenus CAUTOLASIUS Wilson

Lasius subg. Cautolasius Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 13.
Type-species: Formica flava Fabricius. Orig. desig.
fallax Wilson. Mont., Wyo., Colo., Idaho, Utah, Ariz., Wash. Ecology: Colonies have been found

under stones in forest clearings.

Lasius (Cautolasius) fallax Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113:
130-133. 8, 2, do.

flavus (Fabricius). N. S., N. B., Que. s. to Ala. w. to Alta., Wash., Oreg., Calif.; Eurasia.

Ecology: A subterranean ant which nests in various situations but most often under
stones. Known to build mounds in parts of Eurasia. Workers may attend aphids on roots
of grasses.

Formica flava Fabricius, 1781. Species Insectorum 1: 491. 8.

Lasius brevicornis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637. 9, 2, d.

Lasius flavus‘myops Forel, 1894. Soc. Vaud. des Sci. Nat., Bul. 30: 12. 8.

Lasius flavus myops var. flavoides Forel, 1894. Soc. Vaud. des Sci. Nat., Bul. 30: 12. 8.

Lasius flavus var. fuscoides Ruzsky, 1902. Schrift. Naturforsch. Gesell. Univ. Kasan 38:
281.

Lasius flavus var. odoratus Ruzsky, 1902. Schrift. Naturforsch. Gesell. Univ. Kasan 38:
282-283. &.

Lasius flavus var. flavo-myops Forel, 1915. Mitt. Schweiz. Ent. Gesell. 12: 52.

Lasius (Formicina) brevicornis microps Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.
52: 526. &.

Lasius (Formicina) flavus claripennis Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:
Sale Oar a arOe

Formicina flava var. morbosa Bondroit, 1918. Soc. Ent. de France, Ann. 87: 28-29. 9, 2.

Lasius umbratus var. apennina Menozzi, 1924. Atti Soc. Nat. Mat. Modena 8: 15. 9.

Lasius umbratus ibericus Santschi, 1925. Eos 1: 349-350. &.

Lasius umbratus ibericus var. sancho Santschi, 1925. Eos 1: 350. &.

Lasius flavus var. olivacea Karawajew, 1926. Konowia 5: 194. &.

1438 Hymenoptera in America North of Mexico

Lasius (Chthonolasius) helveolus Cook, 1950. Ants of Calif., p. 327. °.
Lasius (Chthonolasius) helvus Cook, 1950. Ants of Calif., p. 326. %. In figs.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24: 68-69. —Buren, 1944. Iowa State Col., Jour.
Sci. 18: 296. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 152-154 (larva).
—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 396. —Goswald, 1932. Ztschr. Wiss.
Zool. 142: 1-156 (in Germany). —Wesson and Wesson, 1940. Amer. Midland Nat. 24: 100.
—Cole, 1942. Amer. Midland Nat. 28: 375. —Wheeler and Wheeler, 1944. N. Dak. Hist.
Quart. 11: 253. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 126-128.
—Waloff, 1957. Insectes Sociaux 4: 391-408 (effect of number of queens on their survival
and the development of first brood). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
189-191. —Gregg, 1963. Ants of Colo., pp. 465-469. —Marikovsky, 1965. Insectes Sociaux
12: 63. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 117.

nearcticus Wheeler. Que., Ont. s. to N. C., Tenn. w. to S. Dak., Wyo., Colo. Ecology:
Subterranean; prefers dense, moist woodlands here it nests in soil under rocks or fallen
logs.
Formica mellea Provancher, 1881. Nat. Canad. 12: 356. %. Preoce. by Say, 1836.
Lasius flavus nearcticus Wheeler, 1906. Psyche 13: 38. &.

Taxonomy: Cole, 1940. Amer. Midland Nat. 24: 68, 70. —Buren, 1944. Iowa State Col., Jour.
Sci. 18: 296, 297. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 151-152 (larva).
—Francoeur and Beique, 1966. Canad. Ent. 98: 144 (Provancher types).

Biology: Wesson and Wesson, 1940. Amer. Midland Nat. 24: 100. —Headley, 1943. Ohio Jour.
Sci. 43: 29. —Hicks, 1947. Canad. Ent. 79: 170-171 (unusual nesting site). —Wilson, 1955.
Harvard Univ., Mus. Comp. Zool., Bul. 113: 135-136. —Gregg, 1963. Ants of Colo., pp.
469-471.

Genus LASIUS Subgenus CHTHONOLASIUS Ruzsky

Chthonolasius Ruzsky, 1913. Arch. f. Naturgesch. 79: 59-61.
Type-species: Formica umbrata Nylander. Desig. by Emery, 1925.

Ants of this subgenus are more subterranean than those of the subgenus Lasius, and some
build large earthen mounds. Food is largely honeydew derived from subterranean plant lice and
mealybugs. Some forms are temporary parasites on ants of the subgenus Lasius.

atopus Cole. Calif. (3 mi. S. Leggett, Mendocino Co.). Ecology: The nest was found in dry soil
under a stone in an unshaded area.
Lasius (Chthonolasius) atopus Cole, 1958. Tenn. Acad. Sci., Jour. 33: 75-77. 8.
humilis Wheeler. Colo., N. Mex., Nev. Ecology: Colonies were found under stones in moist soil

of a meadow and in moist open woods.
Lasius (Formicina) humilis Wheeler, 1917. Amer. Acad. Arts and Sci., Proce. 52: 528. Genoe

Biology: Gregg, 1963. Ants of Colo., pp. 473-475.
minutus Emery. N. S., Maine s. to Va. w. to Minn., Iowa. Ecology: Most often fouiid in
sphagnum bogs, swampy meadows, or open dry forests. Taken most often in mounds or
masonry domes. Some specimens have been associated with L. alienus.
Lasius umbratus minutus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 37, 641. 8, 2, d.

Taxonomy: Wheeler, 1910. Psyche 17: 237, 238, 241-242.

Biology: Gaige, 1914. Univ. Mich., Mus. Zool., Occas. Papers 5: 3, 4, 21, 23. —Wheeler, 1915.
Psyche 22: 206. —Morris, 1943. Ind. Acad. Sci., Proc. 52: 215. —Wilson, 1955. Harvard
Univ., Mus. Comp. Zool., Bul. 113: 182. —Kannowski, 1959. Insectes Sociaux 6: 136-138,
151-153 (colony founding; possible parasitism on Lasius alienus and L. pallitarsis (=
sitkaensis)). —Kannowski, 1959. Ecology 40: 162-165 (radioactive phosphorus in study of
colony distribution)

Superfamily FORMICOIDEA 1439

nevadensis Cole. Nev. (Kyle Canyon, Charleston Mts.). Ecology: Nests were found in an
unshaded area in open forests; some were under stones and some were in exposed soil
with a scattering of soil around the entrance.
Lasius (Chthonolasius) nevadensis Cole, 1956. Tenn. Acad. Sci., Jour. 31: 26-27. 3, 2, d.
speculiventris Emery. N. J., Pa. w. to Minn., lowa, Kans. Ecology: Nests have been found
under rocks and in rotting wood either in pastures or wooded areas.
Lasius speculiventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 637, 641. 3, d.

Taxonomy: Wheeler, 1910. Psyche 17: 237, 242-243. —Wheeler, 1917. Conn. State Geol. and
Nat. Hist. Survey Bul. 22: 592.

Biology: Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 173. —Kannowski, 1959.
Insectes Sociaux 6: 138-141, 153-154 (parasitism on minutus and possible other Lasius
spp.)-

subumbratus Viereck. N. S., Maine w. to Sask., Wash., Oreg., s. to N. Mex., Ariz., Nev.
Ecology: Nests under stones or rotting logs in meadows and forests. A temporary social
parasite. Host: Lasius pallitarsis (Provancher).
Lasius umbratus subumbratus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 73. °.

Taxonomy: Wheeler, 1910. Psyche 17: 237-239. —Cole, 1942. Amer. Midland Nat. 28: 374.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 528. —Wheeler, 1917. Psyche
24: 167-176. —Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 179-180. —Wheeler
and Wheeler, 1963. Ants of N. Dak., pp. 191-192. —Gregg, 1963. Ants of Colo., pp. 475-478.

umbratus (Nylander). N. S., N. B., Que. s. to Fla. w. to Idaho, Utah, Ariz.; Eurasia. Ecology:

Prefers moist soil and most commonly nests under stones and in or under rotting logs
and stumps. May foster subterranean plant lice and mealybugs; occasionally a
house-infesting ant. A temporary social parasite. Host: Lasius alienus (Foerster), L.
niger (L.), L. neoniger Emery (7).

Formica umbrata Nylander, 1846. Acta. Soc. Sci. Fenn. 2: 1048-1050. 2, d.

Formica mixta Nylander, 1846. Acta. Soc. Sci. Fenn. 2: 1050-1052. 2.

Formica affinis Schenck, 1852. Jahrb. Ver. Nat. Nassau 8: 62-63. %, 2, d.

Formica aphidicola Walsh, 1862. Ent. Soc. Phila., Proc. 1: 310. %, d.

Lasuis umbratus var. mixto-umbratus Forel, 1874. Nouv. Mem. Soc. Helv. Sci. Nat., p. 48.
%.

Lasius umbratus var. exacutus Ruzsky, 1904. Kasan Univ. Obschchestvo
estestvoispytatelei Protokoly Zasiedanii, no. 206, p. 15. &.

Lasius umbratus var. mixto-affinis Ruzsky, 1904. Kasan Univ. Obschchestvo
estestvoispytatelei Protokoly Zasiedanii, no. 206, p. 15. Nomen nudum.

Lasius umbratus var. mixto-bicornis Ruzsky, 1905. Schrift. Naturforsch. Gesell. Univ.
Kasan 38: 292. Nomen nudum.

Lasius umbratus var. affino-umbratus Donisthorpe, 1914. Ent. Rec. 26: 40. 9.

Lasius umbratus var. przewalskii Ruzsky, 1915. Mus. Zool. Acad. Sci. Petrograd 20: 434.
om

Formicina umbrata distinguenda Emery, 1916. Rend. Accad. Bologna, pp. 64-65. 3, °.

Formicina umbrata var. hybrida Emery, 1916. Rend. Accad. Bologna, p. 66.

Formicina umbrata var. nuda Bondroit, 1917. Soc. Ent. de France, Bul. 86: 176.

Formicina umbrata var. sabularum Bondroit, 1918. Soc. Ent. de France, Bul. 87: 31.

Formicina belgarum Bondroit, 1918. Soc. Ent. de France, Bul. 87: 31. %, @.

Lasius bicornis var. citrina Emery, 1922. Soc. Ent. Ital., Bol. 54: 12. 8.

Lasius umbratus var. viehmeyeri Emery, 1922. Soc. Ent. Ital., Bol. 54: 13-15. 3, 2.

Lasius silvestrii Wheeler, 1928. Lab. Zool. Portici, Bol. 20: 120-121. °.

Lasius viehmeyeri var. dalmatica Starcke, 1937. Tijdschr. Ent. 80: 53-54. °.

Lasius umbratus var. hirtiscapus Starcke, 1937. Tijdschr. Ent. 80: 43. °.

Lasius umbratus distinguendus var. cereomicans Starcke, 1937. Tijdschr. Ent. 80: 48-49.
Be Gi

Lasius silvestri (!) var. osakana Santschi, 1941. Mitt. Schweiz. Ent. Ges. 18: 278. °.

Chtonolasius(!) affinis var. nyaradi Roszler, 1943. Zool. Anz. 144: 47-48. 3, °.

Lasius umbratus epinotalis Buren, 1944. Iowa State Col., Jour. Sci. 18: 296, 297. %.

1440 Hymenoptera in America North of Mexico

Taxonomy: Wheeler, 1910. Psyche 17: 237-241. —Wheeler, 1917. Conn. State Geol. and Nat.
Hist. Survey Bul. 22: 592. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 150-151
(larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 397. —Smith, 1928. Ent. News 39:
277-278. —Wesson and Wesson, 1940. Amer. Midland Nat. 24: 90, 101. —Wilson, 1955.
Harvard Univ., Mus. Comp. Zool., Bul. 113: 161-165. —Kannowski, 1959. Insectes Sociaux
6: 141, 154-155 (possible parasitism on other Lasius spp.). —Wheeler and Wheeler, 1963.
Ants of N. Dak., pp. 192-194. —Gregg, 1963. Ants of Colo., pp. 478-481. —Smith, 1965. U.
S. Dept. Agr., Tech. Bul. 1326, pp. 84-86 (economic importance).

Morphology: Blum, et. al, 1968. Ent. Soc. Amer., Ann. 61: 1354-1359 (terpenes in mandibular
glands). —Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).
vestitus Wheeler. B. C., Idaho, Oreg., Calif.
Lasius umbratus vestitus Wheeler, 1910. Psyche 17: 238, 242. 2.
Lasius (Chthonolasius) pilosus Smith, 1934. Ent. Soc. Amer., Ann. 27: 384. %.

Biology: Wilson, 1955. Harvard Univ., Mus. Comp. Zool., Bul. 113: 173-175.

Genus ACANTHOMYOPS Mayr

Acanthomyops Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 699.
Type-species: Formica clavigera Roger. Monotypic.

The ants of this exclusively North American genus nest in the soil, usually beneath objects,
and also in rotting logs and stumps. They are mostly subterranean in habit. The females and
workers have a characteristic citronella or lemon-verbena odor. Species of Acanthomyops are
sometimes of economic importance because they foster honeydew excreting insects and because
of their objectionable habit of nesting around foundations of buildings and beneath basement
floors. The winged forms are often mistaken for termites.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 638, 642. —Wheeler, 1916. Psyche 23:
170-172. —Buren, 1950. Ent. Soc. Wash., Proc. 52: 184-190. —Wing, 1968. Cornell Univ.,
Agr. Expt. Sta., Mem. 405, 173 pp. (also biological notes for each species).

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 154-156 (larvae).
—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 648 (larvae).

Biology: Kannowski, 1963. Pavia Univ., Symp. Genet. Biol. Ital. 12: 74-102 (flight activities).
—Talbot, 1963. Ecology 44: 549-555 (local distribution and flight activities).
arizonicus (Wheeler). S. Ariz. Ecology: Most collections have been from under stones.
Lasius (Acanthomyops) interjectus arizonicus Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 532. 3.
bureni Wing. Wis. (Comstock, Barron Co.).
Acanthomyops bureni Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 135-138.
Orne
californicus (Wheeler). S. Calif. Ecology: Colonies have been found in soil under stones.
Lasius (Acanthomyops) interjectus californicus Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 531. %, 9.
claviger (Roger). Mass., Ont., N. Y. s. to Fla. w. to Minn., Nebr., Kans., Miss. Ecology: Found
in woodlands, pastures, or open fields where they nest under stones, in rotting wood,
and occasionally in exposed soil. A common house pest. Smaller yellow ant.
Formica clavigera Roger, 1862. Berlin. Ent. Ztschr. 6: 241. ?.
Lasius (Acanthomyops) parvula Smith, 1934. Psyche 41: 213. %.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 296. —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 46: 155-156 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 398. —Dennis, 1938. Ent. Soc. Amer.,
Ann. 31: 296, 397. —Rau, 1945. Ent. News 56: 119. —Carter, 1962. Elisha Mitchell Sci. Soc.,
Jour. 78: 150-204. —Talbot, 1963. Ecology 44: 549-557 (local distribution, flight activities).

—Smith, 1965. U.S. Dept. Agr., Tech. Bul. 1326, pp. 86-87 (economic importance). —Talbot,
1973. Great Lakes Ent. 6: 20-21 (S. Mich.).

Superfamily FORMICOIDEA 1441

Morphology: Wheeler and McClendon, 1903. Biol. Bul. 4: 149-155. —Chadha, et al., 1962.
Jour. Ins. Physiol. 8: 175-179 (citronellal and citral in mandibular gland secretion).
—Regnier and Wilson, 1968. Jour. Ins. Physiol. 14: 955-970 (alarm-defense system).

colei Wing. N. Mex., Ariz.
Acanthomyops colei Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 88-89. %, d.
coloradensis (Wheeler). Man., N. Dak., S. Dak., Colo., N. Mex. w. to Alta., Oreg., Utah.
Ecology: Most colonies have been found under stones. Confused with A. claviger in
earlier literature.
Lasius (Acanthomyops) interjectus coloradensis Wheeler, 1917. Amer. Acad. Arts and Sci.,
PTOC25 270524. 9F) OO

Biology: Gregg, 1963. Ants of Colo., pp. 481, 483-484. —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 195-197 (reported as claviger).
creightoni Wing. Utah (Moab, Grand Co.).
Acanthomyops creightoni Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp.
141-143. 8, 2, d.
interjectus (Mayr). Mass., N. Y. s. to Ga. w. to Mont., Idaho, Utah, N. Mex. Ecology: Found in
woodlands, pastures or meadows. They may nest in exposed soil where the nest is
sometimes surmounted by a mound, under stones or other objects, in rotting logs and
stumps, or next to foundation walls of buildings. A frequent house pest. Larger yellow
ant.
Lasius (Acanthomyops) interjectus Mayr, 1866. Zool.-Bot. Gesell. Wien, Verh. 16: 888. ?.

Taxonomy: Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 592, 594.
—Buren, 1944. Iowa State Col., Jour. Sci. 18: 296. —Wheeler and Wheeler, 1953. Ent. Soc.
Amer., Ann. 46: 156 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 397-398. —Smith, 1928. Kans. Ent.
Soc., Jour. 1: 14-18. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 296, 306. —Carter, 1962.
Elisha Mitchell Sci. Soc., Jour. 88: 150-204. —Talbot, 1963. Ecology 44: 549-557 (local
distribution, flight activities). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 197-200.
—Gregg, 1963. Ants of Colo., pp. 484-486. —Smith, 1965. U. S. Dept. Agr., Tech. Bul. 1326,
pp. 88-89 (economic importance). —Talbot, 1973. Great Lakes Ent. 6: 22 (S. Mich.).

latipes (Walsh). Que., Maine w. to B. C. s. to S. C., Tenn., Ill., Iowa, Okla., N. Mex., Ariz., Calif.
Ecology: Found in open woodlands, meadows, or pastures where they may nest in
exposed soil commonly surmounted by a mound, under stones or other objects, or at the
base of stumps.
Formica latipes Walsh, 1862. Ent. Soc. Phila., Proc. 1: 311. 8, 9, d.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 638, 642. —Wheeler, 1917. Conn.
State Geol. and Nat. Hist. Survey Bul. 22: 592. —Buren, 1944. Iowa State Col., Jour. Sci.
18: 296. —Cole, 1954. Tenn. Acad. Sci., Jour. 29: 284. —Wing, 1968. Cornell Univ., Agr.
Expt. Sta, Mem. 405, pp. 98-105 (hybrids described).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 398. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 21: 398. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 41: 79. —Talbot,
1963. Ecology 44: 549-557 (local distribution, flight activities). —Wheeler and Wheeler,
1963. Ants of N. Dak., pp. 200-202. —Gregg, 1963. Ants of Colo., pp. 486-488. —Smith,
1965. U. S. Dept. Agr., Tech. Bul. 1326, pp. 91-93 (economic importance). —Cole, 1966.
Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 20 (Nev. Test Site). —Talbot, 1973. Great
Lakes Ent. 6: 22 (S. Mich).

Morphology: Wheeler and McClendon, 1903. Biol. Bul. 4: 149-155. —Wheeler, 1903. Amer.
Mus. Nat. Hist., Bul. 41: 79.
murphyi (Forel). N. Y., Ont. s. to Ga. w. to Sask., Idaho, n. Calif., Utah, N. Mex. Ecology:
Usually nests under or next to stones in open woodlands or edges of woodlands. Prefers
sandy soil. Widely but sporadically distributed in N. Amer.
Lasius (Acanthomyops) Murphii Forel, 1901. Soc. Ent. de Belg., Ann. 45: 367. 8, 2, d.
Lasius (Acanthomyops) murphyi Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 398.
Emend.

1442 Hymenoptera in America North of Mexico

Taxonomy: Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 13-121 (hybrids
described).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 398. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 530. —Talbot, 1963. Ecology 44: 549-557 (local distribution, flight
activities). —Gregg, 1963. Ants of Colo., pp. 489-490. —Smith, 1965. U. S. Dept. Agr., Tech.
Bul. 1326, pp. 89-91 (economic importance). —Talbot, 1973. Great Lakes Ent. 6: 21 (S.
Mich.).

occidentalis (Wheeler). Man., Minn., Nebr., Colo., N. Mex. w. to B. C., Wash., Wyo., Utah.
Ecology: Nests have been found under stones in dry sandy soil.
Lasius (Acanthomyops) occidentalis Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 83. %, 2, d.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 530. —Gregg, 1963. Ants of
Colo., pp. 490-493.
plumopilosus (Buren). N. C., Mich., Minn., lowa. Ecology: Found under stones and in rotting
logs.
Lasius (Acanthomyops) plumopilosus Buren, 1941. Iowa State Col., Jour. Sci. 15: 231-235.
OF Qe

Biology: Buren, 1944. Iowa State Col., Jour. Sci. 18: 296, 299.
pogonogynus (Buren). Iowa, Colo., Idaho. Listed as a hybrid of murphyi x latipes by Wing
(1968).
Lasius (Acanthomyops) pogonogynus Buren, 1950. Ent. Soc. Wash., Proc. 52: 186. %, 2.

Taxonomy: Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 117-119 (as a hybrid).

Biology: Gregg, 1963. Ants of Colo., pp. 493-494.
pubescens (Buren). Minn. Ecology: Nests with low mounds were found in soil in open woods.
Lasius (Acanthomyops) pubescens Buren, 1942. Iowa State Col., Jour. Sci. 16: 405. %, 9.
subglaber (Emery). Maine s. to Ga., Tenn. w. to Sask., N. Dak., S. Dak. Ecology: Nests in
woodlands or open areas, in mound nests, under stones, or in or under rotting logs and
stumps.
Lasius elvis var. swbglaber Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 642. 8, 2, d.
Lasius (Acanthomyops) clavigeroides Buren, 1942. Iowa State Col., Jour. Sci. 16: 406. &,
One

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 398. —Wheeler, 1917. Amer. Acad.
Arts and Sci., Proc. 52: 533. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 155
(larva). —Wing, 1968. Cornell Univ., Agr. Expt. Sta., Mem. 405, pp. 121-132 (hybrids
described). —Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 648, 649 (larva).

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 623. —Wheeler and Wheeler, 1963.
Ants of N. Dak., pp. 203-204. —Talbot, 1973. Great Lakes Ent. 6: 19-20 (S. Mich.).

Genus PARATRECHINA Motschulsky

Species of this genus often nest in the soil where the entrance may be surrounded by a small
irregular crater of earth, under stones, or beneath moss. They are occasionally house pests.
Because of the differences of opinion in recent literature regarding the status of subgenera, par-
ticularly Nylanderia, the two subgenera are listed as in previous catalogs.

Genus PARATRECHINA Subgenus PARATRECHINA Motschulsky

Paratrechina Motschulsky, 1863. Soc. Nat. Moscou, Bul. 36: 13.
Type-species: Paratrechina currens Motschulsky. Desig. by Wheeler, 1911.
longicornis (Latreille). S. C. s. to Fla. w. to Tex.; Calif.; Pantropical. Ecology: Highly adaptable
and may nest in many situations such as trash, refuse, plant cavities, rotting wood, and
in soil under stones. Workers are omnivorous and feed on live and dead insects, seeds,
honeydew and household foods. Sometimes a household pest. Introduced into the U. S.
Probably native to Africa or the Orient and spread to many parts of the world by

Superfamily FORMICOIDEA 1443

commerce. Sometimes found in northern states in greenhouses or other buildings. Crazy
ant.

Formica longicornis Latreille, 1802. Hist. Nat Fourmis, p. 113. 9.

Formica vagans Jerdon, 1851. Madras Jour. Lit. Sci. 17: 124. 3.

Formica gracilescens Nylander, 1856. Ann. Sci. Nat. Zool. 5: 73. 3.

Paratrechina currens Motschulsky, 1863. Soc. Nat. Moscou, Bul. 36: 14. %.

Taxonomy: Bingham, 1903. Fauna of British India 2: 326-327 (each caste). —Smith, 1936.
Puerto Rico Univ., Jour. Agr. 20: 865, 869-870. —Wheeler and Wheeler 1953. Ent. Soc.
Amer., Ann. 46: 143 (larva). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 87
(Polynesia).

Biology: Marlatt, 1928. U.S. Dept. Agr. Farmers’ Bul. 740: 6-7. —Phillips, 1934. (Hawaii
Univ.) Expt. Sta. Pineapple Prod. Coop. Assoc. Bul. 15: 18-19 —Turner, 1940. Conn. Agr.
Expt. Sta. Bul. 434: 311-312. —Fox and Garcia-Moll, 1961. Jour. Econ. Ent. 54: 1065-1066.
—Brown, 1964. Ent. News 75: 14-15. —Smith, 1965. U. S. Dept. Agr. Tech. Bul. 1326, pp.
74-76 (economic importance).

Genus PARATRECHINA Subgenus NYLANDERIA Emery

Nylanderia Emery, 1906. Soc. Ent. de Belg., Ann. 50: 134.
Type-species: Formica vividula Nylander. Orig. desig.

Members of this subgenus are in need of study. Existing keys are not adequate for species
identification, and a number of the names may be wrongly applied. Species determination has al-
ways been considered difficult and male genitalia are believed to offer help in separating spe-
cies. The segregates listed here are essentially as those proposed by Creighton, 1950. This sub-
genus is much better represented in the tropical regions of the world, and there are many tramp
species which make the taxonomy more confusing.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 402-410.

Biology: Smith, 1965. U. S. Dept. Agr. Tech. Bul. 1326, pp. 76-77 (as house pests).

bourbonica (Forel). S. C., Fla.; Pantropical. Introduced. Probably native to tropical Asia and
spread by commerce to Indian and Pacific Oceans and to New World tropics.
Prenolepis nodifera bourbonica Forel, 1886. Soc. Ent. de Belg., Ann. 30: 210. 8, 2, 6.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 405 (a var. of
bourbonica in U. S. ?). —Wilson and Taylor, 1967. Pacific Ins. Monog. 14: 87-89 (Polynesia).
Biology: Smith, 1930. Fla. Ent. 14: 23-24. —Wheeler, 1932. N. Y. Ent. Soc., Jour. 40: 16.
bruesii (Wheeler). Fla., Ala. w. to Okla., Tex., Ariz. (?).
Prenolepis bruesii Wheeler, 1903. Psyche 10: 106. 3, 2, d.
Taxonomy: Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 211 (larva; Okla; as
bruesi).
Biology: Smith, 1924. Ent. News 35: 122.
fulva (Mayr). S. Tex. s. to Argentina. Sporadically in greenhouses as far north as N. J.
Probably introduced. References to P. fulva pubens (Forel) from the U. S. pertain to
this species.
Prenolepis fulva Mayr, 1862. Zool.-Bot. Gesell. Wien, Verh. 12: 698. 3, 2.
Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636-637. —Creighton, 1950. Harvard
Univ., Mus. Comp. Zool., Bul. 104: 406-407.

Biology: Marlatt, 1922. U. S. Dept. Agr. Farmers’ Bul. 740: 8-9.
melanderi arenivaga (Wheeler). N. J. s. to Fla. w. to Iowa, Tex., Calif.
Prenolepis arenivaga Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 391. 8, d.

Prenolepis (Nylanderia) arenivaga var. faisonensis Forel, 1922. Rev. Suisse de Zool. 30:
98. &.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 295. —Harper, 1965. Calif. Dept. Agr.
Bul., Ann. Rpt. 45, 54: 81 (Calif. records).

Biology: Smith, 1928. Ent. News 39: 278.

1444 Hymenoptera in America North of Mexico

melanderi melanderi (Wheeler). Tenn. w. to Kans., Tex., Ariz.; Mexico.
Prenolepis melanderi Wheeler, 1903. Psyche 10: 104. %, 9, d.

Taxonomy: Emery, 1906. Soc. Ent. de Belg., Ann. 50: 132. —Wheeler and Wheeler, 1953. Ent.
Soc. Amer., Ann. 46: 144 (larva).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 74.
parvula (Mayr). Mass. s. to Fla. w. to N. Dak., Nebr., Kans., Tex., Ariz., Utah (?). Ecology:
Nests under moss, in logs and stumps, beneath stones, or in open grassy areas where
the nest may be surmounted by a small crater.
Prenolepis parvula Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 947. %, 2, d.
Prenolepis (Nylanderia) parvula var. grandula Forel, 1922. Rev. Suisse des Sci. Nat., Bul.
20: 348. &.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636 (worker, male). —Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 100. —Buren, 1944. Iowa State Col., Jour. Sci. 18:
295. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 211 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 390. —Talbot, 1934. Ecology 15:
420-422. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 295, 306. —Cole, 1940. Amer. Midland
Nat. 24: 66. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 175-176.

vividula guatemalensis (Forel). S. Ariz.; Central Amer. Possibly introduced. According to
Snelling (correspondence), the Ariz. record for guatemalensis is based on misidentified
arenivaga.
Prenolepis vividula vividula var. guatemalensis Forel, 1884. Soc. Vaud. des Sci. Nat., Bul.
20: 348. &.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 392 (worker, male).

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 342. —Cole, 1934. Ent. Soc. Amer.,
Ann. 27: 401.
vividula vividula (Nylander). Fla., Miss. Sporadically in greenhouses in northern areas as far

as Canada. Introduced.

Formica vividula Nylander, 1846. Acta Soc. Fenn. 2: 900. 8, 2, d.

Formica perminuta Buckley, 1866. Ent. Soc. Phila., Proc. 6: 162. §. Synonymy
questionable.

Formica picea Buckley, 1866. Ent. Soc. Phila., Proc. 6: 163. §. Synonymy questionable.

Formica (Tapinoma) terricola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 168. %, 2, d.
Synonymy questionable.

Taxonomy: Emery, 1906. Soc. Ent. de Belg., Ann. 50: 130 (each caste). —Emery, 1910. Deut.
Ent. Ztschr., p. 131 (each caste).

Biology: Mitchell and Pierce, 1912. Ent. Soc. Wash., Proc. 14: 74. —Marlatt, 1922. U. S. Dept.
Agr. Farmers’ Bul. 740: 7.

Genus PRENOLEPIS Mayr

Prenolepis Mayr, 1861. Die Europaischen Formiciden, pp. 26, 52.
Type-species: Tapinoma nitens Mayr. Desig. by Bingham, 1903.

These ants usually nest in the soil in exposed situations or under cover in small to moderate
sized colonies. Workers feed on honeydew, secretions of floral and extrafloral nectaries, exu-
dates from galls, earthworms, arthropods, and ripened or decaying fruits. The ants often invade
houses in search of food, and they sometimes forage even at freezing temperatures. Repletes
are common. Males and females overwinter in the nest and are among the first of ants to take
their nuptial flights in the spring.

Revision: Emery, 1898. Zool. Jahrb., Abt. f. System. 7: 635-637. —Wheeler, 1930. Ent. Soc.
Amer., Ann. 23: 1-26.
Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 142 (larvae).
imparis arizonica Wheeler. S. Ariz.
Prenolepis imparis var. arizonica Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 22. 3, 9, 6.

Superfamily FORMICOIDEA 1445

imparis californica Wheeler. Oreg., Nev., Calif. Ecology: Reported to feed on succulent tissue
of fruit and on fruit buds, damaging these by eating out the centers and leaving only
the scales.
Prenolepis imparis var. californica Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 23. %, 2, d.

Taxonomy: Eckert and Mallis, 1937. Calif. Agr. Expt. Sta. Bul. 342: 27.

Biology: Essig, 1926. Ins. West. No. Amer., p. 866. —Mallis, 1941. South. Calif. Acad. Sci.,
Bul. 40: 77-78.
imparis coloradensis Wheeler. Colo., N. Mex.
Prenolepis imparis var. coloradensis Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 22. 9.

Biology: Gregg, 1963. Ants of Colo., pp. 496-498.

imparis imparis (Say). Conn., Ont. s. to Fla. w. to Wis., Iowa, Mo., Okla., Tex., N. Mex., Ariz.

An annoying house infesting ant with habits similar to those given for the genus.

Formica imparis Say, 1836. Boston Jour. Nat. Hist. 1: 287. 9, d.

Tapinoma polita Smith, 1855. Ent. Soc. London, Trans. 3: 112. $. Synonymy questionable.

Formica (Tapinoma) Wichita Buckley, 1866. Ent. Soc. Phila., Proc. 6: 169. §. Synonymy
questionable.

Prenolepis nitens var. americana Forel, 1891. In Grandidier, Hist. Madagascar, v. 20, p.
94. 3.

Prenolepis imparis var. minuta Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636. 9, d.

Prenolepis imparis var. testacea Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 636. 2, d.

Prenolepis imparis var. pumila Wheeler, 1930. Ent. Soc. Amer., Ann. 23: 21. 8, d.

Taxonomy: Wesson and Wesson, 1940. Amer. Midland Nat. 24: 100. —Cole, 1940. Amer.
Midland Nat. 24: 66, 67. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 142-143
(larva).

Biology: Smith, 1924. Ent. News 35: 122. —Smith, 1928. Ent. News 39: 278. —Dennis, 1941.
Ent. Soc. Amer., Ann. 34: 82-86. —Talbot, 1943. Ecology 24: 31-44 (population studies).
—Talbot, 1943. Ecology 24: 345-352. (response to temperature and humidity changes).
—Tarpley, 1965. N. Y. Ent. Soc., Jour. 73: 6 (nuptial flight). —Smith, 1965. U. S. Dept.
Agr., Tech. Bul. 1326, pp. 78-79 (economic importance).

Genus MYRMECOCYSTUS Wesmael

Myrmecocystus Wesmael, 1838. Brussels Acad. Roy. de Belg., Bul. de Cl. des Sci. 5: 769.
Type-species: Myrmecocystus mexicanus Wesmael. Monotypic.

This genus is native to Mexico and western United States and is especially typical of the arid
plains and deserts. The ants nest in soil, the nest being surmounted by a small crater, usually in
medium-sized colonies of 1,000 to 3,000 workers. Some forms are apparently entirely predaceous
or carnivorous; others live on honeydew and nectar obtained from plants or the secretion of
galls. They may be diurnal or nocturnal in their activity. Repletes are known to occur in many
forms. Ants of this genus are commonly called “honey ants” because of the honeylike substance
stored in the gasters of the repletes. Fluid stored in repletes is largely for use by adults of the
colony with little or none for the larvae. Larval food consists of dead insects brought in by
foraging workers. Owing to the high degree of polymorphism in many forms, these ants are not
easily determined specifically without large series of workers, especially major ones.

Since this section was completed, Snelling (1976) published a revision of Myrmecocystus, the
results of which cannot be entirely incorporated into this catalog at this date. Snelling recognizes
27 species in three subgenera, the typical subgenus, subgenus Endiodioctes Snelling
(type-species: Myrmecocystus melliger Forel), and subgenus Eremmocystus Snelling
(type-species: Myrmecocystus creightoni Snelling). Except for their arrangement by subgenus,
all species treated by Snelling that fall within the scope of this catalog are given.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 666-667. —Wheeler, 1908. Amer. Mus.

Nat. Hist., Bul. 24: 345-397. —Wheeler, 1912. Psyche 19: 172-181. —Snelling, 1976. Nat.
Hist. Mus. of Los Angeles Co., Sci. Bul. 24: 1-163 (also biology).

Taxonomy: Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 1-9 (melliger complex).
—Wheeler and Wheeler, 1970. Ent. Soc. Amer., Ann. 63: 651 (larvae).

1446 Hymenoptera in America North of Mexico

Biology: Snelling, 1968. Los Angeles Co. Mus., Natur. Hist. Quart. 7: 14-18.
colei Snelling. S. Calif.
Myrmecocystus (Eremnocystus) colei Snelling, 1976. Nat. Hist. Mus. of Los Angeles Co.,
Sci. Bul. 24: 94-97. %, 2, do.
creightoni Snelling. S. Calif.
Myrmecocystus creightoni Snelling, 1971. Los Angeles Co. Mus., Contrib. Sci. 214: 6-11. %,
OER
Taxonomy: Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva; misidentified as
M. lugubris).
Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 120.
depilis Forel. W. Tex., N. Mex., Ariz., s. Nev.; Mexico. Confused with mimicus in the earlier
literature.
Myrmecocystus melliger var. depilis Forel, 1901. Soc. Ent. de Belg., Ann. 45: 135. 8.
Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 354. —Wheeler, 1912. Psyche 19:
173:
ewarti Snelling. S. Calif.

Myrmecocystus ewarti Snelling, 1971. Los Angeles Co. Mus., Contrib. Sci. 214: 2-6. 8, 2,
3.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 120.
flaviceps Wheeler. Utah, Ariz., s. Nev., s. Calif.; Mexico.
Myrmecocystus yuma var. flaviceps Wheeler, 1912. Psyche 19: 174, 177. %.

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 121-122.
hammettensis Cole. Idaho, Nev., e. central Calif.
Myrmecocystus hammettensis Cole, 1938. Amer. Midland Nat. 19: 678. %, 2, d.
kathjuli Snelling. S. Calif.
Myrmecocystus (Endiodioctes) kathjuli Snelling, 1976. Nat. Hist. Mus. of Los Angeles Co.,
Sci. Bul. 24: 59-62. 8, 2, o.
kennedyi Cole. S. Idaho, s.w. Utah, Ariz., s.e. Oreg., Nev., Calif.; Mexico. This species was
called M. semirufus in the literature prior to Snelling, 1969.
Myrmecocystus melliger semirufus var. kennedyi Cole, 1936. Ent. News 47: 119. ¥, 2, d.

Taxonomy: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 355, 368-369 (semirufus; also
biology). —Wheeler, 1912. Psyche 19: 174, 176 (semirufus; also biology). —Cole, 1938.
Amer. Midland Nat. 20: 371, 372. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61:
213 (larva; as semirufus). —Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 6.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 20 (semirufus). —Wheeler and
Wheeler, 1973. Ants of Deep Canyon, p. 122.
koso Snelling. S. Nev., s. Calif.
Myrmecocystus (Endiodioctes) koso Snelling, 1976. Nat. Hist. Mus. of Los Angeles Co.,
Sci. Bul. 24: 74-78. 8, 2, o.
lugubris Wheeler. S. Nev., s. Calif.
Myrmecocystus lugubris Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 98. &.

Taxonomy: Wheeler, 1912. Psyche 19: 174, 176. —Creighton, 1956. Amer. Mus. Novitates
1807: 1.

Biology: Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 21.
melliger Forel. W. Tex.; Mexico. Used as food and medicine by Indians.
Formica melligera Liave, 1832. Reg. Trim. o. Collect. Mem. Hist. Lit., p. 463. 8.
Questionable placement.
Myrmecocystus melliger Forel, 1886. Soc. Ent. de Belg., Ann. 30: 201. %.
Myrmecocystus melliger mendax var. comatus Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.
24: 352. 9, 9, od.

Superfamily FORMICOIDEA 1447

Taxonomy: Wheeler, 1912. Psyche 19: 173, 175. —Snelling, 1969. Los Angeles Co. Mus.,
Contrib. Sci. 170: 4. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 211-213
(larva).

mendax Wheeler. Tex., Colo., N. Mex., Ariz., s. Nev., s. Calif.; Mexico.

Myrmecocystus melliger mendax Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 351. 3, 2,
3

Myrmecocystus melliger orbiceps Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 349. 3, °.

Taxonomy: Wheeler, 1912. Psyche 19: 173. —Cole, 1954. Tenn. Acad. Sci., Jour. 29: 284.
—Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 2.

Biology: Wheeler, 1910. Ants, p. 376. —Cole, 1942. Amer. Midland Nat. 28: 386. —Gregg,
1963. Ants of Colo., pp. 643-645, 645-648 (misident. as comatus, p. 643-645).
mexicanus Wesmael. W. Tex., Colo., N. Mex., Utah, Ariz., Nev., Calif.; Mexico.

Myrmecocystus mexicanus Wesmael, 1838. Brussels Acad. Roy. de Belg., Bul. Cl. des Sci.
5: 770. 8.

Myrmecocystus Melliger var. hortus-deorum McCook, 1881. Acad. Nat. Sci. Phila., Proc.,
pp. 65, 69. %, replete 3, 2, d.

Taxonomy: Forel, 1886. Soc. Ent. de Belg., Ann. 30: 202. —Emery, 1893. Zool. Jahrb., Abt. f.
System. 7: 666. —Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 356-360. —Wheeler, 1912.
Psyche 19: 173, 178. —Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva).

Biology: McCook, 1881. Acad. Nat. Sci. Phila., Proc., pp. 17-77. —McCook, 1882. The Honey
Ants of the Garden of the Gods and the Occident Ants of the American Plains, pp. 17-74.
—Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 81. —Gregg, 1963. Ants of Colo., pp.
648-651. —Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 122-124.

mimicus Wheeler. S.W. Kans., w. Okla., w. Tex., N. Mex., Ariz., Calif.; Mexico.
Myrmecocystus melliger mimicus Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 353. 3,
Qiao

Myrmecocystus melliger mimicus var. jesuita Wheeler, 1908. Amer. Mus. Nat. Hist., Bul.
24: 354. 8.

Myrmecocystus melliger lomaensis Wheeler, 1912. Psyche 19: 174. ¥, replete §, 2.
Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 446. —Wheeler and
Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva).
Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27: 401-402. —Mallis, 1941. South. Calif. Acad. Sci.,
Bul. 40: 80. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 22 (Nev. Test Site).

—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 124-125. —Holldobler, 1976.
Science 192: 192-194 (tournaments and slavery).

navajo Wheeler. Colo., w. Tex., N. Mex., Utah, Ariz., s. Nev., s. Calif.
Myrmecocystus mexicanus navajo Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 360. &,
oF
Taxonomy: Wheeler, 1912. Psyche 19: 173, 179.
Biology: Cole, 1942. Amer. Midland Nat. 28: 386. —Gregg, 1963. Ants of Colo., pp. 651-653.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 125.
placodops Forel. Tex. N. Mex., Ariz.; Mexico.
Myrmecocystus melliger var. placodops Forel, 1908. Soc. Vaud. des Sci. Nat., Bul. 44: 70.
om
Taxonomy: Wheeler, 1912. Psyche 19: 173. —Snelling, 1969. Los Angeles Co. Mus., Contrib.
Sci. 170: 6.
pyramicus Smith. Idaho, s.e. Oreg., Nev.
Myrmecocystus pyramicus Smith, 1951. Great Basin Nat. 11: 91-94. &.
Taxonomy: Cole, 1957. N. Y. Ent. Soc., Jour. 65: 129-130 (female, male).

romainei Cole. W. Kans., w. Okla., w. Tex., Colo., N. Mex., Utah, Ariz., Nev.; Mexico.
Myrmecocystus melliger semirufus var. romainei Cole, 1936. Ent. News 47: 120. 9.

1448 Hymenoptera in America North of Mexico

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 285 (as semirufa, in part). —Gregg, 1963. Ants
of Colo., pp. 653-655 (as semirufus, in part).
semirufus Emery. S. Calif.; Mexico.
Myrmecocystus melliger var. semirufus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 667.
3.

Taxonomy: Snelling, 1969. Los Angeles Co. Mus., Contrib. Sci. 170: 4-6 (The species
previously considered to be M. semirufus by Wheeler and subsequent authors must now
be known as M. kennedy? Cole.).

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 125-126 (misident. as
placodops Forel).
tenuinodis Snelling. S. Nev., s.w. Ariz., s. Calif.; Mexico.
Myrmecocystus (Eremnocystus) tenuinodis Snelling, 1976. Nat. Hist. Mus. of Los Angeles
Co., Sci. Bul. 24: 107-110. 3, 9.
testaceus Emery. Idaho, Utah, Wash. Oreg., Nev., Calif.; Mexico.
Myrmecocystus melliger var. testaceus Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 667.
3.
Myrmecocystus mexicanus mojave Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 360. &.
Myrmecocystus mexicanus idahoensis Cole, 1936. Ent. News 47: 118. %, 2, 3.

Taxonomy: Wheeler, 1912. Psyche 19: 173, 179-181. —Snelling, 1969. Los Angeles Co. Mus.,
Contrib. Sci. 170: 6. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 213 (larva).

Biology: Wheeler, 1910. Ants, p. 349. —Leonard, 1911. San Diego Soc. Nat. Hist., Trans. 1:
91-92. —Cole, 1934. Ent. Soc. Amer., Ann. 27: 403. —Mallis, 1941. South. Calif. Acad. Sci.,
Bul. 40: 81. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 22 (Nev. Test Site).
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 126-127.

wheeleri Snelling. S. Calif.
Myrmecocystus wheeleri Snelling, 1971. Los Angeles Co. Mus., Contrib. Sci. 214: 11-15. 3,
2G:

Biology: Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 127.
yuma Wheeler. W. Ariz., s. Nev., s. Calif.; Mexico.
Myrmecocystus yuma Wheeler, 1912. Psyche 19: 174, 176. &.

Taxonomy: Creighton, 1956. Amer. Mus. Novitates 1087: 1.
TRIBE FORMICINI

Genus FORMICA Linnaeus

Formica Linnaeus, 1758. Syst. Nat., Ed. 10, v. 1, p. 579.
Type-species: Formica rufa Linnaeus. Desig. by Curtis, 1839.
Serviformica Forel, 1913. Soc. Ent. de Belg., Ann. 57: 361.
Type-species: Formica fusca Linnaeus. Orig. desig.
Raptiformica Forel, 1913. Soc. Ent. de Belg., Ann. 57: 361.
Type-species: Formica sanguinea Latreille. Orig. desig.
Formica subg. Neoformica Wheeler, 1913. N. Y. Acad. Sci., Ann. 23: 82.
Type-species: Formica pallide-fulva Latreille. Desig. by Wheeler, 1913.
Coptoformica Mueller, 1923. Soc. Adriat. di Sci. Nat. Trieste, Bol. 28: 133.
Type-species: Formica exsecta Nylander. Desig. by Donisthorpe, 1941.
Adformica Lomnicki, 1925. Polskie Pismo Ent. 3: 164.
Type-species: Formica exsecta Nylander. Desig. by Donisthorpe, 1941.

This is a large and complex holarctic genus, and the largest genus of ants in America north of
Mexico. Various habit patterns are found in this genus including slave-making, temporary social
parasitism of various types, and various methods of nest founding and nest construction. Sub-
genera are not recognized in the following listing, only species groups which more or less reflect
the groupings by Creighton (1950) and Buren (1968). Proformica, apparently a valid genus found
in Europe, is deleted from the North American fauna, and those species previously listed in that

Superfamily FORMICOIDEA 1449

subgenus along with a number of species excluded from Raptiformica by Buren are here listed
in the species group neogagates. Those species included in the subgenus Raptiformica by Buren
(1968) are listed here in the species group sanguinea. The species previously included in the
subgenus Neoformica are listed here in the pallidefulva species group. The other species
groups, microgyna, exsecta, fusca, and rufa, are essentially the same as before. Further studies
may clarify the species groupings and determine the necessity of establishing subgenera.

Revision: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643-654, 657-665. —Wheeler, 1913.
Harvard Univ., Mus. Comp. Zool., Bul. 53: 387-399, 401-536, 560-565. —Creighton, 1940.
Amer. Mus. Novitates 1055: 1-10 (rwfa subspp.). —Creighton, 1950. Harvard Univ., Mus.
Comp. Zool., Bul. 104:450-552. —Francoeur, 1973. Ent. Soc. du Quebec, Mem. 3, 316 pp.
(fusca group).

Taxonomy: Yarrow, 1950. Internatl. Comm. Zool. Nomencl., Bul. Zool. Nomencl. 4: 408.
—Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 160 (larvae). —Yarrow, 1955.
Internatl. Comm. Zool. Nomencl., Bul. Zool. Nomencl. 9: 313-318 (type-species). —Wilson
and Brown, 1955. Psyche 62: 108-129 (notes on sanguinea and neogagates groups).
—Gregg, 1964. Colo. Univ. Mus., Leaflet No. 13, pp. 59-69 (distribution in mountains of
Colo.). —Gregg, 1969. Ent. Soc. Wash., Proc. 71: 38-49 (geographical distribution of genus
in world). —Buren, 1968. Ga. Ent. Soc., Jour. 3: 25-40 (fundamental taxonomic problems;
subg. Raptiformica).

Biology: Gosswald, 1957. Insectes Sociaux 4: 335-348 (caste determination). —Kannowski,
1959. Insectes Sociaux 6: 147. —Kannowski and Johnson, 1969. Anim. Behaviour 17:
425:429 (male patrolling behavior and sex attraction). —Rosengren, 1971. Acta Zool.
Fennica 133, 106 pp. (route fidelity, visual memory, recruitment behavior).

Morphology: Osman and Kloft, 1961. Insectes Sociaux 8: 383-395 (venom).
SPECIES GROUP NEOGAGATES

Species of this group nest in small colonies in the soil, sometimes under stones or beneath
other objects. They are commonly enslaved by members of the microgyna and sanguinea
groups.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 108-129. —Buren, 1968. Ga. Ent. Soc., Jour.

3: 25-40.
bradleyi Wheeler. Man., Minn., Iowa w. to Alta., Mont., Wyo., Colo. Ecology: Nests only in
sand or sandy soil among grass clumps or with low irregular mounds.
Formica bradleyi Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 389, A23. 9, 6.
Formica (Proformica) neogagates neogagates var. morbida Wheeler, 1913. Harvard Univ.,
Mus. Comp. Zool., Bul. 53: 400, 538. 9, 2.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 535. —Wheeler and Wheeler,
1953. Ent. Soc. Amer., Ann. 46: 168-169 (larva). —Wilson and Brown, 1955. Psyche 62:
126-127.

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 258-259. —Beamer and
Michner, 1950. Kans. Ent. Soc., Jour. 23: 110-113 (relationship with leafhoppers).
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 206-209. —Gregg, 1963. Ants of Colo.,
pp. 602-606. —Snelling, 1969. Ent. Soc. Wash., Proc. 71: 196-197. —Halverson, et al. 1976.
Kans. Ent. Soc., Jour. 49: 280-303 (natural history of the sandhill ant).

lasioides Emery. N. S., Que. w. to B. C. s. to Mass., Mich., S. Dak., Colo., N. Mex., Ariz., Calif.
Ecology: Commonly found in grasslands where it nests under stones or in nests with
exposed entrances or small craters, but also found in other habitats.
Formica lasiodes Emery, 1893. Zool. Jahrb., Abt. f. System 7: 646, 664. °.
Formica lasoides var. picea Emery, 1895. Zool. Jahrb., Abt. f. System. 8: 335. %. Preoce.

by Nylander, 1846.
Formica lasioides var. vetula Wheeler, 1912. Psyche 19: 90. N. name for picea Emery.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 400, 540-541 (each
caste). —Gregg, 1944. Ent. Soc. Amer., Ann. 37: 472. —Buren, 1944. Iowa State Col., Jour.

Sci. 18: 299.

1450 Hymenoptera in America North of Mexico

Biology: Cole, 1942. Amer. Midland Nat. 28: 384. —Wheeler and Wheeler, 1944. N. Dak. Hist.
Quart. 11: 268-269. —Kincaid, 1963. Amer. Micros. Soc., Trans. 82: 101-105 (as pollinators of
plants). —Wheeler and Wheeler, 1963. Ants of N. Dak. pp. 209-211, 502. —Gregg, 1963.
Ants of Colo., pp. 498-500, 502. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7:
23 (Nev. Test Site).

limata Wheeler. Minn., N. Dak., Colo., N. Mex., Utah, Nev. Ecology: Commonly nests in
grasslands under stones or in crater nests; tolerant of dry situations.
Formica (Proformica) limata Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
400, 541. 8.

Biology: Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 268. —Gregg, 1946. Amer.
Midland Nat. 35: 750. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 211-212. —Gregg,
1963. Ants of Colo., pp. 502, 504. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7:
24 (Nev. Test Site).

manni Wheeler. Idaho, Utah, Wash., Oreg., Nev., Calif. Ecology: Nests are often under stones
in gravelly or sandy soil of desert areas.
Formica manni Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 389, 420. 3, °.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 127-128.

Biology: Cole, 1942. Amer. Midland Nat. 28: 376, 378-379.
neogagates Emery. N. S., Que. w. to Alaska s. to N. C., Ill., Iowa, Nebr., N. Mex., Ariz., Calif.

Ecology: Nests most often in grasslands in dry and stony situations, under stones or in
the open with or without an irregular mound or crater.

Formica fusca subpolita var. neogagates Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646,
6615 959).

Formica (Proformica) neogagates neogagates var. vinculans Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool., Bul. 53: 400, 539. ¥, @.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 400, 536-538 (each
caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 596. —Wheeler
and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 158-159 (larva).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 401. —Wesson and Wesson, 1940.
Amer. Midland Nat. 24: 102. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 268.
—Buren, 1944. Iowa State Col., Jour. Sci. 18: 299. —Wheeler and Wheeler, 1963. Ants of
N. Dak., pp. 212-214. —Gregg, 1963. Ants of Colo., pp. 503-505. —Cole, 1966. Brigham
Young Univ., Sci. Bul., Biol. Ser. 7: 26 (Nev. Test Site).

obtusopilosa Emery. Minn. w. to Alta. s. to Nebr., N. Mex., Utah, Nev. Ecology: Small colonies
are found under stones or in exposed soil usually with irregular mounds or craters in

meadows and grasslands.
Formica sanguinea obtusopilosa Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 648. &.

Formica munda Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 267. 8, 2.
Formica munda var. alticola Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 534. $.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 389, 414-418. — Wilson
and Brown, 1955. Psyche 62: 128.

Biology: Cole, 1942. Amer. Midland Nat. 28: 378. —Wheeler and Wheeler, 1963. Ants of N.
Dak., pp. 217-219. —Gregg, 1963. Ants of Colo., pp. 608-611. —Cole, 1966. Brigham Young
Univ., Sci. Bul., Biol. Ser. 7: 26 (Nev. Test Site).

oregonensis Cole. Oreg. (Pendleton).
Formica oregonensis Cole, 1938. Amer. Midland Nat. 20: 368. &.

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31: 212-214.

Biology: Kincaid, 1963. Amer. Micros. Soc., Trans. 82: 101-105 (pollinators of plants).
perpilosa Wheeler. Wyo., Colo., Kans., Okla., Tex. w. to Calif.; Mexico. Ecology: Usually

constructs crater nests in the soil in grasslands and open fields.
Formica fusca subpolita var. perpilosa Wheeler, 1902. Soc. Cient. “Antonio Alzate” Mem.

y Rev. 17: 141. 3, 9°.

Superfamily FORMICOIDEA 1451

Taxonomy: Wheeler, 19138. Harvard Univ., Mus. Comp. Zool., Bul. 53: 389, 421-423 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376-378. —Wilson and Brown, 1955. Psyche
62: 128.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 534. —Cole, 1934. Ent. Soc.
Amer., Ann. 27: 401. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 85. —LaBerge, 1952.
Kans. Ent. Soc., Jour. 25: 59. —Gregg, 1963. Ants of Colo., pp. 611-613.

SPECIES GROUP PALLIDEFULVA

Members of this group nest in the soil under stones or at the base of tufts of grass. There is
usually no mound or crater marking the nest. They serve as hosts of slave-making species of
Formica.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 543-552 (subg.
Neoformica).
archboldi Smith. Ga., Ala., Fla.
Formica pallide-fulva archboldi Smith, 1944. Fla. Ent. 27: 16. 9.

Biology: Schneirla, 1944. Amer. Mus. Novitates 1261: 1-2.
pallidefulva nitidiventris Emery. Ont., Que. s. to Ga. w. to Wis., S. Dak., Wyo., Colo., N. Mex.

Formica pallide-fulva nitidiventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 656.
Oe OS:

Formica pallide-fulva schaufussi var. incerta Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
645, 655. 8, 2, d.

Formica pallide-fulva fuscata Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 656. &.

Formica pallide-fulva delicata Cole, 1938. Amer. Midland Nat. 20: 369. %, 9.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 401. —Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool., Bul. 53: 401, 554-557. —Wheeler, 1917. Conn. State Geol. and Nat.
Hist. Survey Bul. 22: 595, 598. —Buren, 1944. Iowa State Col., Jour. Sci. 18: 299, 309-310.
—Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 159-160 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 370-371. —Wheeler, 1917. Ind. Acad.
Sci., Proc. 26: 465. —Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 211-212. —Wesson and
Wesson, 1940. Amer. Midland Nat. 24: 102. —Talbot, 1946. Ecology 27: 65-70. —Schread
and Chapman, 1948. Conn. (State) Agr. Expt. Sta. Bul. 515: 18. —Talbot, 1948. Ecology 29:
316-325. —Gregg, 1963. Ants of Colo., pp. 628-630.

pallidefulva pallidefulva Latreille. N. Y., N. J. s. to Fla. w. to Colo., Okla., Tex.
Formica pallide-fulva Latreille, 1802. Hist. Nat. Fourmis, p. 174. &.
Formica pallide-fulva var. succinea Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 369. 8.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 656-657. — Wheeler, 1913.
Harvard Univ., Mus. Comp. Zool., Bul. 53: 401, 548-552.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 298, 306. —Cole, 1940. Amer. Midland Nat.
24: 81. —Schneirla, 1944. Amer Mus. Novitates 1261: 2-3. —Gregg, 1963. Ants of Colo., pp.
625-628.

schaufussi dolosa Wheeler. Va. s. to Fla. w. to Iowa, Colo., Tex.
Formica pallide-fulva schaufussi var. meridionalis Wheeler, 1904. Amer. Mus. Nat. Hist.,
Bul. 20: 370. 3. Preoce. by Ruzsky, 1889.
Formica pallide-fulva schaufussi var. dolosa Wheeler, 1912. Psyche 19: 90. N. name for
meridionalis Wheeler.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 401, 554 (worker,
female). —Cole, 1940. Amer. Midland Nat. 24: 73, 79. —Buren, 1944. Iowa State Col., Jour.
Sci. 18: 299.

Biology: Dennis, 1938. Ent. Soc. Amer., Ann. 31: 299, 306. —Gregg, 1963. Ants of Colo., pp.
630-632.

schaufussi schaufussi Mayr. Ont. s. to N. C., Tenn. w. to Wis., Iowa.
Formica Schaufussi Mayr, 1886. Akad. der Wiss. Wien, Math.-Nat. K1. Sitzber. 53: 493. 9.

1452 Hymenoptera in America North of Mexico

Taxonomy: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 370. —Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool, Bul. 53: 401, 552-553 (each caste). —Cole, 1940. Amer. Midland
Nat. 24: 73.

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 400. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bul. 22: 598. —Dennis, 1938. Ent. Soc. Amer., Ann. 31: 298,
306.

SPECIES GROUP FUSCA

Members of this group nest in the soil, and the nest is commonly started under objects or at
the base of tufts of grass. Excavated soil may be spread out in the form of an irregular crater or
low mound about the nest openings. Because of their ubiquity and usual docility, they are com-
mon hosts for many of the slave-making species of Formica.

Revision: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, 316 pp.
accreta Francoeur. B. C., Mont., Idaho, Wash., Oreg., Calif.

Formica accreta Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 182-189. 8, 2, do.
aerata Francoeur. Oreg., Nev., Calif. Ecology: Found in sandy soil where it nests under rocks.
Formica aerata Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 116-122. 3, 2.

altipetens Wheeler. Mont., Wyo., Colo., N. Mex., Idaho, Utah, Ariz., Calif. Ecology: Nests in
forested and open areas in the soil; low mounds are sometimes built, but nests are also
under objects.
Formica cinerea cinerea var. altipetens Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bul. 53: 399, 523. 3, 9, d.
Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 52-60.
Biology: Gregg, 1963. Ants of Colo., pp. 507-509. —Funk, 1975. Lepidop. Soc., Jour. 29:
261-262 (association with Lycaenidae).
argentea Wheeler. Que. w. to B. C. s. to S. C., Ohio, Ill., Iowa, S. Dak., N. Mex., Ariz., Calif.
Ecology: Found in open or semi-open situations usually in sandy soil under rocks or

with a low mound.
Formica fusca var. argentata Wheeler, 1902. Amer. Nat. 36: 952. %. Preocc. by Fabricius,

1804.

Formica fusca var. argentea Wheeler, 1912. Psyche 19: 90. N. name for argentata
Wheeler.

Formica fusca var. blanda Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 398,
510. @.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 141-152 (also ecology).

Biology: Gregg, 1963. Ants of Colo., pp. 521-525.
canadensis Santschi. Man., Sask., N. Dak., S. Dak., Kans., Alta., Wyo., Colo., N. Mex., Idaho,
Utah, Ariz., Calif. Ecology: Nests in open or wooded areas in the soil, sometimes with a
low mound. This species has been confused with lepida in much of the literature.
Formica cinerea var. canadensis Santschi, 1913. Soc. Ent. de Belg., Ann. 57: 435. 3, 2.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 60-67.

foreliana Wheeler. S. Ariz.
Formica foreliana Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 391, 451. 8.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 234-238.
fusca Linnaeus. Newfoundland (insular) w. to Yukon s. to N. C., Ill., Iowa, S. Dak., N. Mex.,

Ariz., Calif.; Holarctic. Ecology: Nests in a wide variety of situations, in forests or open
areas, under rocks, logs, in soil, or in rotting wood. Two forms are recognized by
Francoeur (1973), form swbaenescens which is mostly eastern and form marcida which
is mostly western in distribution.

Formica fusca Linnaeus, 1758. Syst. Nat., Ed. 10, v. 1, p. 580.

Formica fusca var. subaenescens Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646,
659-660. 3, °.

Formica fusca fusca var. marcida Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul.
53: 398, 503. 3, 2.

Superfamily FORMICOIDEA 1453

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 532. —Wheeler and
Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 164-165 (larva). —Yarrow, 1954. Soc. Brit. Ent.,
Trans. 11: 229-244. —Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 189-201.

Biology: King, 1949. Iowa Acad. Sci., Proc. 56: 367-370 (in mixed colony with Formica reflexa
Buren). —King, 1951. Iowa Acad. Sci., Proc. 58: 487-489. — Kannowski, 1959. Insectes
Sociaux 6: 141-142. —Wallis, 1960. Insectes Sociaux 7: 187-190 (spinning movements of
larvae). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 264-268. —Gregg, 1963. Ants of
Colo., pp. 514-525. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 23 (Nev. Test
Site).

glacialis Wheeler. Newfoundland (insular) s. to N. Y. w. to Man., N. Dak., Ill. Ecology: Found
in woods or open fields where it builds its nests in the soil and with a low mound
commonly covered with grass. This species has been confused with fusca L. in much of
the literature.
Formica fusca var. glacialis Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 624. 3, 9, d.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 151-161 (also ecology).
gnava Buckley. Tex., Colo., N. Mex., Utah, Ariz., Nev., Calif.; Mexico. Ecology: Found in desert
and semi-desert areas or open woods where it nests in the soil, normally under rocks.
Formica gnava Buckley, 1866. Ent. Soc. Phila., Proc. 6: 156. 3, 9, d.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 390, 518-521 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 377, 380, 383. —Francoeur, 1973. Soc. Ent. du
Quebec, Mem. no. 3, pp. 238-245.

Biology: Wheeler, 1902. Tex. Acad. Sci., Trans. 4: 20. —Wheeler, 1917. Amer. Acad. Arts and
Sci., Proc. 52: 541, 550. —Cole, 1934. Ent. Soc. Amer., Ann. 27: 388. —Mallis, 1941. South.
Calif. Acad. Sci., Bul. 40: 86. —Gregg, 1963. Ants of Colo., pp. 539-541.

hewitti Wheeler. Que., Maine w. to B. C. s. to Minn., N. Mex., Utah, Calif. Ecology: Found in
open or semi-open woods and forests where it nests in the soil under rocks or in rotting
wood.
Formica hewitti Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 552. 3, 2, d.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 208-215.

Biology: Gregg, 1963. Ants of Colo., pp. 526-527.
lepida Wheeler. Calif. Has been found only in the north coastal region.
Formica cinerea cinerea var. lepida Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul.
53: 399, 526. %.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 122-125.

longipilosa Francoeur. Calif. (Mendocino Co.).
Formica longipilosa Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 125-128. 9.
microphthalma Francoeur. Calif. Ecology: In mountains; may construct small crater nests in
the soil.
Formica microphthalma Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 201-207. 8,
é.
moki Wheeler. Utah, Ariz., Nev., Calif. Ecology: Nests in soil under rocks; the nest is
unmarked and the entrance is usually a hole near a bush or tree. Francoeur (1973) chose
to use the name occidua for this species because occidwa has been used more in the
literature; however, the case has not been referred to the Commission.
Formica moki Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 343. ?.
Formica rufibarbis var. occidentalis Wheeler, 1910. Ants, p. 570. Nomen nudum. Preocc.
by Buckley, 1866.
Formica rufibarbis var. occidua Wheeler, 1912. Psyche 19: 90. Nomen nudum. N. name for
occidentalis Wheeler.
Formica rufibarbis var. occidua Wheeler, 1913. Harvard Univ., Mus. Comp. Zool. Bul. 53:
398, 517. 3, 2.

Taxonomy: Smith, 1939. Ent. Soc. Amer., Ann. 32: 582-583. —Cole, 1943. Amer. Midland Nat.
29: 183. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 24-25 (male, also

1454 Hymenoptera in America North of Mexico

biological notes). —Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 254-259 (as
occidua).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 550. —Eckert and Mallis, 1937.
Calif. Agr. Expt. Sta. Cir. 342: 30. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 86.
—Cole, 1942. Amer. Midland Nat. 28: 385. —Wenner, 1959. Amer. Midland Nat. 62: 174.
—Wheeler and Wheeler, 1973. Ants of Deep Canyon, pp. 128-129.

montana Emery. Ohio, Wis., Ill., Minn., Iowa, Man., N. Dak., S. Dak., Nebr., Kans., Colo.

Ecology: A prairie species; nests in earthen mounds, often in natural hummocks,
commonly covered with grass. Sometimes uses thatching.

Formica fusca subpolita var. ? montana Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646,
663. 8.

Formica cinerea var. neocinerea Wheeler, 1910. Ants, p. 571. Nomen nudum.

Formica cinerea var. neocinerea Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
524. 8, 2, do.

Formica cinerea cinerea var. rutilans Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bul. 53: 399, 525. 3.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 67-74.

Biology: Wheeler, 1902. Amer. Nat. 36: 948-952. —Wheeler, 1910. Ants, pp. 201, 203, 460-461,
475. —Wheeler, 1917. Psyche 22: 206. —Amstutz, 1943. Ohio Jour. Sci. 43: 172. —Gregg,
1948. Ent. Soc. Wash., Proc. 50: 183-186. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 263-264 (also as altipetens, pp. 260-261). —Gregg, 1963. Ants of Colo., pp. 513-514.

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 455 (chromosome number).
neoclara Emery. N. Dak., S. Dak., Iowa, Kans., Tex. w. and northwest to N. W. T., B. C.,
Wash., Oreg., Calif. Ecology: Found in grasslands or open woods where it nests in the
soil, usually preferring sandy soil; nests are sometimes at the base of plants and
sometimes have loose mounds of vegetable debris or excavated soil.
Formica fusca var. neoclara Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646, 661. &.
Formica fusca pruinosa Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 548. 8, 2, d.

Formica fusca pruinosa var. lutescens Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52:
549. 8.

Taxonomy: Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 165 (larva). —Francoeur,
1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 84-94.

Biology: Wheeler, 1910. Ants, pp. 201, 460-461, 463. —LaBerge, 1952. Kans. Ent. Soc., Jour.
25: 59. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 268-271. —Gregg, 1963. Ants of
Colo., pp. 526, 528-530 (also as pruniosa, pp. 537-539).

neorufibarbis Emery. Newfoundland (Labrador) w. to Alaska s. to Mass., Mich., Minn., S. Dak.,
N. Mex., Ariz., Calif. Ecology: A dominant ant in the boreal and alpine forests of North
America; usually nests in rotting wood but occasionally in the soil under rocks.
Formica fusca var. neorufibarbis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646, 660. %.
Formica fusca var. algida Wheeler, 1915. Psyche 22: 205. ¥, 2.

Formica fusca fusca var. gelida Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
399, 505. ¥, 2, d.

Taxonomy: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 546-547. —Wheeler and
Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 165 (larva). —Francoeur, 1973. Soc. Ent. du
Quebec, Mem. no. 3, pp. 215-228.

Biology: Brown, 1955. Ent. News 66: 47-50. —Kannowski, 1959. Insectes Sociaux 6: 142.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 271-273. —Gregg, 1963. Ants of Colo.,
pp. 530-536. —Cole, 1966. Brigham Young Univ, Sci. Bul., Biol. Ser. 7: 26 (Nev. Test Site).
occulta Francoeur. Wyo., Colo., N. Mex., Utah, Ane Oreg.

Formica occulta Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 94-101. ¥, 2, d.
pacifica Francoeur. B. C., Wash., Oreg., Calif.

Formica pacifica Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 131-135. ¥, @.

Superfamily FORMICOIDEA 1455

pilicornis Emery. Calif.; Mexico. Ecology: Found at low elevations in the coast ranges north to
the San Francisco area; makes crater or mound nests in sandy soil, sometimes under
rocks.
Formica pilicornis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 646, 664. 3, 9, d.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 74-80.

Biology: Cole, 1934. Ent. Soc. Amer., Ann. 27: 401. —Eckert and Mallis, 1937. Calif. Agr.
Expt. Sta. Cir. 342: 7, 29. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 85.
podzolica Francoeur. N. S., Que. w. to Alaska s. to Pa., Wis., Iowa, S. Dak., N. Mex., Ariz.,
Calif. Ecology: A species of the boreal and alpine forests of North America. Nests are
in the soil, commonly sandy soil on beaches or shores and are craterlike or moundlike.
Has been confused with F’. fusca L. in the literature.
Formica podzolica Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 167-172. 3, 2, d.
sibylla Wheeler. Oreg., Nev., Calif. Ecology: Makes crater nests in the soil, usually in
woodlands.
Formica sibylla Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 399, 530. 8, 2.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 263-268.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 551. —Mallis, 1941. South. Calif.
Acad Sci., Bul. 40: 87.
subelongata Francoeur. Calif. (Burney Junction, Shasta Co.).
Formica subelongata Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 268-270. 8.
subpolita Mayr. B. C., Idaho, Wash., Oreg., Nev., Calif. Ecology: Usually found in semi-desert
areas where it nests in sandy or gravelly soil; nests may be craterlike or moundlike.
Formica fusca var. subpolita Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 426. 9, 2.
Formica rufiventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 665. d. Preoce. by
Fabricius, 1804.
Formica flammiventris Wheeler, 1912. Psyche 19: 90. N. name for rufiventris Emery.
Formica subpolita var. camponoticeps Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bul. 53: 399, 535. 8.
Formica subpolita var. ficticia Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
561. ¥, 2, 6.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 106-115.

Biology: Wheeler, 1910. Ants, pp. 201, 460. —Wheeler, 1917. Amer. Acad. Arts and Sci., Proc.
52: 553-554. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 87. —Cole, 1942. Amer.
Midland Nat. 28: 384, 377. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 26
(Nev. Test Site). —Wheeler and Wheeler, 1973. Ants of Deep Canyon, p. 129.

subsericea Say. N. B., Que. s. to Fla. w. to Man., Mont., Iowa, Kans., Mo., Miss. Ecology:
Occurs in open deciduous woodlands where it nests in the soil under stones or leaf
litter; sometimes builds low mounds which may be covered with debris. This species has
been confused with F’. fusca L. in the literature.
Formica subsericea Say, 1836. Boston Jour. Nat. Hist. 1: 289. 9, d.
Formica lecontei Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 542. 9, 2, d.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 172-181.

Biology: Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 542. —Wilson and Francoeur,

1974. Fla. Ent. 57: 115-116 (Fla.).

transmontanis Francoeur. Alta., B. C., Mont., Idaho, Calif.
Formica transmontanis Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 135-140. &,

Oe

xerophila Smith. Wash., Utah, Ariz., Calif.
Formica (Neoformica) moki xerophila Smith, 1939. Ent. Soc. Amer., Ann. 32: 583. °.
Formica moki grundmanni Cole, 1943. Amer. Midland Nat. 29: 184. °.

Taxonomy: Francoeur, 1973. Soc. Ent. du Quebec, Mem. no. 3, pp. 259-263.

1456 Hymenoptera in America North of Mexico

SPECIES GROUP EXSECTA

Ants of this group build large conspicuous mounds in fields, woods, or at the edge of woods.
The nest founding female may behave as a temporary social parasite, but colonies are also
founded by budding.

Taxonomy: Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 510-515.
exsectoides Forel. N. S., Ont. s. to Ga. w. to Wis., Iowa, Kans., Colo., N. Mex. Ecology: A large
aggregation of mounds of this species may occur in a single locality. Sometimes damages
bark and cambium of small trees and shrubs. Host: Formica fusca L. Often locally
called the Allegheny mound ant.
Formica exsectoides Forel, 1886. Soc. Ent. de Belg., Ann. (C. R.) 30: 38. 3, @.
Formica exsectoides exsectoides var. davisi Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 396, 484. 3, 2.
Formica exsectoides exsectoides var. hesperia Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 396, 484. §.

Taxonomy: Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 653 (worker, male). —Wheeler,
1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 396, 481-483 (each caste). —Buren, 1944.
Iowa State Col., Jour. Sci. 18: 300. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46:
164 (larva). —Cole, 1954. Tenn. Acad. Sci., Jour. 29: 166.

Biology: McCook, 1877. Amer. Ent. Soc., Trans. 6: 253-295. —Pierson, 1922. Jour. Forestry
20: 325-336. —Manter, 1925. Jour. Econ. Ent. 48: 348-351. —Andrews, 1925. Psyche 32:
75-87 (growth of mounds). —Andrews, 1926. Psyche 33: 127-150. —Andrews, 1929. Ent.
Soe. Amer., Ann. 22: 369-391 (association with treehoppers). —Andrews, 1929. Quart. Rev.
Biol. 4: 248-257. —Andrews, 1941. Sci. Monthly 43: 530-533. —Haviland, 1947. Jour. Econ.
Ent. 40: 413-418 (biology and control). —Schread, 1949. Jour. Econ. Ent. 42: 501. —Gregg,
1963. Ants of Colo., pp. 597-599. —Christensen and Quick, 1970. lowa Acad. Sci., Proc. 77:
207-209.

Morphology: Eisner and Happ, 1964. Science 134: 329-331 (haemorrhage of a coccinellid
beetle and its repellent effect on ants).
opaciventris Emery. N. Dak., Mont., Wyo., Colo., N. Mex. Ecology: May construct either
earthen mounds or thatched nests.
Formica exsectoides var. opaciventris Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643,
653. 8.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 484-485 (worker, male).
—Gregg, 1952. Psyche 59: 14-16 (female).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 166. —Scherba, 1961. N. Y. Ent. Soc., Jour. 69:
71-87 (reproduction, nest structure). —Scherba, 1963. N. Y. Ent. Soc., Jour. 71: 219-231.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 599-601. —Gregg, 1963. Ants of Colo.,
pp. 599-601. —Scherba, 1964. N. Y. Ent. Soc., Jour. 72: 231.

ulkei Emery. N. S. w. to Man. s. to Ohio, Ind., Ill, lowa, N. Dak., Wyo. Ecology: The large
mounds may be found in woods or grasslands and occasionally have a thin layer of plant
debris on their surfaces. Host: Formica fusca L.
Formica ulkei Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 653. 3.
Formica ulkei var. hebescens Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
396, 487. 9.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 396, 485-487 (each
caste). —Buren, 1944. Iowa State Col., Jour. Sci. 18: 300. —Gregg, 1944. Ent. Soc. Amer.,
Ann. 37: 472. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 168 (larva).

Biology: Holmquist, 1928. Ecology 9: 70-87. —Holmquist, 1928. Physiol Zool. 1: 325-327.
—Park, 1929. Psyche 36: 195-215. —Dreyer and Park, 1932. Psyche 39: 127-133.
—Creighton, 1934. Psyche 41: 185-200. —Park, 1936. Psyche 42: 216-231. —Scherba, 1958.
Insectes Sociaux 5: 201-213 (reproduction, mounds). —Kannowski, 1959. Insectes Sociaux
6: 142-143. —Talbot, 1959. Amer. Midland Nat. 61: 128-132 (flight activities). —Scherba,
1959. Amer. Midland Nat. 61: 499-508 (moisture regulation in mounds). —Talbot, 1961.

Superfamily FORMICOIDEA 1457

Ecology 42: 202-205 (mounds). —Scherba, 1962. Amer. Midland Nat. 67: 373-385 (mound
temperatures). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 256-258.

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).
SPECIES GROUP RUFA

Nests of members of this group are varied, some species build mounds and others make con-
siderable use of thatching. Nests are usually started beneath objects. Nest founding females of
some species may behave as temporary social parasites.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 1-10 (N. Amer. variants of Formica
rufa). —Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 472-496.
aterrima Cresson. Colo. Known only from the type male which has not been associated with
any other described form.
Formica aterrima Cresson, 1865. Ent. Soc. Phila., Proc. 4: 426. ¢.

Taxonomy: Brown, 1947. Ent. News 58: 8-9. —Gregg, 1963. Ants of Colo., pp. 544, 546.
calviceps Cole. N. Mex. (Capulin Mt. Natl. Mon.). Ecology: Colony found beneath large stones
which were banked with detritus.
Formica (Formica) calviceps Cole, 1954. Tenn. Acad. Sci., Jour. 29: 164-165. 9.
ciliata Mayr. Minn., N. Dak., Mont., Wyo., Colo., Utah. Ecology: Nests in meadows or open
woods; sometimes utilizes thatching.
Formica ciliata Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 428. 8.

Taxonomy: Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19: 639-643 (each caste). —Wheeler,
1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392-393, 452-454 (each caste).

Biology: Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 88. —Wheeler, 1910. Ants, pp. 114, 120, 205,
351, 444-445, 450. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 233-234. —Gregg,
1963. Ants of Colo., pp. 545-547.

comata Wheeler. S. Dak., Mont., Colo. Ecology: Nests under stones which are banked with
thatch.
Formica comata Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 85. 8, 2, d.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 393, 454-456 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376, 379 (also biological notes).

Biology: Gregg, 1963. Ants of Colo., pp. 547-548, 550.
criniventris Wheeler. N. Dak., S. Dak., Mont. s. to Colo., Utah. Ecology: Found in meadows
and open forests where it nests under stones banked with thatch.
Formica crinita Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 87. 3, 2. Preoce. by Smith, 1858.
Formica criniventris Wheeler, 1912. Psyche 19: 90. N. name for crinita Wheeler.

Taxonomy: Wheeler 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 393, 457-458.
—Cole, 1942. Amer. Midland Nat. 28: 376, 379.

Biology: Wheeler, 1910. Ants, pp. 114, 444-445, 450. —Wheeler and Wheeler, 1944. N. Dak
Hist. Quart. 11: 260. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 234-236. —Gregg,
1963. Ants of Colo., pp. 549-551.

dakotensis Emery. N. S., Ont. w. to Alaska, B. C. s. to Ohio, Iowa, N. Mex., Nev. Ecology:

Usually found in grasslands where it nests in earthen mounds or under stones, about
roots of plants, and in grass clumps banked with considerable detritus. Host: Formica
fusca L., F. lepida Wheeler, F. montana Emery, F. pallidefulva Latreille. Some authors
have recognized two subspecies, dakotensis east of the Rockies and montigena in the
rockies and westward.

Formica dakotensis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 644, 652. 3.

Formica fusca subpolita var. ? specularis Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
663. 2.

Formica dacotensis var. Wasmanni Forel, 1904. Soc. Ent. de Belg., Ann. 48: 153. 8, 2, d.

Formica montigena Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 374. Bis de

Formica dakotensis var. saturata Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 542.

3.

1458 Hymenoptera in America North of Mexico

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 391, 393, 394, 462-465.
—Buren, 1944. Iowa State Col., Jour. Sci. 18: 299. —Brown, 1957. Ent. News 68: 167.

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 364-365. —Wheeler, 1910. Ants, pp.
118, 205, 444-445. —Abbott, 1926. Ent. News 37: 210-211. —Wheeler and Wheeler, 1963.
Ants of N. Dak., pp. 236-238. —Gregg, 1963. Ants of Colo., pp. 551-554.

Morphology: Abbott, 1937. Ent. Soc. Amer., Ann. 20: 117-122 (physiology).
ferocula Wheeler. Ill. (Rockford). Ecology: Type specimens were found nesting in a dry open
field in small crater nests.
Formica ferocula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 461. &.
fossaceps Buren. Que., Iowa, N. Dak. Ecology: Found in grasslands and wooded pastures
nesting under stones or fallen logs banked with thatch or in low earthen mounds
covered with fine thatch of grass materials.
Formica fossaceps Buren, 1942. Iowa State Col., Jour. Sci. 16: 402. 3, 2, 3.

Biology: Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 302. —King, 1949. Iowa Acad. Sci.,
Proc. 56: 367-370 (mixed colony with F’. obscuriventris clivia). —King, 1951. Iowa Acad.
Sci., Proc. 58: 487-489. —King, 1952. Iowa Acad. Sci., Proc. 59: 469-474 (macropseudogynes,
or ? pterergates). —King and Sallee, 1957. Iowa Acad. Sci., Jour. 64: 667-669 (mixed colony
with F’. obscuriventris clivia). —King and Sallee, 1959. Iowa Acad. Sci., Proc. 66: 472-473
(slave of F. rubicunda). —King and Sallee, 1962. Iowa Acad. Sci., Proc. 69: 531-539 (mixed
colony with F. obscuriventris clivia). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
238-239. —Finnegan, 1973. Canad. Ent. 105: 441-444 (diurnal foraging activity).

haemorrhoidalis Emery. N. Dak., S. Dak., Colo. w. to B. C., Wash., Calif. Ecology: Found in

areas of moderate to sparse cover where nests are usually started under logs or stones
and later with moderate use of thatching.

Formica rufa integra var. haemorrhoidalis Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
644, 652. 8.

Formica truncicola integroides var. ravida Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 560. %, 2.

Formica truncicola integroides var. tahoensis Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 538. 3, 2.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 394, 441-442 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376. —Brown, 1965. Ent. News 76: 181-186.
—Cole, 1956. Tenn. Acad. Sci., Jour. 3: 260. —Wheeler and Wheeler, 1963. Ants. of N. Dak.,
p. 230 (forms B, C, and D).

Biology: Wheeler, 1910. Ants, pp. 206, 444. —Creighton, 1940. Amer. Mus. Novitates 1055:
6-7. —Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 88-89. —Gregg, 1963. Ants of Colo., pp.
554-557, 579-581.

integra Nylander. N.S. s. to Ga. w. to Mich., S. Dak., Ill, Miss. Ecology: Large colonies are
found in open forests and woods where they nest in stumps and logs or under stones;

plant debris commonly covers the nest.
Formica integra Nylander, 1856. Ann. des Sci. Nat., Zool. 5: 62. 8.
Formica integra var. similis Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 425. 8, 2, d.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 394, 444-445 (each
caste).

Biology: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 399. —Wheeler, 1910. Ants, pp.
204-206, 222, 351, 444. —Davis and Bequaert, 1922. Brooklyn Ent. Soc., Bul. 17: 18-19.
—Creighton, 1940. Amer. Mus. Novitates 1055: 6. —Kloft, et al., 1973. Fla. Ent. 56: 67-76
(habitat, nest structure, polygamy, biometry).

integroides coloradensis Wheeler. Colo., N. Mex., Idaho, Utah. Ecology: Found in forests at
high elevations (mostly 8,000 to 10,000 feet); nests are started under stumps or logs and
later may be large and dome-shaped with extensive use of thatch.
Formica truncicola integroides var. coloradensis Wheeler, 1913. Harvard Univ., Mus.
Comp. Zool., Bul. 53: 393, 440. ¥, 2.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 377, 381.

Superfamily FORMICOIDEA 1459

Biology: Creighton, 1940. Amer. Mus. Novitates 1055: 6-7. —Gregg, 1963. Ants of Colo. pp.
557-560.
integroides integroides Emery. Wash., Oreg., Calif. Ecology: Inhabits open woods in the

coastal mountains and west slopes of the Cascades and Sierras where it nests under
logs and stumps banked with plant debris.

Formica rufa obscuriventris var. integroides Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
644, 649. 8.

Formica truncicola integra var. subcaviceps Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 540. %, do.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 393, 394, 438-439.
—Creighton, 1940. Amer. Mus. Novitates 1055: 6-7.

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 537. —Miller, 1957. Insectes
Sociaux 4: 253.
integroides planipilis Creighton. N. Dak., Colo., Nev. Ecology: Found in grasslands and open
forests; nests are frequently started at bases of plants and extensive use is made of
thatching; completed nests are moundlike.
Formica rufa planipilis Creighton, 1940. Amer. Mus. Novitates 1055: 7, 9. .

Taxonomy: Cole, 1956. Tenn. Acad. Sci., Jour. 31: 259 (female).

Biology: Wheeler and Wheeler, 1963. Ants. of N. Dak., pp. 248-249. —Gregg, 1963. Ants of
Colo., pp. 560-561. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 23 (Nev. Test
Site).

integroides propinqua Wheeler. Colo., Utah, Wash., Calif. Ecology: Found on the east slopes
of the Caseades and Sierras in Wash. and Calif.; inhabits forests where nests are
started under logs and stumps and considerable use is made of thatching.
Formica truncicola integroides var. propinqua Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 538. 8.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 6-7. —Cole, 1956. Tenn. Acad. Sci.,
Jour. 31: 257-259 (female, male).

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 88. —Gregg, 1963. Ants of Colo., pp.
562-564.
integroides subfasciata Wheeler. Calif. (Mill Creek Canyon, Wilson Peak, 7500 ft., San
Bernardino Mtns.).
Formica truncicola integroides var. subfasciata Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 539. 8.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 7. —Brown, 1965. Ent. News 76:
185 (possible syn. of integroides Emery).
laeviceps Creighton. N. Dak., S. Dak., Colo., Utah. Ecology: Nests mostly in open areas under
stones and logs which are banked with little debris.
Formica rufa laeviceps Creighton, 1940. Amer. Mus. Novitates 1055: 7, 9. Oe

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 377, 380 (also biological notes).

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 239-240. —Gregg, 1963. Ants of
Colo., pp. 564-565.
lugubris Zetterstedt. Que.; Europe. Ecology: Populous colonies are found in large mound nests
of thatch and debris in open forests. Introduced into Valcartier, Quebec in 1971 from
Italy for use as a predator of the Swaine jack pine sawfly, Neodiprion swainei Midd. It
is apparently established in Quebec.
Formica lugubris Zetterstedt, 1840. Ins. Lapponica, v. 1, p. 449. é.

Taxonomy: Yarrow, 1955. Soc. Brit. Ent., Trans. 12: 5, 10 (synonymy, also biological notes).

Biology: Pavan, 1959. Min. dell’ Agr. e delle For. [Italy] 4: 5 (in Italy). —Pavan, 1963. Pavia
Univ. Symp. Genet. et Biol. Ital. 11: 61-84 (use to protect alpine forests against insect
damage). —Pavan, 1963. Pavia Univ. Symp. Genet. et Biol. Ital. 12: 122-131 (reconstruction
of nest). —Klimetzek, 1970. Ztschr. f. Angew. Ent. 66: 84-95 (environmental factors and

1460 Hymenoptera in America North of Mexico

distribution). —Williamson, 1973. Canad. Dept. Agr., Liberation Bul. no. 36, p. 2. (in Que.).
—Finnegan, 1975. Canad. Ent. 107: 1271-1274 (introduction into eastern Canada).
mucescens Wheeler. Colo., Utah. Ecology: Found in open areas where it nests under stones;
moderate use is made of thatching.
Formica truncicola mucescens Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
392, 394, 442. 8, 2, do.

Taxonomy: Creighton, 1940. Amer. Mus. Novitates 1055: 6-7. —Cole, 1942. Amer. Midland
Nat. 28: 376, 380.

Biology: Gregg, 1963. Ants of Colo., pp. 566-567.

obscuripes Forel. Que.; Mich., Ind., Man. w. to B. C. s. to N. Mex., Utah., Calif. Ecology: Nests

are found in open areas where they are usually started at the base of a plant. Extensive
use is made of thatching and the finished nest is a large mound of detritus. Probably
the most common thatching ant of the western states. Introduced into Quebec from
Manitoba for use as a predator of the Swaine jack pine sawfly, Neodiprion swainei
Midd.

Formica rufa obscuripes Forel, 1886. Soc. Ent. de Belg., Ann. (C. R.) 30: 39. 8.

Formica rufa obscuriventris var. melanotica Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
644, 650. 8.

Formica rufa obscuriventris var. rubiginosa Emery, 1893. Zool. Jahrb., Abt. f. System. 7:
644, 650. “2” = %. Preoce. by Guillou, 1841.

Formica rufa aggerans Wheeler, 1912. Psyche 19: 90. N. name for rubiginosa Emery.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 394, 432-433 (each
caste). —Creighton, 1940. Amer. Mus. Novitates 1055: 7. —Buren, 1944. Iowa State Col.,
Jour. Sci. 18: 300. —Wheeler and Wheeler, 1953. Ent. Soc. Amer., Ann. 46: 165-166 (larva).

Biology: Treherne, 1915. Canad. Ent. 47: 104. —Burrill and Smith, 1919. Ohio Jour. Sci. 19:
286. —Cole, 1932. Psyche 39: 30-33. —Weber, 1935. Ecol. Monog. 5: 165-206 (in N. Dak.).
—KEssig, 1926. Ins. West. No. Amer., p. 867. —Eckert and Mallis, 1937. Calif. Agr. Expt.
Sta. Cir. 342: 30. —Weber, 1941. Canad. Ent. 73: 140-141. —Cole, 1942. Amer. Midland Nat.
28: 380. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 263-266. —Sallee and King,
1947. Iowa Acad. Sci., Proc. 54: 349-352. —King and Walters, 1950. Iowa Acad. Sci., Proc.
57: 469-47. —King and Sallee, 1953. Iowa Acad. Sci., Proc. 60: 656-659. —King and Sallee,
1957. Iowa Acad. Sci., Proc. 63: 721-723. —Talbot, 1959. Amer. Midland Nat. 61: 124-128
(flight activities). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 240-244. —Gregg,
1963. Ants of Colo., pp. 566-572. —Bradley, 1972. Canad. Ent. 104: 245-249 (transplanting
colonies in jack pine stands in Man.). —Talbot, 1972. Kans. Ent. Soc., Jour. 45: 254-258
(flights and swarms). —Alpert and Akre, 1973. Ent. Soc. Amer., Ann. 66: 753-760
(association with inquiline, Leptothorax diversipilosus Smith, in Wash.). —Williamson,
1973. Canad. Dept. Agr., Liberation Bul. no. 36, p. 2 (introduction into Que.).

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).
obscuriventris clivia Creighton. Man., Wis., Iowa w. to B. C. s. to N. Mex., Utah. Ecology:

Nests in forests under logs and stones; moderate use is made of thatching.
Formica rufa clivia Creighton, 1940. Amer. Mus. Novitates 1055: 8, 9. ¥, 2, d.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 377. —Buren, 1944. Iowa State Col., Jour. Sci.
18: 300, 302.

Biology: King, 1949. Iowa Acad. Sci., Proc. 56: 367-370 (mixed colony with F. fossaceps
Buren). —King and Sallee, 1951. Iowa Acad. Sci., Proc. 58: 487-489. —King, 1955. lowa
Acad. Sci., Proc. 62: 509-513 (winged workers). —King and Sallee, 1959. Iowa Acad. Sci.,
Proc. 66: 472-473 (slave of F. rubicunda Emery). —King and Sallee, 1962. Iowa Acad. Sci.,
Proc. 69: 531-539 (mixed colonies with F’. fossaceps Buren). —Gregg, 1963. Ants of Colo.,
pp. 574-577.

obscuriventris obscuriventris Mayr. Que., Maine s. to Va. w. to N. Dak., lowa, Colo. Ecology:
Nests in woods and grasslands under logs and makes moderate use of thatching.
Formica truncicola var. obscuriventris Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 951.
3.
Formica dryas Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 268. 8, °.

Superfamily FORMICOIDEA 1461

Formica dryas var. gymnomma Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 269. %.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 448. —Creighton,
1940. Amer. Mus. Novitates 1055: 8.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 623. —Wheeler, 1917. Conn. State
Geol. and Nat. Hist. Survey Bul. 22: 595, 597. —Buren, 1941. Iowa State Col., Jour. Sci. 15:
115. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 244-246. —Gregg, 1963. Ants of
Colo., pp. 572-574. —Talbot, 1964. Anim. Behaviour 12: 154 (nest structure and
flights).—Ayre, 1968. Canad. Ent. 100: 165-172 (prey finding, capture and transport).
oreas comptula Wheeler. Sask., N. Dak., Iowa w. to Alta., Wash., Idaho, Utah. Ecology: Nests
in wooded areas or grasslands under stones or logs banked with detritus.

Formica oreas var. comptula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53:
391, 393, 460. §, 2.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 376, 380.

Biology: Cole, 1934. Psyche 41: 227. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
246-248.
oreas oreas Wheeler. Mont., Wyo., Colo., N. Mex., Utah. Ecology: Nests in open woods or
meadows under rocks or logs covered with detritus. Colonies may be abundant and have
sizeable mounds.
Formica oreas Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19: 643. ¥, 2, d.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 391, 393, 458-460.
—Cole, 1942. Amer. Midland Nat. 28: 376, 379.

Biology: Wheeler, 1910. Ants, pp. 114, 205, 351, 444. —Gregg, 1963. Ants of Colo., pp. 577-579.
prociliata Kennedy and Dennis. Ohio, Wis., Iowa. Ecology: Nests of the type specimens were
under stones on ledges in open woods and lacked thatching.
Formica prociliata Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 531. %, 2, d.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 303.
reflexa Buren. Que., Minn., Iowa, N. Dak. Ecology: Has only been found in association with the
host; a permanent inquiline ?. Host: Formica fusca L.
Formica reflexa Buren, 1942. Iowa State Col., Jour. Sci. 16: 399. 3, 2, d.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 299, 303.

Biology: King, 1949. Iowa Acad. Sci., Proc. 56: 367-379 (mixed colony with F. fusca). —King,
1951. Iowa Acad. Sci., Proc. 58: 487-489. —Wheeler and Wheeler, 1963. Ants of N. Dak., p.
249-250. —Francoeur and Beique, 1968. Nat. Canad. 95: 228 (Que.).

subnitens Creighton. N. Dak., Wyo., Colo., N. Mex., B. C., Oreg., Calif. Ecology: Usually nests
in grasslands, under stones banked with debris or in dome-shaped mounds of thatch or
detritus.
Formica rufa subnitens Creighton, 1940. Amer. Mus. Novitates 1055: 7, 10. 9.

Taxonomy: Cole, 1955. Tenn. Acad. Sci., Jour. 30: 50. —Miller, 1957. Insectes Sociaux 4:
253-257 (each caste).

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 165. —Ayre, 1957. Insectes Sociaux 4: 173-176
(ecological notes). —Ayre, 1958. Insectes Sociaux 5: 1-7. —Ayre, 1958. Insectes Sociaux 5:
147-157 (meterological factors affecting foraging). —Ayre, 1959. Insectes Sociaux 6:
105-114 (food habits). —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 250-252. —Gregg,
1963. Ants of Colo., pp. 581-584.

SPECIES GROUP MICROGYNA

Species of this group are believed to be temporary social parasites of other species of For-
mica. The female in some way is adopted by workers of the host species. Host workers may
remain in the colony after the intruding queen has established her own brood, but the host wor-
kers eventually die. Most species are found in open woods or meadows. The nests are usually of
the thatch type, but the thatching is normally scattered about the nest openings and appears as
a flattened disc.

1462 Hymenoptera in America North of Mexico

Taxonomy: Letendre and Huot, 1972. Soc. Ent. Quebec, Ann. 17: 117-132 (preliminary
considerations to a revision of the microgyna group).
densiventris Viereck. Colo., N. Mex., Utah w. to Calif. Ecology: Found in forests and open
forests, under stones and logs; only occasionally with use of thatch.
Formica fusca var. densiventris Viereck, 1903. Amer. Ent. Soc., Trans. 29: 74. §.
Formica microgyna rasilis var. spicata Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bul. 53: 395, 469. %, 2, o.

Formica microgyna rasilis var. pinetorum Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 542. 8.

Taxonomy: Brown, 1947. Ent. News 58: 6-8. —Cole, 1955. Tenn. Acad. Sci., Jour. 30: 50.

Biology: Mallis, 1941. South. Calif. Acad. Sci., Bul. 40: 85. —Cole, 1954. Tenn. Acad. Sci., Jour.
29: 165. —Gregg, 1963. Ants of Colo., pp. 589-593.
difficilis Emery. Mass. s. to Ga. w. to lowa. Ecology: Nests are in woodlands, sometimes under
stones, and are packed with vegetable debris. Host: Formica pallidefulva (Latreille).
Formica rufa difficilis Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 645, 651. 8, 2, d.
Formica difficilis var. consocians Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 371. 8,
5 ele
Formica habrogyna Cole, 1939. Amer. Midland Nat. 22: 413. %, 9.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 395, 477-480 (each
caste). —Buren, 1944. Iowa State Col., Jour. Sci. 18: 306. —Wheeler and Wheeler, 1953.
Ent. Soc. Amer., Ann. 46; 164 (larva).

Biology: Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 347-373. —Wheeler, 1906. Amer. Mus.
Nat. Hist., Bul. 22: 50-64. —Wheeler, 1910. Ants, pp. 113-114, 205-206, 441-444.
dirksi Wing. Maine (Daigle, Aroostook Co.). Ecology: The host nest of the type queen was at
the edge of a clearing under bark of a dry stump. Host: Formica fusca L.
Formica dirksi Wing, 1949. Canad. Ent. 81: 13-15. 2.
impexa Wheeler. Mass., Mich., Minn. Host: Formica fusca L.
Formica impexa Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 273. 8.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp Zool., bul. 53: 395-396, 472-473
(worker, female).

Biology: Wheeler, 1906. Psyche 13: 40. —Wheeler, 1910. Ants, pp. 113, 444.
indianensis Cole. Ind., lowa. Ecology: Nest was in a grassy field under a stone and was
surrounded with grass debris.
Formica indianensis Cole, 1940. Amer. Midland Nat. 23: 224. 3, d.

Taxonomy: Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 306.
knighti Buren. Iowa (Bonaparte). Ecology: Nest was in a pasture under bushes and
surmounted by a low dome of plant debris.
Formica (Formica) knighti Buren, 1944. Iowa State Col., Jour. Sci. 18: 300, 303-305. &.
microgyna Wheeler. Wyo., Colo., N. Mex., Utah, Nev. Ecology: Prefers meadows and open
forests. Nests are usually started under stones which are piled with thatching, the
thatching increasing in size as the nest grows. Host: Formica argentea Wheeler, F.
lasioides Emery, F’. neogagates Emery.
Formica microgyna Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19: 645. 8, 9, d.
Formica microgyna microgyna var. recidiva Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 395, 467. 8, o.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 395-396, 465-467 (each
caste).

Biology: Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 542. —Gregg, 1963. Ants of
Colo., pp. 584-587. —Cole, 1966. Brigham Young Univ., Sci. Bul., Biol. Ser. 7: 24 (Nev. Test
Site).

morsei Wheeler. Mass. (South Natick).
Formica morsei Wheeler, 1906. Psyche 13: 39. &.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp.Zool., Bul. 53: 394, 480-481.

Superfamily FORMICOIDEA 1463

nepticula Wheeler. Mass., Conn., Ill., lowa. Ecology: Nests in open woods under stones or
rotting limbs and are banked with vegetable debris.
Formica nepticula Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 270. %, 2, d.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 394, 396, 475-477 (each
caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 595, 597.
—Buren, 1944. Iowa State Col., Jour. Sci. 18: 30, 306.

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 64. —Wheeler, 1910. Ants, pp. 113,
205, 444.
nevadensis Wheeler. Nev. (Ormsby Co.). Ecology: Nests in open forests under stones and

surmounted by plant debris.
Formica microgyna var. nevadensis Wheeler, 1904. Amer. Mus. Nat. Hist., Bul. 20: 373. 9.

Taxonomy: Wheeler, 1905. Amer. Mus. Nat. Hist., Bul. 21: 272. —Wheeler, 1913. Harvard
Univ., Mus. Comp. Zool., Bul. 53: 396, 470-472 (worker). —Wheeler, 1917. Amer. Acad. Arts
and Sci., Proc. 52: 544. —Cole, 1956. Tenn. Acad. Sci., Jour. 31: 256-257 (also biology).

Biology: Wheeler, 1910. Ants, pp. 11, 444.
postoculata Kennedy and Dennis. Pa., Ind., Ill., lowa. Ecology: Nest of type series from
Indiana was in a pasture between two boulders and was covered with trash.
Formica postoculata Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 540-542. &.
querquetulana Kennedy and Dennis. New England to Ohio. Ecology: Nests found in pastures
and sandy areas usually beneath objects and commonly covered with loose trash. Host:

Formica fusca L.
Formica querquetulana Kennedy and Dennis, 1937. Ent. Soc. Amer., Ann. 30: 536-540. &,
2

rasilis Wheeler. Colo., N. Mex., Utah northwest to Wash. Ecology: Nests in open or semi-open
areas usually under stones. Host: Formica argentea Wheeler, F. fusca L. Creighton
(1950) stated that there is a great deal of intergradation between rasilis and spicata
(see synonymy of densiventris); consequently, rasilis and densiventris may be
conspecific.
Formica microgyna var. rasilis Wheeler, 1903. Amer. Mus. Nat. Hist., Bul. 19: 648. 9, 2,
é.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 395-396, 468-469.
—Creighton, 1950. Harvard Univ., Mus. Comp. Zool., Bul. 104: 506-507.

Biology: Cole, 1942. Amer. Midland Nat. 28: 376, 381. —Gregg, 1963. Ants of Colo., pp.
587-589.
scitula Wheeler. Ga. (Clayton).
Formica microgyna scitula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 395,
470. 2.
spatulata Buren. Minn., Iowa, N. Dak., Mont., B. C. Ecology: Nests under stones. Host:
Formica fusca L. ;
Formica (Formica) microgyna spatulata Buren, 1944. Iowa State Col., Jour. Sci. 18: 300,
305. ¥, 2, d.

Biology: Wheeler and Wheeler, 1963. Ants of N. Dak., p. 253.
whymperi adamsi Wheeler. Mich. to Minn. Ecology: Has been found in drier tamarack
swamps in nests covered with leaves.
Formica adamsi Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 84. 9.
Formica adamsi Wheeler, 1909. Mich. Geol. Survey, Bul. 5: 326. 9.

Taxonomy: Wheeler, 1913. Harvard Univ. Mus. Comp. Zool., Bul. 53: 395, 473-474. — Wheeler,
1917. Amer. Acad. Arts and Sci., Proc. 52: 544.
Biology: Drooz, 1960. U. S. Dept. Agr., Tech. Bul. 1212, p. 36 (predator of larch sawfly).

whymperi alpina Wheeler. Colo., N. Mex., Idaho, Utah. Ecology: Found at high elevations,
mostly above 9,000 feet where it nests in forests under stones or rotting logs and with
some plant debris or thatch around these objects. Host: Formica neorufibarbis Emery.
Formica adamsi var. alpina Wheeler, 1909. N. Y. Ent. Soc., Jour. 17: 85. %.

1464 Hymenoptera in America North of Mexico

Formica adamsi var. alpina Wheeler, 1909. Mich. Geol. Survey, Bul. 5: 327. &.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 395, 475. —Wheeler,
1917. Amer. Acad. Arts and Sci., Proc. 52: 544.

Biology: Cole, 1954. Tenn. Acad. Sci., Jour. 29: 165-166. —Gregg, 1963. Ants of Colo., pp.
593-596 (also description of female).
whymperi californica Wheeler. Mont., Idaho., Wash. s. to Calif.
Formica microgyna californica Wheeler, 1917. Amer. Acad. Arts and Sci., Proc. 52: 543.
3.
Formica microgyna californica var. hybrida Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 543. 8.
whymperi whymperi Forel. Mont. w. to B. C., Wash. Ecology: Nests under stones and logs
which it banks with plant debris. Host: Formica neoclara Emery.
Formica rufa obscuripes var. whymperi Forel, 1904. Soc. Ent. de Belg., Ann. 48: 152. &.
Formica microgyna rasilis var. pullula Wheeler, 1913. Harvard Univ., Mus. Comp. Zool.,
Bul. 53: 562. 8, @.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 392, 434. —Wheeler,
1917. Amer. Acad. Arts and Sci., Proc. 52: 544 (also biological notes).

SPECIES GROUP SANGUINEA

Members of this subgenus are social parasites. Nest founding females forceably enter small
colonies of the host species and kill or drive off the host workers before rearing the host brood
for her own use. Hosts are other species of Formica. Some species are known to regularly con-
duct slave raids, a habit unknown in other Formica species.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 108-129. —Buren, 1968. Ga. Ent. Soc., Jour.
3: 25-40 (key to spp.).

Biology: Wheeler, 1910. Ants, pp. 458-470. —Creighton, 1950. Harvard Univ., Mus. Comp.

Zool., Bul. 104: 460-461.
creightoni Buren. Mich., Ill, lowa. Host: Formica neogagates Emery.

Formica (Raptiformica) creightoni Buren, 1968. Ga. Ent. Soc., Jour. 3: 35-36. &.
curiosa Creighton. B. C., Mont., Idaho, Oreg. Host: Formica lasioides Emery, F. manni

Wheeler.
Formica curiosa Creighton, 1935. Amer. Mus. Novitates 773: 5. %, 2.
Formica parcipappa Cole, 1946. Ent. Soc. Amer., Ann. 36: 616. &.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 123-125. —Buren, 1968. Ga. Ent. Soc., Jour.
3: 32-33. —Snelling, 1969. Ent. Soc. Wash., Proc. 71: 194-196 (syn. of parcipappa; biological
notes).

emeryi Wheeler. Colo. (Bradmoor, Colorado Springs). Host: Formica neogagates Emery.
Formica emeryi Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 389, 419. 3, 9.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 118 (a syn. of subnuda). —Gregg, 1963.
Ants of Colo., pp. 606-608 (valid sp.). —Buren, 1968. Ga. Ent. Soc., Jour. 3: 34-35 (valid sp.).
pergandei Emery. Que., N. H.s. to N. C. w. to N. Dak. S. Dak., Iowa, Colo. Host: Formica
fusca L., F. pallidefulva Latreille (?).
Formica pergandei Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 646. 9.
Formica sanguinea rubicunda var. sublucida Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 390, 408. 8, 2.

Taxonomy: Buren, 1968. Ga. Ent. Soc., Jour. 3: 28-29.

Biology: Wheeler, 1901. Amer. Nat. 35: 722. —Gregg, 1946. Amer. Midland Nat. 35: 752.
—Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 219-220. —Gregg, 1963. Ants of Colo.,
pp. 622-623. —Regnier and Wilson, 1971. Science 172: 267-269 (chemical communication).
—Hung, 1973. Ent. News 84: 253-259 (reproductive biology). —Finnegan, 1973. Canad.
Ent. 105: 441-444 (diurnal foraging activity; as sublucida). ~

Superfamily FORMICOIDEA 1465

puberula Emery. Man., N. Dak., S. Dak., Wyo., Colo., Tex. w. to B. C., Wash., Calif. Host:
Formica altipetens Wheeler, F’. bradleyi Wheeler, F. fusca L., F. hewitti Wheeler, F.
lasioides Emery, F’. lepida Wheeler, F. montana Emery, F. neoclara Emery, F.
neorufibarbis Emery, F. pallidefulva nitidiventris Emery, F. rasilis Wheeler, F.
subpolita Mayr.
Formica sanguinea puberula Emery, 1893. Zool. Jahrb., Abt. f. System 7: 643, 648. °.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 390, 413-414 (each
caste). —Cole, 1942. Amer. Midland Nat. 28: 376. —Buren, 1968. Ga. Ent. Soc., Jour. 3:
30-31.

Biology: Wheeler, 1910. Ants, pp. 458-460. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
220-221. —Gregg, 1963. Ants of Colo., pp. 613-616. —Snelling, 1969. Ent. Soc. Wash., Proc.
71: 196-197.

rubicunda Emery. Ont. s. to N. C., Tenn. w. to Mont., Colo., N. Mex. Host: Formica altipetens
Wheeler, F’. bradleyi Wheeler, F’. fossaceps Buren, F. fusca L., F. lasioides Emery, F.
lepida Wheeler, F. montana Emery, F’. neoclara Emery, F’. neogagates Emery, F.
neorufibarbis Emery, F’. obscuriventris clivia Creighton, F’. pallidefulva nitidiventris
Emery, F’. schaufussi Mayr.
Formica sanguinea rubicunda Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 643, 647. 9, 9.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 390, 406-408 (each
caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 595. —Buren,
1944. Iowa State Col., Jour. Sci. 18: 299. —Buren, 1968. Ga. Ent. Soc., Jour. 3: 30.

Biology: Wheeler, 1910. Ants, pp. 458-470. —Smith, 1928. N. Y. Ent. Soc., Jour. 26: 327-329.
—Talbot, 1934. Ecology 15: 421, 430-434. —King and Sallee, 1959. Iowa Acad. Sci., Proc. 66:
472-473. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 221-223. —Gregg, 1963. Ants
of Colo., pp. 616-618.

subintegra Emery. Newfoundland, N. S., Ont. s. to S. C., Tenn. w. to N. Dak., Iowa, Kans.
Host: Formica fusca L., F. lasioides Emery, F. montana Emery, F’. neogagates Emery,
F. pallidefulva nitidiventris Emery, F. schaufussi Mayr, F’. subpolita Mayr.
Formica sanguinea rubicunda var. subintegra Emery, 1893. Zool. Jahrb., Abt. f. System.
7: 643, 648. 9.
Formica sanguinea subintegra var. gilvescens Wheeler, 1913. Harvard Univ., Mus. Comp.
Zool., Bul. 53: 390, 412. 8.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 390, 410-412.
—Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 595. —Buren, 1944.
Iowa State Col., Jour. Sci. 18: 299. —Wilson and Brown, 1955. Psyche 62: 120. —Buren,
1968. Ga. Ent. Soc., Jour. 3: 29-30.

Biology: Wheeler, 1908. Amer. Mus. Nat. Hist., Bul. 24: 624, 627-631. —Smith, 1928. N. Y.
Ent. Soe. Jour. 36: 323-327. —Talbot and Kennedy, 1940. Ent. Soc. Amer., Ann. 33: 560-577.
—Brown, 1958. Psyche 65: 39-40. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp.
225-227. —Kannowski, 1963. Pavia Univ., Symp. Genet. Biol. Ital. 12: 74-102 (mating
behavior). —Regnier and Wilson, 1971. Science 172: 267-269 (chemical communication).

Morphology: Hung, 1969. Ent. Soc. Amer., Ann. 62: 456 (chromosome number).

subnuda Emery. Newfoundland w. to Yukon, Alaska s. to N. Y., Minn., N. Dak., Colo., N. Mex.,
Ariz., Calif. Host: Formica altipetens Wheeler, F. fusca L., F. montana Emery, F.
neorufibarbis Emery, F’. subpolita Mayr.
Formica sanguinea rubicunda var. subnuda Emery, 1895. Zool. Jahrb., Abt. f. System. 8:

335. 3.
Formica sanguinea aserva Forel, 1901. Soc. Ent. de Belg., Ann. 45: 395. %, @.

Taxonomy: Wheeler, 1913. Harvard Univ., Mus. Comp. Zool., Bul. 53: 389, 404-406, 409-410
(each caste). —Wheeler, 1917. Conn. State Geol. and Nat. Hist. Survey Bul. 22: 595.
—Buren, 1944. Iowa State Col., Jour. Sci. 18: 299, 308. —Wheeler and Wheeler, 1953. Ent.
Soc. Amer., Ann. 46: 170 (larva). —Wilson and Brown, 1955. Psyche 62: 118. —Buren, 1968.

Ga. Ent. Soc., Jour. 3: 33-34.

1466 Hymenoptera in America North of Mexico

Biology: Wheeler, 1906. Amer. Mus. Nat. Hist., Bul. 22: 85. —Wheeler, 1908. Amer. Mus. Nat.
Hist., Bul. 24: 623, 631-633. —Wheeler, 1910. Ants, pp. 458-460, 468. —Wheeler, 1917.
Harvard Univ., Mus. Comp. Zool., Bul. 61: 19. —Cole, 1942. Amer. Midland Nat. 28: 378.
—Gregg, 1947. Colo. Univ. Studies 2: 393. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 223-225. —Gregg, 1963. Ants of Colo., pp. 618-621. —Chapman, 1957. Canad. Ent. 89:
392 (swarming on mountain tops). —Finnegan, 1973. Canad. Ent. 105: 441-444 (diurnal
foraging activity).

wheeleri Creighton. N. Dak., Colo., N. Mex., Utah, Ariz. Host: Formica altipetens Wheeler, F.
bradleyi Wheeler, F’. fusca L., F. lasioides Emery, F.. lepida Wheeler, F’. neogagates
Emery, F. neorufibarbis Emery.

Formica wheeleri Creighton, 1935. Amer. Mus. Novitates 773: 1-5. %, °.

Taxonomy: Wilson and Brown, 1955. Psyche 62: 125-126. —Buren, 1968. Ga. Ent. Soc., Jour.
3: 31-32.

Biology: Cole, 1942. Amer. Midland Nat. 28: 376, 378. —Cole, 1954. Tenn. Acad. Sci., Jour. 29:
104. —Wilson, 1955. Psyche 62: 130-133. —Wheeler and Wheeler, 1963. Ants of N. Dak.,
pp. 227-228. —Gregg, 1963. Ants of Colo., pp. 623-625. —Halverson, et al., 1976. Kans. Ent.
Soc., Jour. 49: 299 (bradley as slave).

Genus POLYERGUS Latreille

Polyergus Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, p. 179.
Type-species: Formica rufescens Latreille. Monotypic.

The species of this genus are obligatory or true slave-making ants. In nest founding, the
female enters a nest of the host species, eventually kills the rightful queen, and uses the host
workers to tend her brood. A colony of Polyergus will conduct slave raids on nests of species of
Formica, and workers of the host are taken and used by the Polyergus colony to feed and rear
the brood and excavate the nest.

Revision: Smith, 1947. Amer. Midland Nat. 38: 150-161.
Taxonomy: Wheeler, 1968. Ent. Soc. Wash., Proc. 70: 156-164 (male genitalia and taxonomy).

Morphology: Forbes and Brassel, 1962. N. Y. Ent. Soc., Jour. 70: 79-87 (male genitalia and
terminal segments).
breviceps Emery. Ont., Mich. w. to B. C. s. to Ind., IIl., Mo., Kans., N. Mex., Ariz., Calif. Host:
Formica altipetens Wheeler, F’. argentea Wheeler, F.. fusca L., F. lepida Wheeler, F.
montana Emery, F. neoclara Emery, F. neorufibarbis Emery, F. pallidefulva
nitidiventris Emery, F. schaufussi schaufussi Mayr, F’. subpolita Mayr.
Polyergus rufescens breviceps Emery, 1893. Zool. Jahrb., Abt. f. System. 7: 666. %.
Polyergus rufescens bicolor Wasmann, 1901. Allg. Ztschr. f. Ent. 6: 369. %, 2, d.
Polyergus rufescens breviceps var. Silvestrii Santschi, 1909. Soc. Ent. Ital., Bol. 41: 7. %,
3

Polyergus rufescens breviceps var. montezuma Wheeler, 1914. N. Y. Ent. Soc., Jour. 22: 56.
Emre

Polyergus rufescens breviceps var. umbratus Wheeler, 1915. Amer. Mus. Nat. Hist., Bul.
34: 419. &.

Polyergus rufescens laeviceps Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 420. &.

Polyergus rufescens breviceps var. fusciventris Wheeler, 1917. Amer. Acad. Arts and Sci.,
Proc. 52: 555. 8.

Taxonomy: Cole, 1942. Amer. Midland Nat. 28: 385. —Buren, 1944. Iowa State Col., Jour. Sci.
18: 310. —Smith, 1947. Amer. Midland Nat. 38: 152, 157-159, 161. —Wheeler, 1968. Ent.
Soc. Wash., Proc. 70: 156-164. —Wheeler and Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 214
(larva, as rufescens).

Biology: Wheeler, 1910. Ants, pp. 475-482. —Wheeler, 1916. N. Y. Ent. Soc., Jour. 24: 107-118.
—Smith, 1928. N. Y. Ent. Soc., Jour. 36: 329-333. —Mallis, 1941. South. Calif. Acad. Sci.,
Bul. 40: 82. —Wheeler and Wheeler, 1944. N. Dak. Hist. Quart. 11: 260. —Gregg, 1946.
Amer. Midland Nat. 35: 754. —Wheeler and Wheeler, 1963. Ants of N. Dak, pp. 275-277.

Superfamily FORMICOIDEA 1467

—Gregg, 1963. Ants of Colo., pp. 635-643. —Hung, 1973. Ent. News 84: 253-259
(reproductive biology).
lucidus longicornis Smith. N. C., S. C., Ga. Host: Formica schaufussi schaufussi Mayr.
Polyergus lucidus longicornis Smith, 1947. Amer. Midland Nat. 38: 155. &.
lucidus lucidus Mayr. Mass. s. to S. C. w. to N. Dak., Iowa, Colo., N. Mex. Host: Formica
lasioides Emery, F’. neogagates Emery, F. pallidefulva nitidiventris Emery, F.
schaufussi schaufussi Mayr.
Polyergus lucidus Mayr, 1870. Zool.-Bot. Gesell. Wien, Verh. 20: 952. %.
Polyergus lucidus montivagus Wheeler, 1915. Amer. Mus. Nat. Hist., Bul. 34: 419. 3, 2,
Gre

Taxonomy: Mayr, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 424. —Buren, 1944. Iowa State
Col., Jour. Sci. 18: 310. —Smith, 1947. Amer. Midland Nat. 38: 152-156. —Wheeler and
Wheeler, 1968. Ent. Soc. Amer., Ann. 61: 214 (larva).

Biology: McCook, 1880. Acad. Nat. Sci. Phila., Proc. 32: 376-384. —Burrill, 1908. N. Y. Ent.
Soe., Jour. 16: 144-151. —Wheeler, 1910. Ants, pp. 482-486. —Smith, 1934. N. Y. Ent. Soc.,
Jour. 42: 360-361. —Wheeler and Wheeler, 1963. Ants of N. Dak., pp. 274-275. —Gregg,
1963. Ants of Colo., pp. 633-635. —Talbot, 1967. Psyche 74: 299-313 (slave raids). —Talbot,
1968. Psyche 75: 46-52 (flights). —Harman, 1968. Ent. News 79: 217-223 (ecology, near
Chicago). —Marlin, 1968. Ill. State Acad. Sci., Trans. 61: 207-209 (colony formation).
—Marlin, 1969. Kans. Ent. Soc., Jour. 42: 108-115 (raiding behavior). —Marlin, 1971. Amer.
Midland Nat. 86: 181-189 (mating, nesting, and ant enemies).

UNPLACED TAXA OF FORMICINAE

Formica arenicola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 160. §. Washington, D. C.

Formica atra Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. $. Washington, D. C.

Formica Connecticutensis Buckley, 1866. Ent. Soc. Phila., Proc. 6: 154. §, 9. Conn., N. Y., D.
C.

Formica lauta Say, 1836. Boston Jour. Nat. Hist. 1: 286. 2, 3.

Formica monticola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 157. 3, 2, 3d. N. Y.

Formica Nortonii Buckley, 1866. Ent. Soc. Phila., Proce. 6: 153. 3, 2. Conn.

Formica Nova Anglae Buckley, 1866. Ent. Soc. Phila., Proc. 6: 153. 3, 2. Maine.

Formica occidentalis Buckley, 1866. Ent. Soc. Phila., Proc. 6: 157. 8, 2. Conn., N. Y.

Formica politurata Buckley, 1866. Ent. Soc. Phila., Proc. 6: 160. 3. Mich.

Formica saxicola Buckley, 1866. Ent. Soc. Phila., Proc. 6: 166. %, 2. Buchanan Co., Tex.

Formica semipunctata Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 262. 2. “Taken on a journey
from New York to Cumberland House.”

Formica septentrionale Buckley, 1866. Ent. Soc. Phila., Proc. 6: 161. %, 2. Mich., Ill.

Formica tenuissima Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. ¥. Central Tex.

Formica Virginiana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 159. %. Washington, D. C.

UNPLACED TAXA OF FORMICIDAE
Formica dislocata Say, 1836. Boston Jour. Nat. Hist. 1: 288. 2, d.
Formica Lincecumii Buckley, 1866. Ent. Soc. Phila., Proc. 6: 163. 3, 2, d. Tex.

Polyergus Texana Buckley, 1866. Ent. Soc. Phila., Proc. 6: 170. 2. Buchanan Co., Tex.
Formica triangularis Say, 1836. Boston Jour. Nat. Hist. 1: 288. °, d.

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1469

Superfamily VESPOIDEA

By KARL V. KROMBEIN

Included in this superfamily are several groups with quite diverse habits. The primitive fami-
ly Masaridae includes the only solitary wasps which store pollen and nectar as food for their lar-
vae rather than paralyzed or dismembered Arthropoda. The solitary eumenid wasps store para-
lyzed lepidopterous or coleopterous, or rarely hymenopterous, larvae as food for their young. All
of the truly social wasps belong to the family Vespidae.

Taxonomy: Ashmead, 1902. Canad. Ent. 34: 163-166, 203-210, 219-221 (keys to families and
genera). —Dalla Torre, 1904. Jn Wytsman, Gen. Ins., fase. 19, pp. 1-108, 6 pls. (keys and
species catalog). —Bequaert, 1928. Ann. and Mag. Nat. Hist. (10)2: 188-176 (notes on
vespoid types in British Museum). —Bequaert, 1928. Brooklyn Ent. Soc., Bul. 23: 53-63
(Fabricius types in Banks coll.). —Reid, 1942. Roy. Ent. Soe. London, Trans. 92: 285-331,
137 figs. (larval classification). —Richards, 1962. Revisional study of masarid wasps, pp.
3-27 (reclassification, key to subfamilies, phylogeny). —Charnley, 1973. Buffalo Soc. Nat.
Sci., Bul. 26: 1-79 (value of propodeal orifice and male genitalia in higher classification).

Biology: Spradbery, 1973. Wasps, 408 pp., 28 pls., 131 text figs. (natural history of British
Vespoidea).
Family MASARIDAE

Revision: Bradley, 1922. Calif. Univ., Pubs. Ent. 1: 361-464, 15 pls. (N. Amer. spp.).
—Richards, 1962. Revisional study of masarid wasps, 294 pp., 241 figs. (species of world).

Taxonomy: Ashmead, 1902. Canad. Ent. 34: 219-221 (key to genera). —Bequaert, 1929.
Psyche 36: 72-76, 81-88 (generic characters, distribution, phylogeny). —Bequaert, 1929.
Psyche 36: 366-368 (fossil record).

Biology: Bequaert, 1929. Psyche 36: 76-80 (nesting habits, larval food).
SUBFAMILY EUPARAGIINAE

This is the most primitive of the masarid wasps. The single included genus Euparagia is the
only masarid known to prey upon insect larvae as food for its young.

Genus EUPARAGIA Cresson

Euparagia Cresson, 1879. Acad. Nat. Sci. Phila., Proc. (Ent. Sect.). 6: vi.
Type-species: Euparagia scutellaris Cresson. Monotypic.

Plesiomasaris Cameron, 1904. Amer. Ent. Soc., Trans. 30: 266.
Type-species: Plesiomasaris maculiceps Cameron. Monotypic.

1470 Hymenoptera in America North of Mexico

The species, which are all rare, are usually collected in association with Eriogonum and
generally on dry hillsides.

Revision: Bohart, 1948. Pan-Pacific Ent. 24: 149-154.

Taxonomy: Bequaert, 1928. Ann. and Mag. Nat. Hist. (10)2: 146 (generic syn.). —Bohart,
1938. Pan-Pacific Ent. 14: 136-139 (synopsis). —Torchio, 1970. Los Angeles Co. Mus.,
Contrib. Sci. 202: 24-25, figs. 30-33 (larva).

boregoensis Bohart. Calif. (southeastern); desert species, Lower Sonoran Zone.
Euparagia boregoensis Bohart, 1948. Pan-Pacifie Ent. 24: 152. d, °.
desertorum Bohart. Calif.; desert species, Lower Sonoran Zone.
Euparagia desertorum Bohart, 1948. Pan-Pacifie Ent. 24: 151. 6, @.
platiniceps Bohart. Calif.; desert species, Lower Sonoran Zone.
Euparagia platiniceps Bohart, 1938. Pan-Pacifie Ent. 14: 138. 3.
richardsi Bohart, n. name. Tex., n. Mex., Ariz.
Psiloglossa simplicipes Rohwer, 1909. Ent. News 20: 357. 2. Preoce. in Euparagia.
scutellaris Cresson. Calif.; Transition Zone. Prey: Ceutorhynchus sp.; Anthonomus sp. Makes
shallow burrows in hard ground, topped with slender curved chimneys, burrows ending
in one or more cells provisioned with weevil larvae.
Euparagia scutellaris Cresson, 1879. Acad. Nat. Sci. Phila., Proc. (Ent. Sect.) 6: vi. d, °.

Biology: Williams, 1927. Pan-Pacifie Ent. 4: 38-39 (nest and prey). —Clement and Grissell,
1968. Pan-Pacifie Ent. 44: 34-37 (nest and prey).
timberlakei Bohart. Calif., Nev., Ariz.; desert species, L. Sonor. Zone.
Euparagia timberlakei Bohart, 1948. Pan-Pacifie Ent. 24: 150. 6, @.

SUBFAMILY MASARINAE

Pseudomasaris is the only masarine genus occurring in North America.

Genus PSEUDOMASARIS Ashmead

Pseudomasaris Ashmead, 1902. Canad. Ent. 34: 221.
Type-species: Masaris occidentalis Cresson. Orig. desig.
Toryna Bradley, 1922. Calif. Univ., Pubs. Ent. 1: 407.
Type-species: Masaris vespoides Cresson. Orig. desig.
Holopticus Bradley, 1922. Calif. Univ., Pubs. Ent. 1: 408.
Type-species: Masaris texanus Cresson. Orig. desig.
Cotyledon Bradley, 1922. Calif. Univ., Pubs. Ent. 1: 408.
Type-species: Masaris edwardsii Cresson. Orig. desig.

Nests are made of sand and mud, or mud alone, and are built on the upper or lower surfaces of
stones or on twigs. Typically, they occur in clumps, and are provisioned with pollen and nectar.
It appears that earlier flower records indicating that Pseudomasaris species are polytropic,
visiting a wide range of flowers, may be erroneous. More recent investigations suggest that at
least some of the species are oligotropic, visiting only a small number of closely related plants
for pollen and nectar.

Revision: Richards, 1963. Calif. Univ., Pubs. Ent. 27: 283-310, 3 pls., 26 text figs.

Taxonomy: Richards, 1962. Revisional study of masarid wasps, pp. 276-281, figs. 236-241 (key
to spp., generic synonymy). —Richards, 1966. Roy. Ent. Soc. London, Proce. 35: 47-55
(revised key).

Biology: Cooper and Bequaert, 1951 (1950). Psyche 57: 137-142 (flower visiting records).
—Cooper, 1952. Amer. Midland Nat. 48: 103-110 (oligotropy, flower visiting records).
—Torchio, 1970. Los Angeles Co. Mus., Contrib. Sci. 202: 25-27 (nests).

basirufus Rohwer. Ariz., Calif. Pollen: Phacelia crenulata, P. pachyphylla.

Pseudomasaris zonalis basirufus Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 452. 9.
Pseudomasaris bariscipus Bradley, 1922. Calif. Univ., Pubs. Ent. 1: 418. ¢.

Taxonomy: Richards, 1963. Calif. Univ., Pubs. Ent. 27: 299 (synonymy).

Superfamily VESPOIDEA 1471

coquilletti Rohwer. Ariz., Utah, Calif., Oreg. in U. Sonoran and Transition Zones. Ecology:
Builds mud nest of 2-5 cells on rocks. Parasite: Anthrax 7. irroratus Say. Pollen:
Phacelia distans, P. californica, P. imbricata, P. ramosissima, P. cicutaria; Ceanothus
sp.; Eriodictyon sp.; Eschscholtzia sp.
Pseudomasaris coquilletti Rohwer, 1911. U. S. Natl. Mus., Proe. 40: 555. 6, @.

Biology: Richards, 1963. Calif. Univ., Pubs. Ent. 27: 295, pls. 1-3 (nest).
edwardsii (Cresson). Wash., Idaho and Wyo., south to Calif. and Ariz., mostly in Transition
Zone; Mexico (Baja California). Ecology: Builds mud nests of 1-10 cells attached to
rocks, usually in somewhat sheltered sites. Parasite: Chrysuwra densa (Cr.). Pollen:
Eriodictyon tomentosum; Phacelia distans, P. crenulata, P. imbricata, P. hastata, P.
sp.; Ceanothus parviflorus, C. sp.; Tamarix gallica; Oenothera clavaeformis var.
aurantiaca, O. sp.; Mentha pulegium, Salvia columbariae; Cryptantha intermedia, C.
muricata, Symphoricarpos sp.; Chaenactis glabriuscula; Eriodictyon trichocalyx var.
lanatum. Predator: Philanthus zebratus nitens (Bks.).
Masaris edwardsui Cresson, 1872. Amer. Ent. Soc., Trans. 4: 87. 3, 9.

Taxonomy: Torchio, 1970. Los Angeles Co. Mus., Contrib. Sci. 202: 23-24, figs. 26-29 (larva,
pupa).

Biology: Hicks, 1929. Canad. Ent. 61: 121-123 (nest). —Torchio, 1970. Los Angeles Co. Mus.,
Contrib. Sci. 202: 2-20, figs. 1-23, 25 (nest, foraging, larval development).
macneilli Bohart. Calif., Utah Pollen: Hydrophyllum occidentale.
Pseudomasaris macneilli Bohart, 1963. In Richards, Calif. Univ., Pubs. Ent. 27: 291. d, 2.
macswaini Bohart. Southern Calif. Pollen: Phacelia sp.
Pseudomasaris macswaini Bohart, 1963. Jn Richards, Calif. Univ., Pubs. Ent. 27: 301. 6,
OF
maculifrons (Fox). Calif., Nev., Utah, Ariz., N. Mex.; Mexico (Sonora, Baja California).
Ecology: Builds delicate tubular mud cells on under surface of stones. Pollen: Phacelia
ramosissima, P. crenulata, P. distans, P. congesta var. rupestris, P. pachyphylla, P. sp.;
Prunus sp.; Astragalus sp.; Sphaeralcea sp.; Eucnide urens; Cryptantha sp.
Masaris maculifrons Fox, 1894. Calif. Acad. Sci., Proce. (2) 4: 12. 9.
Pseudomasaris albifrons Rohwer, 1912. U.S. Natl. Mus., Proe. 41: 451. d.
Pseudomasaris zonalis neomexicanus Rohwer, 1912. U. S. Natl. Mus., Proce. 41: 452. °.
Pseudomasaris (Holopticus) rohweri Bradley, 1922. Calif. Univ., Pubs. Ent. 1: 417. d.

Taxonomy: Richards, 1963. Calif. Univ., Pubs. Ent. 27: 297 (synonymy).

Biology: Parker, 1967. Pan-Pacific Ent. 43: 213-214, figs. b, c, e, f (nest, life history).
marginalis (Cresson). B. C. and Alta. south to Calif. and N. Mex. in Canadian, Transition and
U. Sonoran Zones. Pollen: Phacelia heterophylla, P. sericea, P. sp.
Masaris marginalis Cresson, 1864. Ent. Soc. Phila., Proc. 3: 677. °.
micheneri Bohart. Calif. (Inyo Co.).
Pseudomasaris micheneri Bohart, 1963. In Richards, Univ. Calif., Pubs. Ent. 27: 298, figs.
AN 2 oraoe
occidentalis (Cresson). Tex., Kans., N. Mex.
Masaris occidentalis Cresson, 1871. Amer. Ent. Soc., Trans. 3: 348. 2.
phaceliae Rohwer. Tex., N. Mex., Ariz. Ecology: Builds nests under or on side of stones.
Pollen: Phacelia congesta var. rupestris, P. popei var. similis, P. neomexicana, P. sp.
Pseudomasaris phaceliae Rohwer, 1912. U. S. Natl. Mus., Proc. 41: 450. d, °.

Taxonomy: Richards, 1966. Roy. Ent. Soe. London, Proc. 35: 49 (redescription male, female).

Biology: Parker, 1967. Pan-Pacific Ent. 43: 214-215, fig. a (nest). —Torchio, 1970. Los
Angeles Co. Mus., Contrib. Sci. 202: 21 (nest).
texanus (Cresson). Tex. Pollen: Phacelia popei.
Masaris texanus Cresson, 1871. Amer. Ent. Soc., Trans. 3: 348. d, 2.

Taxonomy: Richards, 1966. Roy. Ent. Soc. London, Trans. 35: 48-49 (redescription male,
female; clarification of distribution).

1472 Hymenoptera in America North of Mexico

vespoides (Cresson). Wash. to S. Dak. south to Calif. and N. Mex. in Transition and U. Sonoran

Zones; Mexico (Baja California). Ecology: Builds nest of hard mud with 1-13 cells placed
vertically with tops in a straight line, or occasionally of sandy material on twigs. Pollen:
Penstemon secundiflorus, P. heterophyllus P. alpinus, P. unilateralis, P. azurea, P.
glaber, P. comarrhenus, P. payetensis, p. cyaneus, P. venustus, P. spectabilis, P. laetus,
P. lyalli, P. attenuatus, P. gracilis, P. sp.; Salvia carduacea; Ranunculus sp.;
Platystemon californicum; Clarkia sp.; Nama parryi; Phacelia sp.; Aster sp., thistle.

Masaris vespoides Cresson, 1863. Ent. Soc. Phila., Proc. 2: 287. 3d, 2.

Pseudomasaris vespoides robertsoni Cockerell, 1913. Ent. Soc. Wash., Proc. 15: 107. 9.

Biology: Cockerell, 1913. Ent. Soc. Wash., Proc. 15: 107 (nest on stem). —Davidson, 1913.
South. Calif. Acad. Sci., Bul. 12: 17-18 (nests on rocks and stems). —Hicks, 1927. Canad.
Ent. 59: 75-79 (nests on rock). —Hicks, 1929. Canad. Ent. 61: 123-125 (behavior on flowers,
nests on rocks and plant stems). —Cooper, 1952. Amer. Midland Nat. 48: 103-110
(vespoides probably oligolectic on Penstemon spp.). —Torchio, 1970. Los Angeles Co. Mus.,
Contrib. Sci. 202: 21, fig. 24 (nest on peach twig). —Torchio, 1974. Pan-Pacifie Ent. 50:
226-234, 1 fig. (pollination mechanism in Penstemon).

wheeleri Bequaert. Calif. in Transition and U. Sonoran Zones; Mexico (Baja California). Pollen:
Eriodictyon tomentosum, E. crassifolium, E. californicum, E. trichocalyx var. lanatum,
E. sp.; Penstemon spectabilis, P. sp.; Yucca sp.; Hyptis emoryi; Larrea glutinosa;
Peucephyllum schotti.
Pseudomasaris wheeleri Bequaert, 1929. Psyche 36: 61. 6, 2.
zonalis (Cresson). B. C., Wash. to Mont. and Nebr., south to Calif. and Colo. in Transition,
Canadian and Hudsonian Zones. Ecology: Nest of 4 cells beneath rock overhang, the
entrance upward, the cells of sandy matrix covered with hard mud. Parasite: Chrysura
densa (Cr.). Pollen: Phacelia heterophylla, P. frigida, P. hastata, P. humilis, P.
nemoralis, P. sp.; Besseya plantaginea; Ranunculus sp.; Ceanothus parviflorus;
Penstemon sp.; Arnica sp., Encelia farinosa, Grindelia sp.
Masaris zonalis Cresson, 1864. Ent. Soe. Phila., Proc. 3: 674. 6, @.
Pseudomasaris zonalis albopictus Bohart, 1950. Biol. Soc. Wash., Proc. 63: 78. d, 9.

Taxonomy: Richards, 1963. Calif. Univ., Pubs. Ent. 27: 292 (synonymy).
Biology: Parker, 1967. Pan-Pacifie Ent. 43: 215-216, fig. ¢ (nest).

Family EUMENIDAE

The arrangement of genera adopted here is that used by van der Vecht and Fischer for the
Palaearctic fauna with interpolation in the appropriate position of the genera restricted to North
America or to the New World.

Taxonomy: Ashmead, 1902. Canad. Ent. 34: 203-210 (key to genera). —Isely, 1917. Ent. Soc.
Amer., Ann. 10: 364-366 (synopsis of N. Amer. spp.). —Bohart, 1939. Pan-Pacifiec Ent. 15:
97-98 (key to Odynerus, sens. lat. subgenera). —Bohart, 1965. Pan-Pacifie Ent. 41: 107-113
(status of spp. described by Cameron). —Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5:
173-174 (key to genera). —van der Vecht and Fischer, 1972. Hym. Cat. 8: 1-199
(Palaearctic spp.). —Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 364-372, 9 figs. (use of
chromosomes in classification).

Biology: Krombein, 1961. Wash. Acad. Sci., Jour. 51: 89-93, 6 figs. (symbiosis between
saproglyphid mites and eumenid wasps). —Cooper, 1967 (1966). Psyche 73: 238-250, 11 figs.
(egg hatch, number of instars, adult emergence from nest).

SUBFAMILY EUMENINAE
Genus ODYNERUS Latreille

Genus ODYNERUS Subgenus ODYNERUS Latreille

Odynerus Latreille, 1802-1803. Hist. Nat. Crust. Ins., v. 3, p. 362.
Type-species: Vespa spinipes Linnaeus. Desig. by Shuckard, 1837.
Odynera Illiger, 1807. Mag. Insektenk. 6: 196. Emend.

Superfamily VESPOIDEA 1473

Epipone Kirby and Spence, 1815. Introduct. to Ent., v. 1, p. 349. Preoce.

Type-species: Vespa spinipes Linnaeus. Monotypic.

Oplopus Wesmael, 1836. Acad. Sci. Belg., Bul. 3: 45. Preoce.

Type-species: Vespa spinipes Linnaeus. Desig. by Girard, 1879.
Oplomerus Westwood, 1840. Introduet. Mod. Class. Ins., v. 2 (Synopsis), p. 84. N. name.
Hoplomerus Agassiz, 1846. Nomencl. Zool., Index Univ., p. 185. Emend.
Hoplopus Agassiz, 1846. Nomencl. Zool., Index Univ., p. 186. Preoce. Emend.
Epiponus Saussure, 1875. Smithsn. Inst. Mise. Collect. 254: 363.

Type-species: Odynerus dilectus Saussure. Monotypic.

Euepipona Dalla Torre, 1904. In Wytsman, Gen. Ins., fase. 19, p. 39.

Type-species: Vespa spinipes Linnaeus. Desig. by Richards, 1937.

The nests are usually made in the ground but O. erythrogaster utilizes twigs of elderberry
(Sambucus).

Revision: Bohart, 1939. Pan-Pacific Ent. 15: 76-84 (N. Amer. spp.)

Taxonomy: Richards and van der Vecht, 1968. Ent. Ber. 28: 196 (type-species of Odynerus).
cinnabarinus Bohart. Calif., Ariz., Utah, Tex. in U. Sonor. Zone.
Odynerus cinnabarinus Bohart, 1939. Pan-Pacifie Ent. 15: 83. d, °.
dilectus Saussure. N. Y., Minn., Mont., Wyo., Colo., N. Mex., Calif., Wash., Oreg., Alaska, Alta.
in Canadian Zone. Ecology: Nests in ground burrows topped by erect turrets fashioned
of mud pellets. Prey: Hypera postica (Gyll.), H. punctata F., larvae; caterpillars.
Odynerus (E pipona) dilectus Saussure, 1870. Rev. Mag. Zool. (2) 22: 141. 6.
Biology: Linsley and Michener, 1942. Pan-Pacific Ent. 18: 27 (nest, prey).
erythrogaster Bohart. Calif. in U. Sonor. and Transit. Zones; Mexico (Baja California).
Ecology: Nests in Sambucus twigs. Parasite: Sarcophagidae sp.; Sphaeropthalma
(Photopsis) sp. Predator: Trichodes ornatus Say.
Monobia bicolor Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 421. °.
Preoce. in Odynerus.
Odynerus erythrogaster Bohart, 1939. Pan-Pacific Ent. 15: 81. d, 9. N. name.
Biology: Parker and Bohart, 1968. Pan-Pacific Ent. 44: 3 (nest, parasites, predator).

margaretellus Rohwer. Colo., Oreg., Wash.
Odynerus margaretellus Rohwer, 1915. U.S. Natl. Mus., Proc. 49: 242. d, °.

Genus PTEROCHEILUS Klug

The species are all ground nesting so far as is known and carry earth from their nests in
“baskets” formed by the long, fringed labial palpi. Caterpillars are used in provisioning.

Revision: Bohart, 1940. Ent. Soc. Amer., Ann. 33: 162-208.
Taxonomy: van der Vecht, 1966. Ent. Ber. 26: 161-162 (type-species of Pterocheilus and
Nannopterocheilus).
Genus PTEROCHEILUS Subgenus PTEROCHEILUS Klug

Pterocheilus Klug, 1805. Beitr. z. Naturkunde, v. 1, p. 143.
Type-species: Vespa phalerata Panzer. Desig. by Blanchard, 1840.
Nannopterocheilus Bluethgen, 1961. Akad. Wiss. Berlin, Abhandl. 1961: 62.
Type-species: Vespa phalerata Panzer. Orig. desig.

The typical subgenus does not occur in the New World.
Genus PTEROCHEILUS Subgenus MEGAPTEROCHEILUS Bohart

Pterocheilus subg. Megapterocheilus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 173.
Type-species: Pterochilus mirandus Cresson. Orig. desig.

This subgenus may be a synonym of Cephalochilus Bluethgen, 1939, which van der Vecht con-
siders to be of generic rank. Bohart noted that the Palaearctie gra ndis Lep. might belong to
Megapterocheilus, it is the type-species of Cephalochilus.

1474 Hymenoptera in America North of Mexico

arizonicus Bohart. Calif., Nev., Ariz., N. Mex., Tex. in L. Sonor. Zone deserts.
Pterocheilus (Megapterocheilus) arizonicus Bohart, 1940. Ent. Soe. Amer., Ann. 33: 174. 6,

OF
biplagiatus Cresson. Calif., Wash., Wyo. in U. Sonor. and Transit. Zones. Usually collected at
Phacelia.
Pterochilus (!) biplagiatus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proe., p. xix. 2? (d
misdet.).

bradleyi Bohart. Ariz. (Nogales); Mexico (Sonora).
Pterocheilus (Megapterocheilus) bradleyi Bohart, 1950. Biol. Soc. Wash., Proc. 63: 195. 3,
2 (2).
crispocornis Bohart. Calif., Nev., Utah, Ariz. in L. Sonor. Zone.
Pterocheilus (Megapterocheilus) crispocornis Bohart, 1940. Ent. Soc. Amer., Ann. 33: 178.
One
decorus decorus Cresson. Oreg., Mont., Wyo., Colo., Nebr., Ariz., Utah, Nev., Calif. in Canadian
and Transition Zones.
Pterochilus (!) decorus Cresson, 1879. Amer. Ent. Soce., Trans. 7: Proc., p. xviii. 2.
Pterochilus (!) zonatus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xviii.
Odynerus (Pachodynerus) cosmiogaster Cameron, 1905. Invertebrata Pacifica 1: 123. d.
decorus leucotaenius Rohwer. Alta., Wash., Oreg., Calif., Nev., Wyo., in Transition and
Canadian Zones.
Pterochilus (!) leucotaenius Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 554. °.
denticulatus (Saussure). Ariz., N. Mex.; Mexico.
Leptochilus denticulatus Saussure, 1855. Rev. Mag. Zool. (2) 7: 373. ¢.
Pterochilus (!) lewisii Cresson, 1868. Amer. Ent. Soc., Trans. 1: 382. °.
Pterochilus (!) aztecus Saussure, 1870. Rev. Mag. Zool. (2) 22: 141. @.
linsleyi Bohart. Western Tex., Ariz.; Mexico (Nuevo Leon).
Pterocheilus (Megapterocheilus) linsleyi Bohart, 1940. Ent. Soc. Amer., Ann. 33: 176. d, 9.
mirandus Cresson. Utah, Nev., Calif. in U. Sonor. and Transit. Zones.
Pterochilus (!) mirandus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xvii. d, 2.
Pterochilus (!) luteicollis Cameron, 1909. Pomona Col. Jour. Ent. 1: 84. 2.
nigricaudus Bohart. Calif., Ariz., Nev.
Pterocheilus (Megapterocheilus) nigricaudus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 181.
OF
Pterocheilus (Megapterocheilus) inyoensis Bohart, 1940. Ent. Soc. Amer., Ann. 33: 186. d.
oregonensis Bohart. Oreg. (Lakeview, Lake Co.).
Pterocheilus (Megapterocheilus) oregonensis Bohart, 1940. Ent. Soc. Amer., Ann. 33: 182.
9.
pedicellatus Bohart. Idaho, Calif., Ariz., Utah, Colo., N. Mex., Tex. in U. Sonoran Zone; Mexico.
Pterocheilus (Megapterocheilus) pedicellatus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 188.
Gree
quinquefasciatus Say. B. C. Alta., Wash., Oreg., Idaho, Mont., Wyo., S. Dak. to Tex., N. Mex.
Ecology: Nests in sand; the obliquely vertical burrow is divided into two horizontal
galleries ending in 1-3 cells. Prey: Noctuidae sp.
Pterochilus (!) 5-fasciatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 347. °.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. 8: 294-296, pl. 26, fig. 1 (nest, prey).
texanus Cresson. Tex., Fla., N. C. Ecology: Constructs 1-celled nest at end of vertical burrow
in sandy soil. Prey: Heliophana mitis Grt.
Pterochilus (!) texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 244. 6, @.
Biology: Evans, 1956. Ent. Soe. Wash., Proc. 58: 265-267, figs. 5, 6 (nest, prey).
trichogaster Bohart. Calif., Oreg. in Transition Zone. Ecology: Nests in sandy soil in
aggregations. Parasite: Fedtschenkia anthracina (Ashm.). Prey: Hydriomena sp.
Pterocheilus (Megapterocheilus) trichogaster Bohart, 1940. Ent. Soc. Amer., Ann. 33: 183.
Chmee

Biology: Bohart and Schuster, 1972. Pan-Pacific Ent. 48: 149 (nesting habits, prey, parasite).

Superfamily VESPOIDEA 1475

Genus PTEROCHEILUS Subgenus ONCHOPTEROCHEILUS Bohart

Pterocheilus subg. Onchopterocheilus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 191.
Type-species: Pterochilus(!) comptus Cresson. Orig. desig.
bakeri Cameron. Calif. in Transit. Zone from San Diego Co. to Butte Co.; Mexico (Baja
California).
Pterochilus (!) bakeri Cameron, 1909. Pomona Col. Jour. Ent. 1: 123. @.
comptus Cresson. Calif. in L. Sonor., U. Sonor., and Transit. Zones.
Pterochilus (!) comptus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xviii. °.
hirsutipennis Bohart. Calif., Nev., Ariz. in deserts in L. Sonoran Zone; Mexico (Baja
California).
Pterocheilus (Onchopterocheilus) hirsutipennis Bohart, 1940. Ent. Soc. Amer., Ann. 33:
200. 6, 2.
hurdi Bohart. Calif., Nev.
Pterocheilus (Onchopterocheilus) hurdi Bohart, 1950. Biol. Soc. Wash., Proc. 63: 197. d, °.
laticeps Cresson. Tex., Ariz., Calif. (Owens Valley and southern deserts) in L. Sonoran Zone;
Mexico (Baja California, Sonora, Chihuahua, Coahuila, Durango).
Pterochilus (!) laticeps Cresson, 1872. Amer. Ent. Soc., Trans. 4: 244. °.
Odynerus leucospilus Cameron, 1909. Pomona Col. Jour. Ent. 1: 133. 2. Preoce.
micheneri Bohart. Calif., Nev., Utah, Idaho.
Pterocheilus (Onchopterocheilus) micheneri Bohart, 1940. Ent. Soc. Amer., Ann. 33: 196. d,
Q.
panamintensis Bohart. Calif., Ariz.
Pterocheilus (Onchopterocheilus) panamintensis Bohart, 1940. Ent. Soc. Amer., Ann. 33:
192. 6.
pimorum (Viereck). Southern Calif., Nev., Ariz. in deserts, in L. Sonoran Zone; Mexico (Baja
California).
Odynerus pimorum Viereck, 1908. Amer. Ent. Soc., Trans. 33: 405. d.
sculleni Bohart. Ariz., Colo., Tex.
Pterocheilus (Onchopterocheilus) sculleni Bohart, 1950. Biol. Soc. Wash., Proc. 63: 196. 4,
OF
timberlakei Bohart. Calif. in U. Sonoran and Transition Zones north to Eldorado Co., Ariz.
Pterocheilus (Onchopterocheilus) timberlakei Bohart, 1940. Ent. Soe. Amer., Ann. 33: 194.
One
trachysomus Bohart. Calif. in L. Sonoran Zone of southern deserts and Owens Valley.
Pterocheilus (Onchopterocheilus) trachysomus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 193.
3 (2 misdet.).

Genus PTEROCHEILUS Subgenus MICROPTEROCHEILUS Bohart

Pterocheilus subg. Micropterocheilus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 201.
Type-species: Pterocheilus desertorwm Bohart. Orig. desig.

acuceps Bohart. Calif., Wyo.
Pterocheilus (Micropterocheilus) acuceps Bohart, 1940. Ent. Soe. Amer., Ann. 33: 208. °.

cyathopus Bohart. Calif. in Transition Zone.
Pterocheilus (Micropterocheilus) flavobalteatus cyathopus Bohart, 1940. Ent. Soc. Amer.,
Ann. 33: 206. d, @.
desertorum Bohart. Wash., Calif., Nev., Ariz., N. Mex. in L. Sonoran Zone.
Pterocheilus (Micropterocheilus) desertorum Bohart, 1940. Ent. Soc. Amer., Ann. 33: 202.
Gin, Ve
diversicolor Rohwer. Southern Calif., Nev., Ariz. in Desert of L. Sonoran Zone; Mexico (Baja
California).
Pterochilus (!) diversicolor Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 554. °.
morrisoni Cresson. Oreg., Calif., Nev., Utah, Ariz. in Transition, Canadian and Hudsonian

Zones.
Pterochilus (!) morrisoni Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proe., p. xix. d, 2.
Pterochilus (!) flavobalteatus Cameron, 1909. Pomona Col. Jour. Ent. 1: 84. 2.

1476 Hymenoptera in America North of Mexico

Taxonomy: Bohart, 1965. Pan-Pacific Ent. 41: 109 (synonymy).
paenacuceps Bohart. Wyo., Idaho, Utah.
Pterocheilus (Micropterocheilus) paenacuceps Bohart, 1950. Biol. Soc. Wash., Proc. 63: 200.
OF
provancheri albotinctus Bohart. Calif., Oreg., Wash., Wyo., Nev.
Pterocheilus (Micropterocheilus) provancheri albotinctus Bohart, 1950. Biol. Soc. Wash.,
Proc. 63: 199. ¢, 2.
provancheri provancheri (Huard). Calif., Nev., Oreg., Utah, Transition to Canadian and
Hudsonian Zones.
Odynerus truncatus Provancher, 1895. Nat. Canad. 22: 158. ¢, 2. Preoce.
Odynerus provancheri Huard, 1897. Nat. Canad. 24: 25. N. name.
Leptochilus cratocerus Cameron, 1909. Pomona Col. Jour. Ent. 1: 122. ¢.
pruinosus Cameron. Ariz., Idaho in Transition Zone.
Pterochilus (!) pruinosus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 227. 9.
seneconis Rohwer. Colo., Wyo., Mont., Idaho, Nev. in Canadian Zone.
Pterochilus(!) seneconis Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 553. d, @.
sparsipunctatus Bohart. Calif.
Pterocheilus (Micropterocheilus) sparsipunctatus Bohart, 1950. Biol. Soc. Wash., Proc. 63:
1985Gn ee
tricoloratus Bohart. Ariz. (Maricopa Co.), Calif. (San Bernardino Co.).
Pterocheilus (Micropterocheilus) tricoloratus Bohart, 1940. Ent. Soc. Amer., Ann. 33: 207.
2.

Genus LEPTOCHILOIDES Bohart

Leptochiloides Bohart, 1940. Ent. Soc. Amer., Ann. 33: 165.
Type-species: Leptochiloides utahensis Bohart. Orig. desig.

arizonae Bohart. Ariz.

Leptochiloides arizonae Bohart, 1940. Ent. Soc. Amer., Ann. 33: 166. 3.
brevicornis Bohart. South. Calif.

Leptochiloides brevicornis Bohart, 1940. Ent. Soc. Amer., Ann. 33: 167. ¢.
utahensis Bohart. Utah (Bellevue, Washington Co.).

Leptochiloides utahensis Bohart, 1940. Ent. Soc. Amer., Ann. 33: 166. °.

Genus HYPALASTOROIDES Saussure

Hypalastoroides Saussure, 1856. Etudes sur la famille des Vespides, v. 3, p. 328. Proposed
originally as a division of Alastor subg. Alastoroides Sauss.; validated by Op. 189,
Internat]. Comn. Zool. Nomencl., 1970.

Type-species: Alastor brasiliensis Saussure. Monotypice.

Revision: Bohart, 1948. Calif. Acad. Sci., Proce. (4) 24: 322-323.

Taxonomy: van der Vecht, 1967. Bul. Zool. Nomencl. 24: 31 (request for validation of
Hypalastoroides Sauss., 1856). —Internatl. Comn. Zool. Nomencl., 1970. Bul. Zool.
Nomencl. 26: 187 (validation of Hypalastoroides Sauss., type-species Alastor brasiliensis
Sauss.).
mexicanus (Saussure). Southern Tex.; Mexico to Canal Zone.

Alastor (Alastoroides) mexicanus Saussure, 1870. Rev. Mag. Zool. (2) 22: 141. @.
slevini (Bohart). Ariz. (Baboquivari Mts.); Mexico (Baja California).

Alastoroides slevini Bohart, 1948. Calif. Acad. Sci., Proc. (4) 24: 323. d, °.

Genus MICRODYNERUS Thomson

Microdynerus Thomson, 1874. Hym. Scand., v. 3, p. 58.
Type-species: Odynerus exilis Herrich-Schaeffer. Desig. by Jones, 1937.

Two North American species have been reared from nests in twigs; both species stored cater-
pillars as prey. A few Palaearctic species have been reported to nest in cavities in plants, walls
or rocks, and to provision with larvae of small Curculionidae.

Superfamily VESPOIDEA 1477

Taxonomy: Bohart, 1955. Ent. Soc. Wash., Proce. 57: 287-299, 19 figs. (review of trinodus
group). —Parker, 1970. Pan-Pacifie Ent. 46: 241-253, 52 figs. (key).
arenicolus (Bohart). Calif., Ariz., N. Mex.
Leptochilus arenicolus Bohart, 1955. Ent. Soc. Wash., Proc. 57: 299, fig. 19. d, ?.
bakerianus (Cameron). Oreg., Calif., Nev., Utah, Ariz., N. Mex., Colo., Wyo. Ecology: Nests in
Sambucus stems. Parasite: Toxophora virgata 0. S., Anthrax irroratus Say; Chrysis
parkeri Moore, C. pattoni Aar., C. stenodyneri Krom.; Mutillidae sp. Prey: Lepidoptera
larvae. Predator: Trichodes ornatus Say.
Ancistrocerus bakerianus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 220. 6.
Leptochilus occidentalis Bohart, 1955. Ent. Soc. Wash., Proc. 57: 291, figs. 3, 8, 10. d, 2.

Taxonomy: Bohart, 1965. Pan-Pacific Ent. 41: 108 (synonymy).

Biology: Parker and Bohart, 1968. Pan-Pacific Ent. 44: 3 (parasites, predator). —Parker,

1970. Pan-Pacific Ent. 46: 242-243, figs. 51, 52 (nest, life history).
bechteli (Bohart). Calif., Ariz.; deserts.

Leptochilus bechteli Bohart, 1955. Ent. Soc. Wash., Proce. 57: 294, figs. 1, 12. d, °.
cavatus (Bohart). South. Calif.

Leptochilus cavatus Bohart, 1955. Ent. Soc. Wash., Proce. 57: 295, fig. 18. d, 2.
gibboceps (Bohart). Calif.

Leptochilus gibboceps Bohart, 1955. Ent. Soc. Wash., Proc. 57: 296, fig wisi es
hurdi Parker. N. Mex., Ariz.; Mexico (Sonora, Baja California).

Microdynerus hurdi Parker, 1970. Pan-Pacific Ent. 46: 248, figs. 4, 21, 27, 34. d, 9.
inusitatus Parker. Calif.

Microdynerus inusitatus Parker, 1970. Pan-Pacific Ent. 46: 248. d, °.
lissosomus (Bohart). Calif.

Odynerus lissosomus Bohart, 1940. Pan-Pacific Ent. 16: 88. d, °.
monolobus (Bohart). Tex., Fla.

Odynerus heterospilus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 201. 2. Preoce. by

Cameron, 1907.

Leptochilus monolobus Bohart, 1951. U. S. Dept. Agr., Agr. Monog. 2: 897. N. name.
patagoniae Parker. Ariz. (Patagonia).

Microdynerus patagoniae Parker, 1970. Pan-Pacific Ent. 46: 252, fig. 23. ¢.
sayi (Cameron). Calif., Nev., Oreg., Idaho.

Ancistrocerus sayi Cameron, 1908. Amer. Ent. Soc., Trans. 34: 221. d.

Leptochilus williamsi Bohart, 1955. Ent. Soc. Wash., Proc. 57: 297, fig. 14. d, °.

Taxonomy: Bohart, 1965. Pan-Pacific Ent. 41: 112 (synonymy).
schlingeri (Bohart). Calif.
Leptochilus schlingeri Bohart, 1955. Ent. Soe. Wash., Proce. 57: 298, figs. 4, 5, 18. d, 2.
singulus (Bohart). S. Dak. south to Tex., west to Idaho, Ariz., Calif. Ecology: Nests in
Sambucus stems. Prey: Lepidoptera larvae.
Leptochilus singulus Bohart, 1955. Ent. Soc. Wash., Proc. 57: 293, fig. 11. d, °.

Biology: Parker, 1970. Pan-Pacific Ent. 46: 243 (nest, life history).

trinodus (Bohart). Calif., Ariz., N. Mex., Tex.
Leptochilus trinodus Bohart, 1955. Ent. Soc. Wash., Proc. 57: 289, figs. 2, 6, 7,9. d, 2.

umbifer (Bohart). Calif.
Leptochilus umbifer Bohart, 1955. Ent. Soc. Wash., Proce. 57: 297, fig. 16. d, °.

Genus LEPTOCHILUS Saussure

Genus LEPTOCHILUS Subgenus LEPTOCHILUS Saussure

Leptochilus Saussure, 1853. Etudes sur la famille des Vespides, v. 1, p. 233.
Type-species: Pterochilus (!) mauritanicus Lepeletier. Desig. by Ashmead, 1902.
Zendalia Robertson, 1928. Flowers and Insects, p. 12.
Type-species: Odynerus zendaloides Robertson. Desig. by Bohart, 1951
(=Leptochilus ornatus Saussure).

1478 Hymenoptera in America North of Mexico

Only the typical subgenus occurs in North America. The nests are constructed in pre-existing
cavities, usually in stems or twigs, but a Mexican species is recorded as nesting in a rock crevice,
and the type-species from Europe apparently nests only in empty snail shells. The species are
known to prey upon larvae of Coleoptera or Lepidoptera, or sometimes a combination of both.

Revision: Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 153-229, 239 figs., 18 maps (Nearctic
spp.).
SPECIES GROUP MESOLOBUS
mesolobus Parker. Southern Calif. in Mojave Desert.

Leptochilus mesolobus Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 177, figs. 33, 38, 39, 42,
43, map 16. 6, @.

SPECIES GROUP MINUTISSIMUS

Taxonomy: Bohart, 1940. Pan-Pacific Ent. 16: 87 (group characters).
gibberus Parker. Southern Ariz. in mts.
Leptochilus gibberus Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 180, figs. 34, 37, 40, 44,
45, map l. 6, 9.
minutissimus (Bohart). Tex. to Calif., Nev.; Mexico (Baja California).
Odynerus minutissimus Bohart, 1940. Pan-Pacifie Ent. 16: 87. ¢, 2.
petilus Parker. Southwest Tex.; Mexico south to Oaxaca and Puebla. Ecology: Nest in old
beetle burrow in dead twig. Prey: Leaf-mining larvae of Coleoptera.
Leptochilus petilus Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 178, figs. 35, 36, 41, 46, 47,
map l. 6, °.

Biology: Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 166, fig. 4 (nest, prey).
SPECIES GROUP ERUBESCENS
Both species nest in twigs and use macerated pith rather than mud in nest construction.

Taxonomy: Bohart, 1940. Pan-Pacific Ent. 16: 89 (group characters).
erubescens (Bohart). Wash. and Idaho south to Calif. and Nev. Ecology: Nests in Sambucus
stems. Prey: Larvae of Coleoptera.
Odynerus erubescens Bohart, 1940. Pan-Pacific Ent. 16: 89. 6, @.

Biology: Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 162 (nest, life history).
periallis Parker. Oreg., Calif., Nev., Utah, Ariz. Ecology: Nests in living and dead stems and
twigs. Parasite: Anthrax irroratus Say; Aritranis notata sierrae Tow.;
Microdontomerus anthidii (Ashm.); Ceratochrysis thysana Boh. Prey: Larvae of

Gelechioidea.
Leptochilus periallis Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 182, figs. 50-54, 58, 59,

map 2. d, &.
Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 160-162, fig. 2 (nest, life history).
SPECIES GROUP BELLULUS

autumnus Parker. Western Nev.
Leptochilus autumnus Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 185. 2.
bellulus (Cresson). Ga., Ala., Miss., Ark., Okla., Tex.; Mexico (Tamaulipas, Veracruz,
Guanajuato).
Odynerus bellulus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 243. 9.
Odynerus fedorensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 200. 3.
ellenae Parker. Calif., Ariz. Ecology: Nests in old beetle borings in dead Ephedra branches.
Leptochilus ellenae Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 185, figs. 60, 63-65, 71, 72,
map 3. 6, @.

Biology: Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 162 (nest).

marshi Parker. N. Mex., Ariz.
Leptochilus marshi Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 184, figs. 61, 69, 70, map 3.
on Ge

Superfamily VESPOIDEA 1479

SPECIES GROUP RUFINODUS

The species of this group build the most complex nests in the family Eumenidae. The known
species construct a linear series of cells in old borings in twigs. Each cell is closed with a com-
plex plug consisting of a layer of macerated pith above which is a layer of mud or sand. At matu-
rity the larva moves through the pith layer into the earthen layer and moves the earthen parti-
cles around, spinning them together with silk to form a cocoon.

Taxonomy: Bohart, 1940. Pan-Pacific Ent. 16: 81-82 (key to spp., group characters).
chiricahua Parker. Southern Ariz. and Calif. Ecology: Nests in Sambucus stem traps.
Parasite: Macrosiagon c. cruentum (Germ.); Sphaeropthalma amphion (Fox). Prey:
Larvae of leaf-mining Coleoptera.
Leptochilus chiricahua Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 190, figs. 91, 97, 98,
map 6. 6, °.

Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 166 (nest, prey, parasite).
ornatus Saussure. Mass. to Fla., Nebr., Colo., Kans., Tex. Ecology: Nests in old borings in

twigs of Rhus and Sambucus. Parasite: Toxophora amphitea Wlkr.; Rhydinofoenus
tarsatorius (Say); Epistenia osmiae Ashm., E. coeruleata Westw.; Ceratochrysis
enhuycki Coop. Prey: Chalepus scapularis Oliv.?, C. dorsalis Thunb. ?; Brachys ovatus
complex; Prionomerus calceatus (Say); all are leaf-mining beetle larvae.

Leptochilus Ornatus Saussure, 1853. Etudes sur la famille des Vespides, v. 1, p. 236. 2, d.
Preoce. in Odynerus.

Odynerus republicanus Dalla Torre, 1889. Wien. Ent. Ztg. 8: 125. N. name.

Odynerus zendaloides Robertson, 1901. Amer. Ent. Soc., Trans. 27: 197. 6.

Biology: Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 400-404 (nest, life history). —Krombein,
1959. Ent. News 70: 17-23 (nest, life history).
rubicundulus (Bohart). Calif.; Mexico (Baja California).
Odynerus rubicundulus Bohart, 1940. Pan-Pacifie Ent. 16: 90. d, 2.
rufinodus (Cresson). Wash. to Calif., east through Idaho to Wyo., western Kans. and N. Mex.
Ecology: Nests in stems of Sambucus, Foeniculum, Cirsium, Tetradymia, Nama.
Parasite: Anthrax irroratus Say; Clistopyga canadensis Prov.; Eurytoma stigmi Ashm.;
Epistenia odyneri Ashm., E. sp.; Ceratochrysis tuberella Boh., Chrysis derivata Buyss.,
C. parkeri Moore?
Odynerus rufinodus Cresson, 1868. Amer. Ent. Soc., Trans. 1: 381. °.
Odynerus rufobasilaris Ashmead, 1896. Psyche 7: 335. °.
Odynerus bruesi Cameron, 1909. Pomona Col. Jour. Ent. 1: 81. d.

Biology: Davidson, 1896. Psyche 7: 335-336 (nest). —Parker, 1966. Ent. Soe. Amer., Misc.
Pub. 5: 163-165, figs. 5, 6 (nest, life history).
trachysomus (Bohart). Utah, Ariz., N. Mex., Tex.; Mexico. Ecology: Nests in trap-stems.
Odynerus trachysomus Bohart, 1940. Pan-Pacifie Ent. 16: 84. d, °.
washo Parker. Calif., Nev., Utah, Ariz., N. Mex., western Tex.; Mexico (Chihuahua). Ecology:
Nests in stems of Argemone, Stanleya, Sambucus. Parasite: Macrosiagon c. cruentum
(Germ.); Phaenacra sp.; Anthrax irroratus Say; Sphaeropthalma (Photopsis) sp.;
Chrysis pattoni Aar. Prey: Leaf-mining larvae of Exema sp.
Leptochilus washo Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 189, figs. 79, 80, 87, 88, 93,
map 4. 6, @.
Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 165 (nest, life history).
SPECIES GROUP ACOLHUUS

Taxonomy: Bohart, 1940. Pan-Pacific Ent. 16: 85 (characters of tylocephalus group).
acolhuus (Saussure). N. J. to Fla., Ala., Tex. to Ariz.; Mexico to Costa Rica. Prey: Leaf-mining

larvae of Gracilariidae from Galactica.
Odynerus (Odynerus) acolhuus Saussure, 1857. Rev. Mag. Zool. (2) 9: 280. °.
Odynerus tylocephalus Bohart, 1940. Pan-Pacific Ent. 16: 85. d, °.
Odynerus monotylus Bohart, 1940. Pan-Pacifie Ent. 16: 86. d, °.

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 10 (prey).

1480 Hymenoptera in America North of Mexico

californicus Parker. Southern Calif. in mts.
Leptochilus californicus Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 195, figs. 104, 110,

115, 129, 130, map 7. 6, °.
SPECIES GROUP PAIUTE

paiute Parker. Nev., Calif.
Leptochilus paiute Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 199, figs. 1382, 135, 136, 138,
139, 147, 148, map 9. 3, °.
stangei Parker. N. Mex. (Jemez Springs); Mexico (Guanajuato). Ecology: Nests in hole in rock.
Leptochilus stangei Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 199, figs. 133, 149, 151,
map 9. 6, 2.
Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 166 (nest).
SPECIES GROUP IRWINI
irwini Parker. Southern Tex. to Calif., Nev. Ecology: Nests in old Harmolita galls on Hilaria.
Leptochilus irwini Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 199, figs. 131, 137, 140, 144,
146, map 9. 6, 2.
Biology: Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 166-167 (nest).

krombeini Parker. Fla. (Highlands Co.). Ecology: May nest in dead twigs of live oak.
Leptochilus krombeini Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 201, figs. 134, 141, 143,

map) 9G) Se
Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 167 (nest).
SPECIES GROUP ELECTUS

crocatus Parker. Ariz. (Huachuca Mts.); Mexico (Morelos, Guerrero, Puebla).
Leptochilus crocatus Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 210, figs. 158, 159, map
12iGpas.
electus (Cresson). S. C., Fla., Kans., Colo., Tex., N. Mex., Ariz., Nev., Calif.; Mexico (Chihuahua,
Sonora, Sinaloa, Zacatecas). Ecology: Nests in small branches of dead composite in
desert. Parasite: Microdontomerus anthidii (Ashm.). Prey: Larvae of Lepidoptera.
Odynerus electus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 248. 6.

Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 167, fig. 3 (nest, parasite).
ferrugineus Parker. Ariz. to Tex. in Chihuahuan Desert.
Leptochilus ferrugineus Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 206, figs. 156, 157,
170, map 12. dg, 2.
levinodus Bohart. Calif. (Needles); Mexico (Baja California).
Leptochilus levinodus Bohart, 1948. Calif. Acad. Sci., Proc. (4) 24: 324. d, @.
menkei Parker. Nev., Ariz., Calif.; Mexico (Baja California). Ecology: Nests in beetle burrow
in Ephedra, and in gall on Tetradymia.
Leptochilus menkei Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 206, figs. 1, 152, 153, 164,
166 167e le liosnmapel canon or

Biology: Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 167 (nest).

michelbacheri Bohart. Calif.; Mexico (Baja California).
Leptochilus michelbacheri Bohart, 1948. Calif. Acad. Sci., Proc. (4) 24: 326. d, 2.

tosquineti (Cameron). Calif., Nev. Ecology: Nests in twigs of Sambucus and in gall on
Tetradymia. Prey: Yponomeutidae, larvae.
Odynerus tosquineti Cameron, 1909. Pomona Col. Jour. Ent. 1: 82. ¢.

Biology: Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 167 (nest, prey).
SPECIES GROUP CHICHIMECUS

boharti Parker. Calif.
Leptochilus boharti Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 217, figs. 210, 211, 226,

229, 233, map 18. 6, @.

Superfamily VESPOIDEA 1481

brachialis Parker. Southern Tex.; Mexico (Chihuahua, Queretaro, Zacatecas, Durango).
Leptochilus brachialis Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 215, figs. 214, 215, 230,
Zo0uMapel lace
chichimecus (Saussure). Southern Ariz. and Tex. south to El Salvador.
Odynerus chichimecus Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 280. 2.
dolius Parker. Ariz. in Sonoran Desert; Mexico (Sonora, Sinaloa).
Ln dolius Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 216, figs. 194, 195, 227, map
UZ Big LE
humerus Parker. Calif., Nev., Ariz., Tex.
Leptochilus humerus Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 220, figs. 202, 203, 220,
map 10. 6, °.
labrosus Parker. Southern Ariz. and Calif.
Leptochilus labrosus Parker, 1966. Ent. Soc. Amer., Mise. Pub. 5: 218, figs. 196, 197, 218,
232, map 18. 6, °.
milleri Parker. Southern Calif. in Colorado Desert.
Leptochilus milleri Parker, 1966. Ent. Soc. Amer., Misc. Pub. 5: 222, figs. 200, 201, 228, 231,
map 10. 6, °.
monticolus Parker. Nev., Ariz., N. Mex.
Leptochilus monticolus Parker, 1966. Ent. Soc. Amer. , Mise. Pub. 5: 220, figs. 204, 205,
221, map 10. 6, 9.
propodealis Bohart. Calif.; Mexico (Baja California).
Leptochilus propodealis Bohart, 1948. Calif. Acad. Sci., Proe. (4) 24: 325, figs. 4-6. d, °.

Genus CEPHALODYNERUS Parker

Cephalodynerus Parker, 1965. Ent. Soc. Amer., Ann. 58: 364.
Type-species: Cephalodynerus wnicornis Parker. Orig. desig.

Taxonomy: Parker, 1965. Ent. Soc. Amer., Ann. 58: 365 (key to spp.).
deformiceps (Bohart). Southern Ariz.
Odynerus deformiceps Bohart, 1942. Pan-Pacific Ent. 18: 153, figs. 1, 8, 11, 12. d, @.
russipes (Bohart). Calif., Nev. in U. Sonor. Zone.
Odynerus russipes Bohart, 1942. Pan-Pacific Ent. 18: 152, figs. 18, 15. d, 2.
sculleni Parker. Tex. (Big Bend Natl. Park).
Cephalodynerus sculleni Parker, 1965. Ent. Soc. Amer., Ann. 58: 365, figs. 4, 9, 14. d.
unicornis Parker. Ariz., Utah.
Cephalodynerus unicornis Parker, 1965. Ent. Soc. Amer., Ann. 58: 365, figs. 1, 6, 15, 16, 19.
Oty ae
vanduzeei (Bohart), n. comb. (R. M. Bohart and J. van der Vecht). Western Tex., N. Mex.,
Ariz.; Mexico (Baja California). Ecology: Nests in borings in wood. Prey: Caterpillars.
Stenodynerus vanduzeei Bohart, 1948. Calif. Acad. Sci., Proe. (4) 24: 327. d, °.

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 140 (nest, prey, life history).
Genus SMERINGODYNERUS Snelling

Smeringodynerus Snelling, 1975. Ent. Soc. Wash., Proc. 77: 56.
Type-species: Odynerus morelios Saussure. Orig. desig.

Taxonomy: Snelling, 1975. Ent. Soc. Wash., Proc. 77: 56-58, 7 figs.
morelios (Saussure). Ariz., N. Mex., western Tex.; Mexico (temperate parts).
Odynerus (Odynerus) Morelios Saussure, 1857. Rev. Mag. Zool. (2) 9: 276.
Odynerus Moreliae (!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 186.
Odynerus Morelii Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 299. 6. Emend. of
morelios.
Odynerus Morelos (!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 299.
Ancistrocerus (?) nigro-hirsutus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 203. °.
Odynerus (Stenodynerus) canamexicus Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 448. ¢.

1482 Hymenoptera in America North of Mexico

Genus DOLICHODYNERUS Bohart

Odynerus subg. Dolichodynerus Bohart, 1939. Pan-Pacifie Ent. 15: 101.
Type-species: Odynerus (Dolichodynerus) turgiceps Bohart. Orig. desig.
tanynotus (Cameron). Calif., Ariz., N. Mex., Tex.; Mexico (Nuevo Leon).
Odynerus (?) tanynotus Cameron, 1909. Pomona Col. Jour. Ent. 1: 133. 2.
turgiceps (Bohart). Calif., Ariz., N. Mex.
Odynerus (Dolichodynerus) turgiceps Bohart, 1939. Pan-Pacifie Ent. 15: 102, figs. 4-8. 3,
Ox

vandykei Bohart. Ariz.
Dolichodynerus vandykei Bohart, 1950. Biol. Soc. Wash., Proc. 63: 80. ¢, 2.

Genus MARICOPODYNERUS Viereck

Odynerus subg. Maricopodynerus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 397.
Type-species: Odynerus (Maricopodynerus) maricoporum Viereck. Monotypic.

One species has been reared from stems of Sambucus. It is presumed that all species nest in
abandoned burrows or cavities in stems or twigs. The prey is unknown.

Revision: Bohart, 1950. Brooklyn Ent. Soe., Bul. 45: 17-25.

chisosensis Bohart. Western Tex.

Maricopodynerus-chisosensis Bohart, 1950. Brooklyn Ent. Soc., Bul. 45: 20. 6, °.
decorabilis Bohart. Oreg., Nev., Calif.

Maricopodynerus decorabilis Bohart, 1950. Brooklyn Ent. Soc., Bul. 45: 22. 6, @.
lissus Bohart. Ariz., southeastern Calif.

Maricopodynerus lissus Bohart, 1950. Brooklyn Ent. Soc., Bul. 45: 23. 5, @.
maricoporum (Viereck). Calif., Nev., Ariz., N. Mex., Tex.; Mexico (Baja California).

Odynerus (Maricopodynerus) maricoporum Viereck, 1908. Amer. Ent. Soc., Trans. 33: 397.

é.
Odynerus chelonogastrus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 202. 2.
rudiceps Bohart. Nev., Calif., Idaho, Colo., Mont., Oreg., Wash., Utah, Ariz.
Maricopodynerus rudiceps Bohart, 1950. Brooklyn Ent. Soc., Bul. 45: 21. d, °.
sericifrons Bohart. Southern Calif., Ariz. Ecology: Nests in Sambucus stems.
Maricopodynerus sericifrons Bohart, 1950. Brooklyn Ent. Soc., Bul. 45: 19. d, 2.

Biology: Parker and Bohart, 1968. Pan-Pacific Ent. 44: 2 (nest).

shannoni Bohart. Wash.
Maricopodynerus shannoni Bohart, 1950. Brooklyn Ent. Soc., Bul. 45: 24. d.

Genus STENODYNERUS Saussure

Stenodynerus Saussure, 1863. Soc. Phys. Hist. Nat. Geneve, Mem. 16: 228. Proposed
originally as a division of Odynerus subg. Leionotus Sauss.; validated by Op. 893,
Internat]. Comn. Zool. Nomencl., 1970.

Type-species: Odynerus chinensis Saussure. Desig. by Bohart, 1939.

Nannodynerus Bluethgen, 1938. Konowia 16: 281.

Type-species: Lionotus teutonicus Bluethgen. Orig. desig.

Most species utilize as nesting sites abandoned borings of other insects in twigs, stems or
wood, or old mud-dauber nests. A few make burrows in soil for a nesting site; in at least one spe-
cies a mud turret is constructed over the burrow entrance. Most species prey upon caterpillars,
but two of the ground-nesting species use larvae of both Lepidoptera and Coleoptera.

Revision: Bohart, 1943. Brooklyn Ent. Soc., Bul. 38: 6-11 (Anormis group). —Bohart, 1944.
Pan-Pacific Ent. 20: 69-75 (Fundatus group). —Bohart, 1948. Fla. Ent. 31: 71-77 (Fla. spp.).
—Bohart, 1949. Ent. Soc. Wash., Proce. 51: 237-253 (Ariz. spp.).

Taxonomy: van der Vecht, 1966. Ent. Ber. 26: 163 (generic synonymy). —van der Vecht,
1967. Bul. Zool. Nomencl. 24: 31 (request for validation of Stenodynerus Sauss.).
—Internatl. Comn. Zool. Nomencl., 1970. Bul. Zool. Nomencl. 26: 187 (validation of
Stenodynerus Sauss., type-species Odynerus chinensis Sauss.).

Superfamily VESPOIDEA 1483

ammonia ammonia (Saussure). Fla., S. C.
Odynerus (Ancistrocerus) ammonia Saussure, 1852. Etudes sur la famille des Vespides, v.
ep al22 se
Odsneniis ammonis Smith, 1857. Cat. Hym. Brit. Mus., v. 5, p. 83. Emend.
Odynerus floridanus Robertson, 1901. Amer. Ent. Soc., Trans. 27: 197. °.
ammonia paraensis (Saussure). U. S. west to Tex., Kans., Minn.
Odynerus (Ancistrocerus) paraensis Saussure, 1855. Etudes sur la famille des Vespides, v.
3, p- 207. do.
Odynerus clypeatus Robertson, 1901. Amer. Ent. Soc., Trans. 27: 196. ¢, 2. Preoce. N. syn.
(J. van der Vecht).
Odynerus clypeolatus Dalla Torre, 1904. In Wytsman, Gen. Ins., fase. 19, p. 42. N. name.
N. syn. (J. van der Vecht).
Odynerus bradleyi Robertson, 1925. Amer. Ent. Soc., Trans. 51: 82. N. name. Preoce. N.
syn. (J. van der Vecht).
anatropus Bohart. Calif., southern Oreg., Nev. Ecology: Observed nesting in loose gravelly
dirt on a 45 degree slope, the cells made of dirt stuck to a silk lining.
Stenodynerus anatropus Bohart, 1944. Pan-Pacifie Ent. 20: 71. ¢, °.
anormiformis (Viereck). Southeastern Calif., Ariz.
Odynerus (Stenodynerus) anormiformis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 407. 2.
anormis (Say). Transcontinental and widespread in U. S. and Canada. Ecology: Reared from
cells in a pithy stem, the cells separated by mud partitions. Parasite: Amobia sp. Prey:
Caterpillars.
Eumenes anormis Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 346. 6.
Odynerus oculatus Say, 1837. Boston Jour. Nat. Hist. 1: 385. d.
Odynerus (Odynerus) Persecutor Saussure, 1855. Etudes sur la famille des Vespides, v. 3,
p. 256. 6.
Odynerus (Ancistrocerus) philetas Cameron, 1908. Amer. Ent. Soc., Trans. 34: 214. 2.
Ancistrocerus truncatus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 217. 2. Preoce.
Odynerus approximatus Cameron, 1909. Pomona Col. Jour. Ent. 1: 79. °.
Odynerus crassispinus Cameron, 1909. Pomona Col. Jour. Ent. 1: 130. d.

Biology: Peckham and Peckham, 1905. Wasps, Social and Solitary, p. 91 (nest, life history,
prey). —Rau and Rau, 1918. Wasp Studies Afield, pp. 331-332 (nest, parasite).
apache Bohart. Ariz., N. Mex., Tex., Colo., Wyo., Utah, Nev., Calif., Oreg.; Mexico.
Stenodynerus (Stenodynerus) apache Bohart, 1949. Ent. Soc. Wash., Proc. 51: 244, fig. 1. 6,
oF
australis (Robertson). Fla.
Odynerus australis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 197. d, °.
beameri Bohart. Fla. Ecology: Nests in borings in wood. Parasite: Toxophora amphitea Wlkr.,
Anthrax a. argyropygus Wied. Prey: Lepidoptera larvae.
Stenodynerus (Stenodynerus) beameri Bohart, 1948. Fla. Ent. 31: 75.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 126-128 (nest, prey, life history,
parasites).
blandoides blandoides Bohart. Western U. S. except Ariz., east to N. Dak., Wyo., Colo., N.
Mex. Ecology: Nests in Sambucus stems.
Stenodynerus blandoides blandoides Bohart, 1943. Brooklyn Ent. Soc., Bul. 38: 9. d, 2.

Biology: Parker and Bohart, 1968. Pan-Pacific Ent. 44: 3 (nest).
blandoides owensi Bohart. Calif., Nev.
Stenodynerus blandoides owensi Bohart, 1943. Brooklyn Ent. Soc., Bul. 38: 10. d.

blandus blandus (Saussure). B. C., Wash., Idaho, Oreg., Calif., Ariz., Utah; Mexico (Baja
California). Ecology: Nests in Sambucus stems.
Odynerus (Odynerus) blandus Saussure, 1870. Rev. Mag. Zool. (2) 22: 105. °.
Odynerus mathewi Cameron, 1906. Entomologist 39: 268. d.
Odynerus giffardi Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 237. 9.

Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 93 (nest).

1484 Hymenoptera in America North of Mexico

blandus catalinae Bohart. Calif.
Stenodynerus blandus catalinae Bohart., 1943. Brooklyn Ent. Soe., Bul. 38: 10. 6, 2.
blepharus Bohart. Md. south to Fla., west to W. Va., Tex., Kans.
Stenodynerus (Stenodynerus) blepharus Bohart, 1953. Biol. Soc. Wash., Proc. 66: 185. d, °.
canus canus Bohart. B. C., Wash., Idaho, Mont., Wyo., Colo., Utah, Nev. Ecology: Nests in
crevices in voleanice outcroppings with entrance capped by mud turret. Prey:
Gelechiidae, Gracilariidae.
Stenodynerus canus canus Bohart, 1966. Biol. Soc. Wash., Proc. 79: 73, figs. 1-4. d, 2.

Taxonomy: Clement, 1973 (1972). Pan-Pacific Ent. 48: 274-275, figs. 2-5 (larva).

Biology: Clement, 1973 (1972). Pan-Pacific Ent. 48: 271-274, fig. 1 (nest, prey, life history).
canus helvolus Bohart. Oreg., Calif., Nev., Ariz.
Stenodynerus canus helvolus Bohart, 1966. Biol. Soc. Wash., Proc. 79: 76. 5, @.
chisosensis Bohart. Tex.
Stenodynerus chisosensis Bohart, 1966. Biol. Soc. Wash., Proc. 79: 79. 3, °.
claremontensis (Cameron). Calif., Nev., Oreg., Idaho. Ecology: Nests in ground and constructs
mud turret over entrance. Parasite: Senotainia trilineata (Wulp), Amobia floridensis
(Tns.); Toxophora virgata O. S.; Tetrastichus sp.; Chrysis venustella Boh. Prey:
Smicronyx sp.; Hispinae, probably Xenochalepus sp.; Plutella maculipennis (Curt.);
Carposinidae sp.; Gnorimoschema sp.; Yponomeutidae sp.
Odynerus (Stenodynerus) claremontensis Cameron, 1905. Invertebrata Pacifica 1: 122. °.
Odynerus mediatus Cameron, 1909. Pomona Col. Jour. Ent. 1: 79. °.

Biology: Markin, 1965. Pan-Pacific Ent. 41: 139 (nest, behavior). —Markin and Gittins, 1967.
Univ. Idaho, Res. Bul. 74: 1-24, 13 figs. (nest, prey, life history, parasites).
cochisensis (Viereck). B. C., Wash., Idaho and Nebr., south to Calif., Ariz. and N. Mex.
Ecology: Nests in twigs and Sambucus stems.
Odynerus (Stenodynerus) cochisensis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 395. d, ?.
Ancistrocerus pelias Cameron, 1908. Amer. Ent. Soc., Trans. 34: 208. °.
Ancistrocerus pilias (!) Cameron, 1909. Pomona Col. Jour. Ent. 1: 78.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nest).
coniodes Bohart. Calif., Nev., Utah, Idaho, N. Mex., Tex. Ecology: Nests in Tetradymia stems.
Stenodynerus coniodes Bohart, 1966. Biol. Soc. Wash., Proc. 79: 75.
Stenodynerus conoides (!) Bohart, 1966. Biol. Soc. Wash., Proc. 79: 79, figs. 17-20. 3, °.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest).

foxensis Bohart. Ariz., N. Mex.
Stenodynerus foxensis Bohart, 1944. Pan-Pacifie Ent. 20: 73. d, °.
fundatiformis fundatiformis (Robertson). S. C., Ga., Fla., Ala., Miss., Tex., Ark., Mo. Ecology:
Nests in sandy soil but does not construct a turret above entrance. Parasite:
Bombyliidae sp.; Miltogrammini spp.? Prey: Chlamisus sp., Chlamisinae sp.; Pyralididae
spp.; Gelechiidae sp.; Olethreutidae sp.; Synchlora aerata (F.); Psychidae sp.
Odynerus fundatiformis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 197. 3.

Biology: Evans, 1956. Ent. Soc. Wash., Proc. 58: 268, figs. 3, 4 (nest, prey). —Krombein, 1964.
Amer. Mus. Novitates 2201: 4-10 (nest, prey, life history, parasites).
fundatiformis gonosceles Bohart. U.S. west to Mississippi Valley, mostly in central and
northern states.
Stenodynerus fundatiformis gonosceles Bohart, 1944. Pan-Pacific Ent. 20: 74. 3, °.
histrionalis histrionalis (Robertson). U. S. west to Tex., Okla., Kans. Ecology: Nests in
borings in wood. Parasite: Pseudoxenos robertsoni Pierce; Amobia erythrura (Wulp);
Chrysis stenodyneri Krom. Prey: Olethreutidae sp.; Gelechiidae sp.; Tortricidae sp.
Odynerus histrionalis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 196. d, ?.

Biology: Krombein, 1955. Ent. Soe. Wash., Proc. 57: 147-148 (nest, prey, life history).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 134-136 (nest, prey, life history,
parasites).

Superfamily VESPOIDEA 1485

histrionalis paenevagus (Viereck). Tex., Kans., Colo.
Odynerus paenevagus Viereck, 1906. Amer. Ent. Soc., Trans. 34: 224. 9.
Nortonia (2) basimacula Cameron, 1908. Amer. Ent. Soe., Trans. 34: 224. 9.
histrionalis rufustus Bohart. Fla. Ecology: Nests in borings in wood. Parasite: Amobia
erythrura (Wulp); Chrysis stenodyneri Krom. Prey: Olethreutidae sp.; Gelechiidae sp.;
Tortricidae sp.
Stenodynerus histrionalis rufustus Bohart, 1974. Kans. Ent. Soc., Jour. 47: 468. 6, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 134-136 (nest, prey, parasite,
cocoon, life eycle; misdet. as ammonia histrionalis (Robt.)).
hoferi Bohart. Southern Ariz.
Stenodynerus (Stenodynerus) hoferi Bohart, 1949. Ent. Soc. Wash., Proc. 51: 246, fig. 3. d,
9

hybogaster Bohart. Mo., Kans. and Colo. south to Miss. and N. Mex.
Stenodynerus hybogaster Bohart, 1966. Biol. Soc. Wash., Proc. 79: 76, figs. 5-8. d, °.
innobilis Bohart. Southern Calif.
Stenodynerus innobilis Bohart, 1966. Biol. Soc. Wash., Proc. 79: 77, figs. 15, 16. d, °.
kennicottianus antheus (Cameron). West. U.S. east to Wyo., Utah, N. Mex.
Ancistrocerus antheus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 210. d.
Ancistrocerus tityrus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 211. 2.
Ancistrocerus satyrus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 211. 6.
kennicottianus kennicottianus (Saussure). Transcont. in U. S. and Canada except southwest.
Odynerus (Odynerus) Kennicottianus Saussure, 1870. Rev. Mag. Zool. (2) 22: 104.
krombeini Bohart. N. C. (Kill Devil Hills). Ecology: Nests in wood borings in sandy areas.
Parasite: Pyemotes ventricosus (Newp.); Miltogrammini sp.; Chrysis stenodyneri Krom.
Prey: Gelechiidae sp.; Olethreutidae sp.
Stenodynerus (Stenodynerus) krombeini Bohart, 1953. Biol. Soc. Wash., Proc. 66: 187. 6,
oF

Biology: Krombein, 1955. Ent. Soc. Wash., Proc. 57: 148 (nest, life history, parasite).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 128-130 (nest, prey, life history,
parasites).

lindemanni (Cameron). Tex. (Lee Co.).
Ancistrocerus lindemanni Cameron, 1908. Amer. Ent. Soc., Trans. 34: 219. d.
lineatifrons Bohart. Fla., S. C., N. C. Ecology: Nests in borings in wood in sandy areas.
Parasite: Toxophora amphitea Wlkr.; Chrysis stenodyneri Krom. Prey: Olethreutidae
sp.; Rhyacionia frustrana (Comst.), Tortricidae sp.; Gelechiidae sp.; Olethreutidae sp.
Stenodynerus (Stenodynerus) lineatifrons Bohart, 1948. Fla. Ent. 31: 76. d, °.

Biology: Krombein, 1953. Ent. Soc. Wash., Proc. 55: 115 (prey). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 136-140 (nest, prey, life history, parasites).
lissolobus Bohart. Ariz., N. Mex.; Mexico
Stenodynerus (Stenodynerus) lissolobus Bohart, 1949. Ent. Soc. Wash., Proc. 51: 246, fig. 8.
One:
lixovestis Bohart. Calif. (southern), Ariz., N. Mex., western Tex.
Stenodynerus (Stenodynerus) lixovestis Bohart, 1949. Ent. Soc. Wash., Proc. 51: 247, fig. 4.
Gave
lucidus (Rohwer). Que., Ont., B. C., Calif., Oreg., Wash., Idaho, Utah, Colo., Wyo., Mont., Minn.
Ancistrocerus lucidus Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 235. d.
microstictus (Viereck). U. S. east to Missouri River, north to S. Dak., Wyo., Calif.; Mexico.
Ecology: Nests in ground and constructs an entrance turret to burrow. Prey:
Phaloniidae sp.; Gelechiidae sp.; Cosmopterygidae sp.
Odynerus microstictus Viereck, 1906. Amer. Ent. Soc., Trans. 32: 199. °.
Odynerus (Stenodynerus) gulielmi Viereck, 1908. Amer. Ent. Soc., Trans. 33: 408. 2.
Ancistrocerus (Nortonia) phoenixensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: ZOO Ol
Q.

Biology: Evans, 1956. Ent. Soc. Wash., Proc. 58: 269, figs. 7, 8 (nest, prey, life history)

1486 Hymenoptera in America North of Mexico

noticeps clarki Bohart. Calif.

Stenodynerus noticeps clarki Bohart, 1948. Calif. Acad. Sci., Proc. (4) 24: 331. d, °.
noticeps noticeps Bohart. Calif., Nev., Oreg., Wash., Utah, Idaho, Wyo., Colo.

Stenodynerus noticeps noticeps Bohart, 1948. Calif. Acad. Sci., Proc. (4) 24: 330. d, °.
ochrogonius Bohart. Ariz., N. Mex.

Stenodynerus ochrogonius Bohart, 1944. Pan-Pacific Ent. 20: 74. d, 9.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 155-156, fig. 3 (sleeping habits).
oculeus illinoensis (Robertson). IIl., Minn., Mich., N. Y., D. C., Va., northern Fla., La., Tex.
Odynerus illinoensis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 197. 6, 2.
oculeus oculeus (Robertson). Fla., Ga.
Odynerus oculeus Robertson, 1901. Amer. Ent. Soc., Trans. 27: 197. 3, 2.
opalinus Bohart. Calif., Ariz., N. Mex.
Stenodynerus opalinus Bohart, 1966. Biol. Soc. Wash., Proc. 79: 80, figs. 10, 11. d, 9.
painteri Bohart. Southern Ariz.
Stenodynerus (Stenodynerus) painteri Bohart, 1949. Ent. Soc. Wash., Proc. 51: 248. d.
papagorum papagorum (Viereck). Sask., Idaho, Wyo., Colo., Kans., N. Mex., Utah, Ariz.;
Mexico. Ecology: Nests gregariously in soil, the burrow entrance capped by a mud
turret, stores 14-19 coleopterous leafminers per cell. Prey: Noctuidae sp.; Anthonomini
sp.; Xenochalepus sp. Predator: Philanthus pulcher D. T.
Odynerus (Stenodynerus) papagorum Viereck, 1908. Amer. Ent. Soc., Trans. 33: 394. 6, 9.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 256-271 (nest, prey, life history). —Evans,
1970. Mus. Compar. Zool., Bul. 140: 479 (nest, prey, parasites?).
papagorum tinctifer Bohart. Wyo., Utah, Calif.
Stenodynerus papagorum tinctifer Bohart, 1944. Pan-Pacific Ent. 20: 71. d, °.
patagoniensis Bohart. Southern Ariz.
Stenodynerus (Stenodynerus) patagoniensis Bohart, 1949. Ent. Soc. Wash., Proc. 51: 249,
fig. 7. 3.
percampanulatus (Viereck). Western Tex., Okla., Kans., N. Mex., Ariz., Utah, Wyo., Idaho,
eastern Wash.; Mexico.
Odynerus percampanulatus Viereck, 1906. Amer. Ent. Soe., Trans. 32: 200. d.
Odynerus (Stenodynerus) blawus Rohwer, 1915. U. S. Natl. Mus., Proc. 49: 240. @.
Odynerus (Stenodynerus) odontoschius Rohwer, 1915. U.S. Natl. Mus., Proc. 49: 241. °.
Odynerus (Stenodynerus) odontoschius var. dichrous Rohwer, 1915. U. S. Natl. Mus., Proce.
49: 241. 9.
propinquus (Saussure). Ala., Miss., La., Tex., Ark., Kans., Mo., Okla., N. Mex. Parasite:
Pseudoxenos fundati Pierce.
Odynerus (Odynerus) propinquus Saussure, 1870. Rev. Mag. Zool. (2) 22: 104. 9.
Odynerus fundatus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 242. 2, 6.
pulvinatus pulvinatus Bohart. Mass. south to Fla., Mo., Kans., Mich.
Stenorynerus (!) (Stenodynerus) pulvinatus Bohart, 1953. Biol. Soc. Wash., Proc. 66: 186.
Gq 2.
pulvinatus surrufus Krombein. Fla. Ecology: Nests in borings in wood. Parasite: Anthrax a.
argyropygus Wied.; Miltogrammini sp.; Chrysis inaequidens Dahlb. Prey: Gelechiidae
sp.; Olethreutidae sp.; Salebriaria sp , Phycitinae sp.
Stenodynerus (Stenodynerus) pulvinatus swrrufus Krombein, 1959. Ent. Soe. Wash., Proc.
61: 149. 3, °.

Biology: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 150 (nest, prey, life-history, parasite).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 130-133 (nest, prey, life history,
parasites).

pulvivestis Bohart. Ariz., Calif. in desert; Mexico.
Stenodynerus (Stenodynerus) pulvivestis Bohart, 1949. Ent. Soc. Wash., Proc. 51: 249, fig.
Os Cy Se
rudus Bohart. Ariz., southern Calif., eastern Wash., Wyo.
Stenodynerus (Stenodynerus) rudus Bohart, 1949. Ent. Soe. Wash., Proce. 51: 250. 6, @.

Superfamily VESPOIDEA 1487

sonoitensis Bohart. Southern Ariz.
Stenodynerus (Stenodynerus) sonoitensis Bohart, 1949. Ent. Soe. Wash., Proe. 51: 251, fig.
ONGane
superpendentis Bohart. Ariz., Idaho, western Tex., Nev.
Stenodynerus (Stenodynerus) superpendentis Bohart, 1949. Ent. Soe. Wash., Proc. 51: 252,
fie 2 Ga oe
toas (Cresson). Kans., Nebr., S. Dak., Wyo., Colo., Tex., N. Mex., Ariz., Calif. (San Diego Co.);
Mexico. Parasite: Pseudoxenos neomexicanus Pierce. Predator: Philanthus zebratus
nitens (Bks.).
Odynerus toas Cresson, 1868. Amer. Ent. Soc., Trans. 1: 381. 6, °.
Odynerus taos (!) Cresson, 1872. Amer. Ent. Soc., Trans. 4: 243. Emend.
Odynerus (Pachyodynerus (!)) cressoni Cameron, 1908. Amer. Ent. Soc., Trans. 34: 198. 2,
OF
Odynerus vegasensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 210. 3.
Odynerus pallidipictus Cameron, 1909. Pomona Col. Jour. Ent. 1: 83. 2.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 156 (sleeping habits).
valliceps Bohart. U.S. east to Minn., Kans., western Tex.; Mexico (Sinaloa).
Stenodynerus valliceps Bohart, 1948. Calif. Acad. Sci., Proce. (4) 24: 329. 3, 9.
ventones (Cameron). Tex. (Fedor).
Odynerus ventones Cameron, 1908. Amer. Ent. Soc., Trans. 34: 210. ¢.
williamsi Bohart. Ariz., N. Mex.
Stenodynerus (Stenodynerus) williamsi Bohart, 1949. Ent. Soc. Wash., Proe. 51: 253, fig. 5.
Grime
xanthianus (Saussure). Southern Calif.; Mexico (Baja California).
Odynerus (Odynerus) Xanthianus Saussure, 1870. Rev. Mag. Zool. (2) 22: 103. 9.
Odynerus Xantianus (!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 204.

Genus PARANCISTROCERUS Bequaert

Ancistrocerus subg. Parancistrocerus Bequaert, 1925. Amer. Ent. Soc., Trans. 51: 64.
Type-species: Odynerus (Ancistrocerus) fulvipes Saussure. Orig. desig.

Most species make their nests in abandoned borings of other insects in twigs, stems or wood,
in artificial borings in wood, in abandoned mud-dauber nests and insect galls, and in abandoned
borings of ground-nesting wasps or bees in the ground. One species constructs multicellular mud
nests attached to branches or twigs. The species have been reported as preying only upon larvae
of Lepidoptera.

All species have a complex symbiotic relationship with saproglyphid mites, each wasp having a
host-specific mite. Each species of wasp has developed an acarinarium, a chamber at the base of
the second abdominal tergum which is covered by the apex of the first tergum. The hypopial
stage of the mite congregates in large numbers in the acarinarium.

Revision: Bohart, 1948. Fla. Ent. 31: 71-74, 77-80 (Fla. spp.). —Bohart, 1949. Ent. Soe. Wash.,
Proc. 51: 237-241, 253-259 (Ariz. spp.). —Bohart, 1952. Ent. Soc. Wash., Proc. 54: 38-53
(Calif. spp., key to U.S. spp.).

acarigaster (Bohart). Wash., Oreg., Idaho, Utah, Calif., Nev., Ariz. Ecology: Nests in
Sambucus stems and in abandoned mud-dauber nests. Parasite: Chrysis parkeri Moore.
Stenodynerus (Parancistrocerus) acarigaster Bohart, 1952. Ent. Soc. Wash. Proce. 54: 49,
1H (sh Cy Se
Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nests, parasite).
acarophorus (Bohart). Wash., Oreg., Calif., Nev. Ecology: Nests in Sambucus stems. Parasite:
Chrysis coerulans F.
Stenodynerus (Parancistrocerus) acarophorus Bohart, 1952. Ent. Soc. Wash., Proce. 54: 45,
figs Zuo ee
Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nest, parasite).

austrinus (Cresson), n. comb. Tex., Okla., Kans., N. Mex.; Mexico.
Odynerus austrinus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 248. d, 2.

1488 Hymenoptera in America North of Mexico

Odynerus excentralis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 198. °.
bicornis bicornis (Robertson), n. comb. Fla.
Odynerus bicornis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 196. d, °.
bicornis ceanothi (Rohwer), n. comb. N. Y., Pa., Va., S. C., Mich., Tenn., Ark.
Ancistrocerus (Stenancistrocerus) ceanothi Rohwer, 1912. U.S. Natl. Mus., Proce. 51: 449.
Onna
bicornis cushmani (Bohart), n. comb. Western Tex., N. Mex., southern Ariz. Ecology: Nests in
borings in wood. Parasite: Saproglyphidae sp.; Chrysididae sp.
Stenodynerus (Parancistrocerus) bicornis cushmani Bohart, 1949. Ent. Soe. Wash., Proc.
51: 253. 3, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 143-144 (nest, life history,
parasites).
coronado (Bohart), n. comb. Ariz., N. Mex., Utah, Colo., western Tex.
Stenodynerus (Parancistrocerus) coronado Bohart, 1949. Ent. Soc. Wash., Proce. 51: 255. 6,
OF
cotti (Bohart), n. comb. Calif., Oreg.
Stenodynerus (Parancistrocerus) cotti Bohart, 1952. Ent. Soc. Wash., Proc. 54: 52, fig. 11.
Onace
declivatus (Bohart), n. comb. South. Calif.; Mexico (Baja California).
Stenodynerus declivatus Bohart, 1948. Calif. Acad. Sci., Proc. (4) 24: 331, pl. 12, figs. 7, 8.
Onnes
Taxonomy: Bohart, 1952. Ent. Soc. Wash., Proc. 54: 46, fig. 3.
fulvipes fulvipes (Saussure), n. comb. U.S. west to Mich., Ill., Kans., Okla., Tex. Ecology:
Nests in borings in wood, in old mud-dauber nests, in abandoned borings of
ground-nesting bees, and also constructs its own burrows in ground. Parasite: Anthrax
a. argyropygus Wied.; Vespacarus fulvipes Bak. and Cunl. Prey: Cymolomia sp.,
Tortricidae sp.; Characoma nilotica Rog.
Odynerus flavipes Lepeletier, 1841. Hist. Nat. Ins. Hym., v. 2, p. 659. d. Preoce.
Odynerus (Ancistrocerus) fulvipes Saussure, 1855. Etudes sur la famille des Vespides, v. 3,
p. 205. d, 2. N. name.

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 340-344 (nests, prey). —Rau, 1935.
Brooklyn Ent. Soe., Bul. 30: 112 (nest, prey). —Krombein, 1967. Trap-nesting wasps and
bees, pp. 154-156, pl. 21, figs. 102, 104-107, pl. 22, figs. 109, 110 (nest, prey, life history,
parasites).

fulvipes rufovestis (Bohart), n. comb. Fla., S. C. Ecology: Nests in borings in wood. Parasite:

Vespacarus rufovestis Bak. and Cunl.; Senotainia trilineata (Wulp) (?). Prey: Pyrausta
tyralis (Guen.), Pyraustidae sp.; Trichotaphe sp.; sp. of Olethreutidae or Phaloniidae.

Stenodynerus (Parancistrocerus) fulvipes rufovestis Bohart, 1948. Fla. Ent. 31: 78. d, ?.

Biology: Evans, 1956. Ent. Soc. Wash., Proc. 58: 270 (nest, prey). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 156-157 (nest, life history, prey, parasites).
histrio (Lepeletier), n. comb. D. C. and Va. to Fla., La. Ecology: Nests in borings in wood.
Parasite: Vespacarus histrio Bak. and Cunl.; Pseudoxenos louisianae (Pierce); Chrysis
stenodyneri Krom. Prey: Caterpillars.
Odynerus histrio Lepeletier, 1841. Hist. Nat. Ins. Hym., v. 2, p. 638. d.

Biology: Krombein, 1955. Ent. Soc. Wash., Proc. 57: 148-149 (nest, prey, life history, parasite).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 145-146 (nest, prey, life history,
parasite).

leionotus leionotus (Viereck). Ont., U. S. except Pacific Coast states.
Odynerus leionotus Viereck, 1906. Amer. Ent. Soc., Trans. 32: 198. @.

leionotus yumus (Viereck). Western Tex., N. Mex., Ariz., Colo.
Odynerus (Stenodynerus) ywmus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 397. ¢.
Odynerus (Ancistrocerus) arizonaensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 208. ?.

macfarlandi (Cameron). Ariz., N. Mex.; Mexico.
Odynerus macfarlandi Cameron, 1909. Pomona Col. Jour. Ent. 1: 79. 2.

Superfamily VESPOIDEA 1489

Stenodynerus (Parancistrocerus) chiricahuae Bohart, 1949. Ent. Soe. Wash., Proce. 51: 254.
Cras
mcclayi (Bohart), n. comb. Calif., Idaho, N. Mex.
Stenodynerus (Parancistrocerus) meclayi Bohart, 1952. Ent. Soc. Wash., Proc. 54: 51, fig.
LOMAS
minimoferus (Bohart), n. comb. Oreg., Calif., Ariz., Nev., Utah, Wyo., Tex. Ecology: Nests in
old Sceliphron cells.
Stenodynerus (Parancistrocerus) minimoferus Bohart, 1949. Ent. Soc. Wash., Proce. 51:
ZDGsOn) ee

Taxonomy: Bohart, 1952. Ent. Soc. Wash., Proc. 54: 47, fig. 5 (male redescription).
parapedestris arenosus (Bohart), n. comb. Calif., Nev., Ariz.
Stenodynerus (Parancistrocerus) parapedestris arenosus Bohart, 1952. Ent. Soc. Wash.,
Proc. 54: 48. 6, @.
parapedestris parapedestris (Bohart), n. comb. South. Calif.
Stenodynerus (Parancistrocerus) parapedestris Bohart, 1952. Ent. Soc. Wash., Proc. 54: 47,
fIpOmore ce
pedestris bifurcus (Robertson). Fla., Ga. Ecology: Nests in borings in wood.
Odynerus bifurcus Robertson, 1901. Amer. Ent. Soc., Trans. 27: 196. 6, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 142 (nest, life history).
pedestris pedestris (Saussure). Que., U. S. west to N. Mex., Kans., Minn. Ecology: Nests in

borings in wood, sumac and elder. Parasite: Vespacarus pedestris Bak. and Cunl.;
Pseudoxenos louisianae (Pierce); Toxophora amphitea Wlkr. Prey: Gelechiidae sp.

Odynerus (Ancistrocerus) Fuscipes Saussure, 1853. Etudes sur la famille des Vespides, v.
1, p. 143. 6. Preoce.

Odynerus (Ancistrocerus) pedestris Saussure, 1855. Etudes sur la famille des Vespides, v.
3, p. 206. N. name.

Odynerus (Ancistrocerus) Conformis Saussure, 1855. Etudes sur la famille des Vespides,
Vooapaclonig:

Odynerus (Ancistrocerus) proximus Saussure, 1857. Rev. Mag. Zool. (2) 9: 274. d.

Ancistrocerus parvispinosus Cameron, 1906. Amer. Ent. Soe., Trans. 32: 332. °.

Ancistrocerus Foxeanus Cameron, 1906. Amer. Ent. Soc., Trans. 32: 333. d.

Odynerus (Ancistrocerus) acanthopus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 223. °.

Taxonomy: Giordani Soika, 1941. Soc. Veneziana Stor. Nat., Bol. (n. s.) 2: 272 (generic
assignment).

Biology: Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 388-395 (nests, life history, parasite).
—Rau, 1935. Brooklyn Ent. Soc., Bul. 30: 110 (nest). —Krombein, 1967. Trap-nesting wasps
and bees, pp. 140-142 (nest, life history, prey, parasite).

pensylvanicus ignotatus (Bohart), n. comb. Calif., Oreg., Wash., Idaho, Minn., Mich., N. B.
Stenodynerus (Parancistrocerus) pensylvanicus ignotatus Bohart, 1952. Ent. Soe. Wash.,
Proc. 54: 46, fig. 4. d, °.
pensylvanicus pensylvanicus (Saussure), n. comb. Canada and north. U.S. south to Oreg.,
Colo., Kans., Mo., N. C., Tex., N. Mex. Ecology: Nests in borings in twigs. Parasite:
Bombyliidae sp. Prey: Tortricidae sp.; Olethreutidae sp.
Odynerus (Odynerus) Pensylvanicus Saussure, 1855. Etudes sur la famille des Vespides,
Veron ps 2ote oe
Odynerus (Odynerus) Huro Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p.
20 eaoe
Odynerus (Odynerus) Mohicanus Saussure, 1855. Etudes sur la famille des Vespides, v. 3,
p. 297. 2.
Odynerus (Odynerus) Pennsylvanicus Saussure, 1875. Smithsn. Inst., Mise. Collect. 254:
327. 6, 2. Emend.

Biology: Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 395-398 (nest, life history). —Reinhard,
1929. Witchery of wasps, pp. 221-223 (nest, prey). —Rau, 1935. Brooklyn Ent. Soce., Bul. 30:
110 (nest, parasite).

1490 Hymenoptera in America North of Mexico

perennis anacardivora (Rohwer). N. Y., Md., Va., N. C., S. C., Ga., Fla., La. Ecology: Nests in
borings in wood and twigs. Parasite: Vespacarus anacardivorus Bak. and Cunl. Prey:
Olethreutidae spp.; Gelechiidae sp.
Odynerus (Stenodynerus) anacardivora Rohwer, 1915. U.S. Natl. Mus., Proc. 49: 241. 2.
Biology: Krombein, 1955. Jn Krombein and Evans, Ent. Soc. Wash., Proc. 57: 228-229 (nest,
prey, life history, parasite). —Krombein, 1967. Trap-nesting wasps and bees, pp. 151-153
(nest, prey, life history, parasite).
perennis perennis (Saussure). U. S. west to Mississippi River, also Kans., Mo. Ecology: Nests
in borings in twigs and sumac. Prey: Blastobasidae sp.
Odynerus (Odynerus) perennis Saussure, 1857. Rev. Mag. Zool. (2) 9: 277.
Odynerus (Ancistrocerus) minnesotaensis Cameron, 1908. Amer. Ent. Soe., Trans. 34: 231.
3.

Biology: Rau, 1935. Brooklyn Ent. Soc., Bul. 30: 110 (nest). —Krombein, 1960. Ent. News 71:
33. (nest, prey, life history).

polingi (Bohart), n. comb. Southern Ariz., N. Mex.
Stenodynerus (Parancistrocerus) polingi Bohart, 1949. Ent. Soc. Wash., Proc. 51: 257. 3,
Os

rectangulis frazieri (Bohart), n. comb. Calif., Oreg.
Stenodynerus (Parancistrocerus) rectangulis frazieri Bohart, 1952. Ent. Soc. Wash., Proc.
54: 45, fig. 1. 6; 9.
rectangulis rectangulis (Viereck). Ariz., N. Mex.; Mexico. Ecology: Nests in borings in wood
and Sambucus stems. Parasite: Toxophora virgata O. S. Prey: Caterpillars.
Odynerus (Stenodynerus) rectangulis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 393. 6, @.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 153-154 (nest, prey, life history).
—Parker and Bohart, 1968. Pan-Pacific Ent. 44: 3 (nest, parasite).
salcularis rufulus (Bohart). Fla. Ecology: Nests in borings in wood and in old galls on scrub
oak. Parasite: Vespacarus saecularis Bak. and Cunl.; Megaselia aletiae (Comst.) ?;
Amobia ? sp.; Anthrax a. argyropygus Wied., Toxophora amphitea Wlkr.; Melittobia
chalybii Ashm.; Chrysis coerulans F., C. inaequidens Dahlb. Prey: Olethreutidae sp.;
Gelechiidae sp.; Rhyacionia sp., Platynota rostrana (Wlkr.), P. sp., Tortricidae sp.;
Phaloniidae sp.; Pyrausta tyralis (Guen.), Pyraustinae sp., Homoeosoma sp., Phycitinae
sp., Epipaschiinae sp. Predator: Lepidophora lepidocera (Wied.).
Stenodynerus (Parancistrocerus) saecularis (!) rufulus Bohart, 1948. Fla. Ent. 31: 79. 3,

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 146-151 (nest, prey, life history,
parasites, predator).
salcularis salcularis (Saussure). Tex., Ga., N. C., Va., D. C., Md., N. J.
Odynerus (Ancistrocerus) salcularis Saussure, 1852. Etudes sur la famille des Vespides, v.
pel22:
Odynerus (Ancistrocerus) Saecularis (!) Saussure, 1853. Etudes sur la famille des
Vespides, v. 1, p. 142. Misspelled secularis, p. 145.
Ancistrocerus leensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 215. 3.
siccus (Bohart), n. comb. Colo., Utah, Wash., Calif., Ariz.; Mexico (Baja California).
Stenodynerus (Parancistrocerus) siccus Bohart, 1952. Ent. Soc. Wash., Proc. 54: 50, fig. 9.
On eis
subtoltecus (Viereck), n. comb. Kans. (Hamilton Co.).
Odynerus subtoltecus Viereck, 1906. Amer. Ent. Soc. Trans. 32: 197. 2, (d misdet.). Type
possibly lost.
texensis (Saussure). Tex., N. Mex., Ariz., southern Calif.; Mexico. Ecology: Nests in borings in
wood and in old mud-dauber nests. Parasite: Saproglyphidae sp.
Odynerus (Odynerus) texensis Saussure, 1870. Rev. Mag. Zool. (2) 22: 104. ¢.
Odynerus (Ancistrocerus) lacunus Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 111. @.

Taxonomy: Bohart, 1952. Ent. Soc. Wash., Proc. 54: 49, fig. 7 (male characters).

Superfamily VESPOIDEA 1491

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 144-145 (nest, life history,
parasite).
toltecus (Saussure). Wash., Oreg., Idaho, Utah, Nev., Calif., Ariz., N. Mex., western Tex.:

Mexico. Ecology: Nests in borings in wood and Sambucus. Parasite: Vespacarus
toltecus Bak. and Cunl.; Toxophora virgata 0. S.; Melittobia chalybii Ashm.; Chrysis
barri Boh., C. arizonica Boh. Prey: Gelechiidae spp.; Phycitinae sp.

Odynerus (Odynerus) toltecus Saussure, 1857. Rev. Mag. Zool. (2) 9: 277. 2, d.

Odynerus Packardi Cameron, 1906. Amer. Ent. Soc., Trans. 32: 335. °.

Taxonomy: Bohart, 1952. Ent. Soc. Wash., Proc. 54: 53, fig. 12 (male characters).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nests, parasites). —Krombein,
1967. Trap-nesting wasps and bees, pp. 157-160 (nest, prey, life history, parasites).
vagus slossonae (Bohart), n. comb. Fla.
Stenodynerus vagus slossonae Bohart, 1948. Fla. Ent. 31: 77. 2.
vagus vagus (Saussure), n. comb. Ont., B. C., U. S. except Pacific Coast states. Ecology: Builds
multicellular mud nests attached to branches. Parasite: Psewdoxenos lowisianae (Pierce),
P. jonesi (Pierce); Chrysis sp.; Monodontomerus mexicanus Gahan.
Odynerus (Odynerus) vagus Saussure, 1857. Rev. Mag. Zool. (2) 9: 277.
Odynerus (Odynerus) collega Saussure, 1870. Rev. Mag. Zool. (2) 22: 61. ¢.
Odynerus colon Cresson, 1872. Amer. Ent. Soc., Trans. 4: 241. d, °.
Odynerus delodontus Viereck, 1906. Amer. Ent. Soc., Trans. 32: 196. °.

Biology: Evans, 1956. Ent. Soc. Wash., Proc. 58: 269-270 (nest, parasite).
vogti (Krombein), n. comb. Md. (Plummers Island). Ecology: Nests in borings in wood.
Parasite: Saproglyphidae sp. Prey: Gelechiidae spp.; Tortricidae sp.
Stenodynerus (Parancistrocerus) vogti Krombein, 1962. Biol. Soc. Wash., Proc. 75: 6. °.

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 151 (nest, prey, life history,
parasite).

Genus PSEUDEPIPONA Saussure

Genus PSEUDEPIPONA Subgenus PSEUDEPIPONA Saussure

Pseudepipona Saussure, 1856. Etudes sur la famille des Vespides, v. 3, p. 309. Proposed
originally as a division of Odynerus subg. Epipona Shuckard; validated by Op. 893,
Internat]. Comn. Zool. Nomencl., 1970.

Type-species: Odynerus herrichii Saussure. Monotypic.

Only the typical subgenus occurs in North America. The Palaearctic h. herrichii (Sauss.) nests
gregariously, making short burrows in firm sandy soil and preys upon larvae of Microlepidop-
tera.

Taxonomy: van der Vecht, 1967. Bul. Zool. Nomencl. 24: 31 (request for validation of
Pseudepipona Sauss., 1856). —Internatl. Comn. Zool. Nomencl., 1970 (1969). Bul. Zool.
Nomencl. 26: 187, 189, 191 (validation of Pseudepipona Saussure (misspelled Pseudopipona
on p. 187, and corrected in 1973 to Pseudepipona in Bul. Zool. Nomencl. 30: 66),
type-species Odynerus herrichii Sauss.)).

herrichii aldrichi (Fox). Alaska, B. C., Alta., Wash., Idaho, Mont., Wyo., Colo., Utah, N. Mex.
in Canadian and Transition Zones. Typical herrichii (Sauss.) is widespread in the
Palaearctic Region.
Odynerus aldrichi Fox, 1892. Ent. News 3: 197. 6, °.

Genus EUODYNERUS Dalla Torre

Some of our species belong to the subgenera Pareuodynerus and typical Huodynerus. Others
belong to one or more apparently undescribed subgenera. So far as known the subgenera
Knemodynerus, Syneuodynerus and Xanthodynerus do not occur in the New World.

Our species have quite diverse nestiag habits. A few species make original mud nests on rocks
and a few dig burrows in the soil. The majority of species nest in cavities in twigs, stems, struc-

1492 Hymenoptera in America North of Mexico

tural lumber, in old mud-dauber nests, and sometimes in old Polistes cells. The prey in all known
cases consists of caterpillars.

Euodynerus Dalla Torre, 1904. Gen. Ins., fase. 19, p. 38. Proposed originally for Sect. II of
Div. III of Odynerus subg. Leionotus Sauss.; validated by Op. 893, Internatl. Comn.
Zool. Nomencl., 1970 (1969).

Type-species: Vespa dantici Rossi. Desig. by Bluethgen, 1938.

Euodynerus subg. Pareuodynerus Bluethgen, 1938 (1937). Konowia 16: 278.

Type-species: Vespa notata Jurine. Orig. desig.
Euodynerus subg. Knemodynerus Bluethgen, 1940. Ent. Tidskr. 61: 43.
Type-species: Odynerus (Lionotus) excellens Perez. Orig. desig.
Euodynerus subg. Syneuodynerus Bluethgen, 1951. Soc. Ent. Ital., Bol. 81: 75.
Type-species: Odynerus egregius Herrich-Schaeffer. Orig. desig.
Euodynerus subg. Xanthodynerus Bluethgen, 1954. Deut. Ent. Ztschr. 1: 265.
Type-species: Odynerus (Rhynchium) octavus Giordani Soika. Orig. desig.

Taxonomy: Bequaert, 1939. Ent. Soc. Amer., Ann. 32: 61-65 (color forms of annectens
(Sauss.)). —Bohart, 1939. Brooklyn Ent. Soc., Bul. 34: 245-251 (Boscii Group). —Bohart,
1942. Pan-Pacifie Ent. 18: 145-152 (Congressus Group). —Bohart, 1948. Brooklyn Ent. Soc.,
Bul. 43: 81-87 (subspp. of foraminatus (Sauss.), fusus (Cr.), tempiferus (Vier.)). —van der
Vecht, 1967. Bul. Zool. Nomencl. 24: 29-30 (genotype). —Internatl. Comn. Zool. Nomencl.,
1970. Bul. Zool. Nomencl. 26: 187-188 (validation of Euodynerus D. T., type-species Vespa
dantici Rossi).

alvarado alvarado (Saussure). Ariz., N. Mex.; Mexico.
Odynerus (Odynerus) Alvarado Saussure, 1857. Rev. Mag. Zool. (2) 9: 276.
Odynerus Alvaradi Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 268. d, ¢. Emend.
Odynerus (Pachodynerus) halmus Cameron, 1905. Amer. Ent. Soc., Trans. 31: 389. d, °.
alvarado safranus (Bohart). Western Tex.
Rygchium alvarado safranum Bohart, 1948. Brooklyn Ent. Soc., Bul. 43: 86. d, 2.

annectens (Saussure). Va., Ga., Fla. Ecology: The only nest described was of coarse sandy clay
with 21 kidney-shaped cells surrounding a twig.
Odynerus (Odynerus) annectens Saussure, 1870. Rev. Mag. Zool. (2) 22: 59. @.
Odynerus tempiferus var. macio Bequaert, 1936. U.S. Natl. Mus., Proc. 84: 81. d, 9.

Biology: Clark and Sandhouse, 1936. U.S. Natl. Mus., Proc. 84: 89-95 (nest).

annulatus annulatus (Say). Ariz., Utah, Wyo. east to Tex., Kans., N. Dak., Alta.; north.

Mexico. Ecology: Nests in ground and builds curved mud tube above entrance; the cells
diverge leaflike from the main burrow. Parasite: Pseudoxenos hookeri (Pierce). Prey:
Loxostege sticticalis (L.).

Odynerus annulatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 348. d, °.

Odynerus (Odynerus) Bairdi Saussure, 1858. Rev. Mag. Zool. (1) 10: 169. ¢.

Odynerus verus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 237. d, 2.

Odynerus ruficandis Cameron, 1906. Amer. Ent. Soc., Trans. 32: 328. °.

Pterochilus (!) maculifrons Cameron, 1908. Amer. Ent. Soc., Trans. 34: 226. 2.

Odynerus annulatus var. birkmanni Cameron, 1909. Pomona Col. Jour. Ent. 1: 124. d, °.

Odynerus annulatus var. oslari Cameron, 1909. Pomona Col. Jour. Ent. 1: 125.

Odynerus sulciventris Cameron, 1909. Pomona Col. Jour. Ent. 1: 130. 2.

Biology: Hungerford and Williams, 1912. Ent. News 23: 250-253, pl. 14, figs. 5, 6 (nest, prey).
—Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 277-281 (nest, prey). —Rau and Rau, 1918. Wasp
Studies Afield, pp. 300-312 (nest, prey).

annulatus arvensis (Saussure). U. S. west to Tex., Okla., Kans., Nebr., S. Dak., N. Dak.

Ecology: Makes vertical ground burrows with one to six cells topped by a thick-walled
erect or bent chimney about 1 inch long. Parasite: Psewdoxenos hookeri (Pierce). Prey:
Lozxostege sticticalis (L.), Elasmopalpus furfurellus Hlst., Pyralididae spp.; Noctuidae
sp. Predator: Solenopsis sp.

Odynerus (Odynerus) arvensis Saussure, 1870. Rev. Mag. Zool. (2) 22: 59. d, °.

Odynerus cultus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 236. d.

Odynerus firmus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 239. d, 2.

Odynerus geminus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 240. 9.

Superfamily VESPOIDEA 1493

Odynerus orasus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 196. d, °.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 271-276 (nest, prey, predator). —Evans, 1956.
Ent. Soe. Wash., Proc. 58: 267-268, figs. 1, 2 (nest, prey).
annulatus evectus (Cresson). Southwest U. S. including Tex., N. Mex., Ariz., Calif.; Mexico
(Baja California).
Odynerus evectus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 235. °.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 153-154 (sleeping habits).
annulatus imperialis (Bohart). Southern Calif., Nev., Ariz., southwestern Utah; Mexico.
Rygchium sulphureum imperialis Bohart, 1945. Ent. Soc. Wash., Proc. 47: 48. d, °.
annulatus sulphureus (Saussure). Calif., Oreg., Wash., B. C., Idaho, Nev., Utah, Ariz. Parasite:
Pseudoxenos hookeri (Pierce).
Odynerus (Odynerus) sulphureus Saussure, 1858. Rev. Mag. Zool. (2) 10: 170. 9.
auranus albivestis (Bohart). Alta., Wash., Idaho, eastern Oreg., N. Dak., S. Dak., Wyo., Colo.
Odynerus boscii albivestis Bohart, 1939. Brooklyn Ent. Soc., Bul. 34: 249. d, °.
auranus aquilus Bohart. West. Kans. to west. Tex., west to Ariz.; Mexico (Coahuila, Durango).
Euodynerus auranus aquilus Bohart, 1974. Kans. Ent. Soc., Jour. 47: 463. d, °.
auranus auranus (Cameron). Calif., Oreg., Wash., Nev., Utah, Wyo., Colo. Ecology: Nests in
the ground, making clumps of five or six complete jug-shaped mud pots. Parasite:
Sphaeropthalma orestes (Fox).
Odynerus auranus Cameron, 1906. Invertebrata Pacifica, v. 1, p. 148. 6.
auranus azotopus (Bohart). Southern Nev., Ariz., southeastern Calif.; Mexico (Baja California).
Odynerus boscii azotopus Bohart, 1939. Brooklyn Ent. Soc., Bul. 34: 248. d, 2.
barberi (Bohart), n. comb. Tex. (Brownsville).
Rygchium barberi Bohart, 1945. Ent. Soc. Wash., Proc. 47: 56. °.
bidens (Saussure), n. comb. Fla., S. C., Tenn., N. J., Minn.
Odynerus (Odynerus) bidens Saussure, 1870. Rev. Mag. Zool. (2) 22: 58. d, °.
boscii boharti (Krombein). Fla.
Rygchium molestum boharti Krombein, 1959. Ent. Soc. Wash., Proc. 61: 145, fig. 1. d, °.
boscii boscii (Lepeletier). N. C. to northern Fla.
Odynerus boscii Lepeletier, 1841. Hist. Nat. Ins. Hym. 2: 687. ¢d.
boscii molestus (Saussure). Eastern U. S. to northern Fla., west to Kans., Okla. and Tex.
Ecology: Nests in borings in wood, and in burrows in structural lumber. Parasite:
Macrosiagon c. cruentum (Germ.); Chrysididae sp. Prey: Desmia funeralis (Hbn.),
Framinghamia helvalis (Wlkr.), Pyraustinae sp., Tetralopha sp., Epipaschiinae sp.
Odynerus (Odynerus) turpis Saussure, 1870. Rev. Mag. Zool. (2) 22: 60. °.
Odynerus (Odynerus) molestus Saussure, 1870. Rev. Mag. Zool. (2) 22: 61. d.
Odynerus manifestus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 240. d, 2. Preoce.
Odynerus primus Dalla Torre, 1889. Wien. Ent. Ztg. 8: 125. N. name.

Taxonomy: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 148-149 (as first reviser, synonymized
turpis under molestus).

Biology: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 101 (prey, nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 83-84 (nest, prey, life history, parasites).
castigatus castigatus (Saussure). Most of U. S., particularly more southern states; Mexico.
Predator: Philanthus zebratus nitens (Bks.).
Odynerus (Leionotus) Castigatus Saussure, 1853. Etudes sur la famille des Vespides, v. 1,
Desa oanGs
Odynerus fusus Cresson, 1872. Amer. Ent. Soe., Trans. 4: 238. d, °.
Odynerus fuscus (!) Dalla Torre, 1904. Gen. Ins., fase. 19, p. 45.
castigatus rubrivestis (Bohart). Fla., Ga.
Rygchium fusum rubrivestis Bohart, 1948. Brooklyn Ent. Soc., Bul. 4: 84. 6, °.

castigatus sanneovestis (Bohart). Calif. (Owens Valley).
Rygchium fusum sanneovestis Bonart, 1948. Brooklyn Ent. Soc., Bul. 4: 84. 6, °.

1494. Hymenoptera in America North of Mexico

cockerelli (Cameron). Pacific Coast states, Idaho, in Upper Sonoran and Transition Zones.
Ecology: Constructs vertical ground burrows as nesting sites.
Ancistrocerus cockerelli Cameron, 1908. Amer. Ent. Soc., Trans. 34: 220. @.
Odynerus infuscipennis Bohart, 1942. Pan-Pacifie Ent. 18: 146. 6, °.
congressus (Viereck), n. comb. Tex., N. Mex., Ariz., Calif., Nev., L. Sonor. Zone; Mexico.
Odynerus (Stenodynerus) congressus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 405. °.
crypticus balteatus (Say). N. Y., Pa., N. J., Va., Ind., Ohio, Mich., IIl., Tex., Utah, Ont.
Rygchium balteatum Say, 1837. Boston Jour. Nat. Hist. 1: 383. °.
Monobia Sylvatica Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 95. 2.
crypticus crypticus (Say). Ont., Alta., U. S. west to S. Dak., Wyo., Colo., Utah, Ariz.; Mexico.
Ecology: Nests in small colonies, making vertical ground burrows usually with about
three cells, one above the other, separated by clay partitions. Parasite: Bombyliidae sp.;
Miltogrammini sp.; Chrysis sp. Prey: Pyrgus c. communis (Grt.), Pholisora catullus (F.).
Odynerus crypticus Say, 1823. West. Quart. Rptr. 2: 81.
Rygchium Louisianum Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 106. 9.
Odynerus (Odynerus) Iturbide Saussure, 1857. Rev. Mag. Zool. (2) 9: 276.
Odynerus designatus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 235. °.
Odynerus Iturbidi (!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 265. d.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 281-294 (nest, prey, parasites). —Rau and
Rau, 1918. Wasp Studies Afield, pp. 312-331 (nest, prey, life history, parasite). —Turner,
1922. Biol. Bul. 42: 153-172. —Rau, 1931. Brooklyn Ent. Soc., Bul. 26: 5-6. —Vest, 1936.
Utah Acad. Sci. Arts and Letters, Proc. 13: 207-209. —Rau, 1945. Brooklyn Ent. Soc., Bul.
40: 29-30 (carnivorous habit, adult).

crypticus stricklandi (Bequaert). Alta., Wyo., S. Dak., Nebr., Colo.
Odynerus dorsalis stricklandi Bequaert, 1940. Canad. Ent. 72: 55. d, °.

delicatus (Cresson), n. comb. Tex.
Odynerus delicatus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 236. 2.
digiticornis (Bohart), n. comb. Ariz., Utah; Mexico (Baja California).
Odynerus canaliculatus Viereck, 1906. Amer. Ent. Soc., Trans. 33: 392. 6. Preoce.
Rygchium digiticornis Bohart, 1945. Ent. Soe. Wash., Proc. 47: 49. 6. N. name.
discogaster (Bequaert), n. comb. Calif., Oreg., Wash., B. C., Idaho, Nev., Ariz., Colo., Wyo.
Ecology: Makes mud nests in cracks between rocks, with about three cells in a clump.
Odynerus discogaster Bequaert, 1939. Ent. Soc. Amer., Ann. 32: 65. d, 9.

exoglyphus albovittatus (Bohart), n. comb. Great Basin area including Wash., Oreg., Calif.,
Nev., Idaho, Wyo., Colo., N. Mex.
Odynerus exoglyphus albovittatus Bohart, 1939. Brooklyn Ent. Soc., Bul. 34: 251. 3, °.
exoglyphus exoglyphus (Bohart), n. comb. Calif., Oreg., Idaho, Utah, Wyo., northern Ariz.
Odynerus exoglyphus Bohart, 1939. Brooklyn Ent. Soc., Bul. 34: 250. 6, @.
foraminatus aequalis (Cameron). Western Tex. to Ariz., Colo., Utah, Idaho, Oreg. and Wash.
in Upper Sonoran Zone. Ecology: Nests in old mud-dauber cells.
Odynerus aequalis Cameron, 1906. Amer. Ent. Soc., Trans. 32: 329. °.
Odynerus sapellvensis Cameron, 1906. Amer. Ent. Soc., Trans. 32: 331.
Odynerus sulfuritinctus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 389. d, °.
Odynerus spectabiliformis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 390. d, 2.
Odynerus hidalgiformis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 391. °.
Odynerus spectabiformis (!) Viereck, 1908. Amer. Ent. Soc., Trans. 33: 391.
Odynerus sapelloensis (!) Cameron, 1909. Pomona Col. Jour. Ent. 1: 125. d, 2.
Odynerus deficiens Cameron, 1909. Pomona Col. Jour. Ent. 1:126. ¢.
Odynerus jeromensis Cameron, 1909. Pomona Col. Jour. Ent. 1: 131. ¢.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 299 (nest).
foraminatus apopkensis (Robertson). Central to southern Fla. Ecology: Nests in borings in
wood. Parasite: Pyemotes sp.; Pseudoxenos hookeri (Pierce); Macrosiagon c. cruentwm
(Germ.); Toxophora sp.; Amobia floridensis (Tns.), A. erythrura (Wulp); Chrysis
coerulans F., C. inaequidens Dahlb., C. derivata Buyss. Prey: Olethreutidae sp.

Superfamily VESPOIDEA 1495

primarily; Platynota sp., Tortricidae sp., occasionally. Predator: Lepidophora lepidocera
(Wied.) ?

Odynerus apopkensis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 196. d.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 62-70, pl. 25, fig. 119 (nest, prey,
life history, parasites, predator).
foraminatus blakeanus (Cameron). Tex. (Lee Co.).
Odynerus blakeanus Cameron, 1909. Pomona Col. Jour. Ent. 1: 80. 2.
Rygchium rugosum fedoris Bohart, 1945. Ent. Soe. Wash., Proc. 47: 46. d, 9.
foraminatus foraminatus (Saussure). Southern Canada, U.S. west to Rocky Mts. Ecology:

Nests in hollow twigs, burrows in logs, borings in wood, and old Polistes cells. Parasite:
Anthrax irroratus Say; Megaselia aletiae (Comst.); Pseudoxenos hookeri (Pierce), P.
foraminati Pierce; Pimpla spatulata Tow.; Melittobia chalybii Ashm.; Lycogaster
pullata Shuck.; Chrysis coerulans F., C. nitidula F., C. inaequidens Dahlb. ?. Prey:
Gnorimoschema solidaginis (Riley), Gelechiidae spp.; Oecophoridae spp.; Olethreutidae
sp.; Archips sp., Enarmonia sp., Tortricidae spp.; Thyris maculata Harr.; Desmia
funeralis (Hbn.), Loxostege sp., Pyraustinae spp., Acrobasis sp., Pyralidinae sp.,
Epipaschia superatilis Clem.

Odynerus (Odynerus) Rugosus Saussure, 1853. Etudes sur la famille des Vespides, v. 1, p.
179. 3.

Odynerus (Odynerus) Foraminatus Saussure, 1853. Etudes sur la famille des Vespides, v.
Te py 180 set Se

Odynerus (Odynerus) flavopictus Saussure, 1857. Rev. Mag. Zool. (2)9: 276.

Odynerus Harringtoni Cameron, 1906. Amer. Ent. Soc., Trans. 32: 327. 2.

Odynerus santa-feae Cameron, 1906. Amer. Ent. Soc., Trans. 32: 330. 9.

Biology: Hungerford and Williams, 1912. Ent. News 23: 255 (nest). —Rau and Rau, 1918.
Wasp Studies Afield, PP. 334-340 (nest, prey, parasite). —Rau, 1922. Acad. Sci. St. Louis,
Trans. 24, no. 7: 17-18 (nest). —Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 398-400 (nest, life
history). —Rau, 1932. Ent. News 53: 119-121 (nest). —Rau, 1935. Brooklyn Ent. Soe., Bul.
30: 110-111 (cocoon). —Rau, 1944. Canad. Ent. 76: 129 (nest). —Hartman, 1944. Psyche 51:
1-4 (egg-laying). —Cooper, 1954. Ent. Soc. Wash., Proc. 56: 281-282 (nest, parasite).
—Medler, 1964. Ent. Soc. Amer., Ann. 57: 56-60 (nest, prey, life history, parasites).
—Markin, 1965. Ent. Soc. Amer., Ann. 58: 132-133, fig. 1 (nest, life history). —Krombein,
1967. Trap-nesting wasps and bees, pp. 56-62 (nest, prey, life history, parasites, predator).

foraminatus parvirudis (Bohart). Fla., S. C.
Rygchium foraminatum parvirudis Bohart, 1948. Brooklyn Ent. Soc., Bul. 4: 83. d, °.
foraminatus scutellaris (Saussure). Idaho, Oreg., Calif., Nev. Ecology: Nests in cavities in
Sambucus and in twigs, and in old mud-dauber nests. Parasite: Psewdoxenos hookeri
(Pierce); Anthrax irroratus Say; Amobia floridensis (Tns.); Epistenia sp.; Lycogaster
pullata nevadensis (Cr.); Chrysis coerulans F., C. derivata Buyss., C. parkeri Moore, C.
inaequidens Dahlb. Predator: Trichodes ornatus Say.
Odynerus (Odynerus) scutellaris Saussure, 1870. Rev. Mag. Zool. (2) 22: 58. 6.
Odynerus visellus Cameron, 1906. Invertebrata Pacifica 1: 147. ¢d.
Ancistrocerus rivularis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 215. d.
Odynerus blandinus Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 236. d, 2.

Taxonomy: Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 364-372, figs. 7, 8 (chromosome
number).
Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nest, parasites). —Parker and
Bohart, 1968. Pan-Pacifie Ent. 44: 2 (nest, predator).
guerrero (Saussure). Western Tex., N. Mex., Ariz.; Mexico (Baja California). Ecology: Nests in
borings in wood. Parasite: Chrysis arizonica Boh., C. inflata Aar. Prey: Pyraustinae sp.;
Noctuidae sp.
Odynerus (Odynerus) Guerrero Saussure, 1857. Rev. Mag. Zool. (2) 9: 277.
Odynerus Guerreri Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 294. d, °. Emend.
Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 81-83, pl. 9, fig. 41 (nest, prey, life
history, parasites).

1496 Hymenoptera in America North of Mexico

hidalgo boreoorientalis (Bequaert). East Coast states, Ohio, Mo., eastern Kans. Ecology:
Nests in borings in wood. Prey: Homoeosoma electellwm (Hlst.).
Odynerus hidalgo var. boreo-orientalis Bequaert, 1937. Pan-Pacifie Ent. 12: 11. d, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 77-79 (nest, prey, life history).
hidalgo hidalgo (Saussure). U. S. east to Miss., north to Mo., Kans., Colo., Calif.; Mexico.
Ecology: Nests in abandoned bee burrows in a cliff, in a Sceliphron mud nest, and in an
old Polistes nest. Prey: Caterpillars.
Odynerus (Odynerus) Hidalgo Saussure, 1857. Rev. Mag. Zool. (2) 9: 275.
Odynerus ductus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 238. 6, 2.
Odynerus Hidalgi Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 252. 6, 2. Emend.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 296-299 (nest, prey). —Bequaert, 1939. Ent.
Soe. Amer., Ann. 32: 68-69 (nest). —Rau, 1943. Ent. Soe. Amer., Ann. 36: 533 (nest).
—Linsley, 1962. Ent. Soc. Amer., Ann. 55: 154 (sleeping habits).

hidalgo viereckii (Cameron). Calif., Nev., Utah, Idaho, Wash., Colo. Ecology: Nests in old
Sceliphron mud nests, dividing each cell transversely in two.
Odynerus viereckii Cameron, 1909. Pomona Col. Jour. Ent. 1: 127. @.
Odynerus hidalgo var. boreo-occidentalis Bequaert, 1937. Pan-Pacific Ent. 12: 13. d, @.
leucomelas leucomelas (Saussure). North. and west. U. S. and Canada south to mountains of
Tenn., N. Mex., Ariz. Ecology: Nests in cavities in sumac twigs. Parasite: Amobza
distorta (Allen); Chrysis coerulans F., C. nitidula F. ?. Prey: Sarrothiripus sp., Palthis
angulalis (Hbn.); Rheumaptera hastata L. ?; Framinghamia helvalis (Wlkr.),
Tetralopha spp., Acrobasis spp., Dioryctria reniculella Grt., D. spp.; Pterophoridae sp.;
Olethreutidae sp.; Badebecia urticana Hbn., Exartema spp., Griselda radicana Walsh,
Rhopobota naevana geminana Steph., Pseudexentera spp., Sparganothis reticulatana
Clem., S. sp., Platynota sp., Pandemis spp., Archips argyrospilus Wlkr., A. spp., C.
rosaceana Harr., C. fumiferana (Clem.), C. pinus Free., Argyrotaenia velutinana Wlkr.
?, Acleris variana Fern., A. logiana placidana Rob., A. sp.; Recurvaria spp., Eucordylea
spp., Trichotaphe levisella Fyles ?, T. spp., Gelechiidae spp.
Odynerus (Odynerus) Leucomelas Saussure, 1855. Etudes sur la famille des Vespides, v. 3,
p: 255.6579.

Biology: Medler, 1964. Ent. News 75: 26-27 (nest, life history, parasite). —Fye, 1965. Canad.
Ent. 97: 718-722 (nest, prey, life history, parasites).
leucomelas oregonensis (Bohart). Wash., Oreg., Nev., Utah, Wyo.
Rygchium foraminatum oregonense Bohart, 1948. Brooklyn Ent. Soc., Bul. 43: 82. d, °.
macswaini (Bohart), n. comb. Calif. in Transit. Zone.
Rygchium macswaini Bohart, 1948. Brooklyn Ent. Soc., Bul. 48: 80. d, °.
martini (Bohart), n. comb. Tex., Colo., N. Mex., Ariz., Utah.
Odynerus martini Bohart, 1942. Pan-Pacifie Ent. 18: 149. d, °.
megaera (Lepeletier). East. and south. U.S. north to N. Y., Il. Pa., west to eastern Tex., Okla.
Ecology: Nests in borings in wood. Parasite: Toxophora amphitea Wlkr., Anthrax
aterrimus (Big.); Chrysis coerulans F., C. inaequidens Dahlb. Prey: Archips sp.,
Platynota sp., Tortricidae spp.; Desmia funeralis (Hbn.), Framinghamia helvalis
(Wlkr.), Tetralopha sp., Epipaschiinae sp., Nephopteryx nyssaecolella (Dyar), N. wvinella
(Rag.), Phycitinae sp.; Psilocorsis sp.; Olethreutidae sp.; Palthis angulalis Hbn.
Odynerus megaera Lepeletier, 1841. Hist. Nat. Ins. Hym., v. 2, p. 636. 2.

Biology: Krombein, 1955. Ent. Soc. Wash., Proc. 57: 146-147 (nest, prey, life history).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 70-74, pl. 9, fig. 40, pl. 26, figs. 128,
124 (nest, prey, life history, parasites).
oslarensis (Cameron). Calif., Ariz. Ecology: Nests in borings in Sambucus stems and wood.
Prey: Phycitinae sp.; Gelechiidae sp.; Cosmopterygidae sp.
Odynerus (Pachyodernus (!)) oslarensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 199.
oe

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 84-85 (nest, life history, prey).

Superfamily VESPOIDEA 1497

planitarsis (Bohart), n. comb. South. Canada, Maine, N. H., Mich., Wis., Minn., Colo.
Rygchium planitarsis Bohart, 1945. Ent. Soc. Wash., Proc. 47: 45. 3, °.
pratensis brumalis (Bequaert). Wash., Oreg., northeastern Calif., Idaho, Nev.
Odynerus pratensis var. brumalis Bequaert, 1936. U. S. Natl. Mus., Proc. 84: 86. d.
pratensis pratensis (Saussure). Ont., Southwest. U. S. east to Kans., Tex., west to Calif.;
Mexico. Ecology: Nests in Sambucus stems and wood. Parasite: Macrosiagon c.
cruentum (Germ.) ?; Toxophora sp.; Trichrysis mucronata (Br.). Prey: Caterpillars.
Odynerus (Odynerus) pratensis Saussure, 1870. Rev. Mag. Zool. (2) 22: 61. d, 9.
Odynerus clusinus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 234. 6, °.
Odynerus egregius Viereck, 1906. Amer. Ent. Soc., Trans. 32: 197. ¢. Preoce.
Odynerus congressensis Cameron, 1909. Pomona Col. Jour. Ent. 1: 132. d, °.

Biology: Parker and Bohart, 1966. Pan-Pacifiec Ent. 42: 93 (nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 79-81 (nest, prey, life history, parasites). —Parker and
Bohart, 1968. Pan-Pacifie Ent. 44: 2 (nest).

provisoreus (Viereck), n. comb. Ariz., southern Colo.
Odynerus (Stenodynerus) provisoreus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 396. d.
russatus (Bohart), n. comb. Calif., Ariz., Utah, N. Mex., Tex. in L. Sonor. Zone; Mexico
(Coahuila).
Odynerus russatus Bohart, 1942. Pan-Pacific Ent. 18: 150, figs. 2, 9, 14. d, @.
schwarzi (Krombein). N.J., Pa., Md., D. C., Va., Okla. Ecology: Nests in borings in wood.
Parasite: Toxophora amphitea Wlkr.; Melittobia chalybii Ashm.; Chrysis nitidula F.
Prey: Gelechia albisparsella (Chamb.); Psilocorsis sp.; Tortricidae sp.; Loxostege
mancalis (Led.).
Rygchium schwarzi Krombein, 1962. Biol. Soc. Wash., Proc. 75: 5. d, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 74-77, pl. 9, fig. 39 (nest, prey, life
history, parasites).
tempiferus birepandus (Bohart). Nev. (Charleston Mountain Park).
Rygchium tempiferum birepandum Bohart, 1948. Brooklyn Ent. Soe. Bul. 43: 86. 6.
tempiferus eldoradensis (Rohwer). B. C., Northwest. U.S. including Calif., Oreg., Wash., Nev.,
Idaho, Mont., Wyo., Colo.
Odynerus eldoradensis Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 238. d.
Odynerus robustus Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 238. 2. Preoce.
tempiferus pritchardi (Bequaert). Okla.
Odynerus annectens var. pritchardi Bequaert, 1939. Ent. Soc. Amer., Ann. 32: 63. 2.
tempiferus subrubeus (Bohart). Utah, Colo.
Rygchium tempiferum subrubeum Bohart, 1948. Brooklyn Ent. Soce., Bul. 43: 85. d, °.
tempiferus tempiferus (Viereck). Ariz., N. Mex., southern Colo., Utah, Wyo., S. Dak.
Odynerus (Stenodynerus) tempiferus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 392. d.
Odynerus trichiosomus Cameron, 1909. Pomona Col. Jour. Ent. 1: 127. d.
tetralobus (Bohart), n. comb. Calif.
Odynerus tetralobus Bohart, 1942. Pan-Pacifie Ent. 18: 148. d, 2.

Genus MONOBIA Saussure

Monobia Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 94.

Type-species: Vespa quadridens Linnaeus Desig. by Ashmead, 1902.
Triarthra Dalla Torre, 1904. Gen. Ins., fase. 19, p. 28. Preoce.

Type-species: Odynerus cyanipennis Guerin. Desig. by Bequaert, 1940.
Tetrarthra Dalla Torre, 1904. Gen. Ins., fase. 19, p. 28.

Type-species: Vespa quadridens Linnaeus. Desig. by Bequaert, 1940.

Most species occur in Central and South America. Nesting habits have been recorded for only
a few species. The preferred nesting sites are in pre-existing cavities. Caterpillars are stored as
prey.

Revision: Bequaert, 1940. Rev. de Ent. 11: 822-842 (New World spp.)

1498 Hymenoptera in America North of Mexico

quadridens (Linnaeus). N. H. to Fla., west to Wis., Ill., Kans. and N. Mex. Ecology: Nesting

sites are in pre-existing cavities such as carpenter bee tunnels, hollow stems, old
mud-dauber cells, and occasionally in old burrows of ground-nesting bees. The nests
consist of a linear series of cells separated by mud partitions; usually there is an empty
intercalary cell between each stored cell. Parasite: Tortonia quadridens Bak.;
Monobiacarus quadridens Bak. and Cunl.; Anthrax aterrimus (Big.); Amobia erythrura
(Wulp); Megaselia aletiae (Comst.); Pseudoxenos bishoppi (Pierce); Melittobia chalybii
Ashm.; Chrysis smaragdula F., C. inaequidens Dahlb. Prey: Nephopteryx uvinella
(Rag.), N. sp., Phycitinae sp., Epipaschia superatilis Clem., E. sp., Tetralopha
asperatella (Clem.), T. sp., Epipaschiinae sp., Desmia funeralis (Hbn.), Pyraustinae sp.;
Stenoma schlaegeri Zell., S. sp., Stenomidae sp.; Psilocorsis sp.; Gelechiidae spp.;
Platynota sp., Tortricidae spp. Predator: Lecontella cancellata (LeC.), Cleridae sp.;
Dermestidae sp.

Vespa quadridens Linnaeus, 1763. Centuria Ins. Rar., p. 31.

Vespa quadricornis (!) Degeer, 1773. Mem. pour Servir a l’Hist. des Ins., v. 3, p. 584.
Lapsus.

Vespa cincta Degeer, 1773. Mem. pour Servir a !’Hist. des Ins., v. 3, p. 584. Preoce.

Vespa uncinata Fabricius, 1775. Systema Ent. p. 367.

Vespa incincta (!) Fabricius, 1804. Systema Piezatorum, p. 259. Lapsus.

Biology: Ashmead, 1894. Psyche 7: 76-78 (nest). —Tandy, 1908. Ent. News 19: 231-232 (nest).
—Rau and Rau, 1918. Wasp Studies Afield, pp. 346-354 (nest, prey, life history). —Rau,
1922. Acad. Sci. St. Louis, Trans. 24, no. 7: 16 (nest). —Rau, 1926. Acad. Sci. st. Louis,
Trans. 25: 199-200 (nest). —Reinhard, 1929. Witchery of Wasps, p. 86 (nest). —Rau, 1981.
Brooklyn Ent. Soc., Bul. 26: 4-6 (lack of cocoon). —Rau, 1935. Ent. News 46: 57-58
(courtship, mating). —Frost, 1944. Ent. News 55: 10-14 (nest, prey, life history).
—Krombein, 1958. Ent. Soc. Wash., Proc. 60: 101 (prey). —Krombein, 1967. Trap-nesting
Wasps and Bees, pp. 46-56, pl. 7, figs. 24-29, pl. 8, figs. 30-38 (nest, prey, life history,
associates). —Byers, 1972. Kans. Ent. Soc., Jour. 45: 235-238 (supersedure in Osmia
lignaria Say nests, nest, prey).

texana (Cresson). Tex., Ariz.; Mexico (Baja California, Sonora, Veracruz).
Odynerus texanus Cresson, 1872. Amer. Ent. Soe., Trans. 4: 234. 2.
Monobia Californica Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 129. 6.
Monobia variabilis Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 137. 9.

Genus MONTEZUMIA Saussure

Montezumia Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 87.
Type-species: Montezwmia rufidentata Saussure. Desig. by Ashmead, 1902.
Alpha Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 160.
Type-species: Montezumia rufidentata Saussure. Desig. by Bohart, 1951.
Beta Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 162.
Type-species: Montezumia morosa Saussure. Desig. by Bequaert, 1921.
Metazumia Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 114.
Type-species: Montezumia huasteca Saussure. Desig. by Bequaert, 1921.
Eumontezumia Dalla Torre, 1904. Jn Wytsman, Gen. Ins., fase. 19, p. 27. N. name.

Most species are restricted to America south of U. 8. Two of the Neotropical species are sup-
posed to build mud nests, but it has been suggested for one of them that it merely used aban-
doned mud-dauber nests. One species is reported as preying upon caterpillars.

Taxonomy: Bequaert, 1921. Rev. Zool. Afric. 9: 242-247 (diagnosis of typical subgenus, list of

spp.). —Bequaert, 1940. Ent. Soc. Amer., Ann. 33: 96 (raises typical subgenus to generic
rank).

arizonensis Bequaert. Southern Ariz.
Montezumia arizonensis Bequaert, 1940. Ent. Soc. Amer., Ann. 33: 96. °.

Superfamily VESPOIDEA 1499

Genus PACHODYNERUS Saussure

Pachodynerus Saussure, 1870. Rev. Mag. Zool. (2) 22: 56. Proposed originally as a division
of Odynerus subg. Leionotus Sauss.; validated by Op. 893, Internat]. Comn. Zool.
Nomencl., 1970.

Type-species: Odynerus (Odynerus) californicus Saussure. Desig. by Bohart, 1951.

Most species nest in pre-existing cavities such as in stems, twigs, abandoned larval bagworm
cases, and abandoned mud-dauber cells, but one species (perhaps erroneously) has been reported
as also building mud nests attached to foliage. Caterpillars are stored as prey.

Taxonomy: van der Vecht, 1967. Bul. Zool. Nomencl. 24: 31 (request for validation of
Pachodynerus Sauss., 1870). —Internatl. Comn. Zool. Nomencl., 1970. Bul. Zool. Nomencl.
26: 187 (validation of Pachodynerus Sauss., type-species Odynerus californicus Sauss.).

astraeus (Cameron). Southwestern Tex. to southern Calif.; Mexico (Baja California, Hidalgo).

Ecology: Nests in cavities in wood, Sambucus stems, abandoned mud-dauber cells,
abandoned larval bagworm cases; it is also reported, perhaps in error, as building mud
nests attached to leaves of growing plants. Parasite: Sphaeropthalma (Photopsioides)
uro (Bl.). Prey: Caterpillars. Predator: Macrosiagon c. cruentum (Germ.).

Odynerus (Pachodynerus) astraeus Cameron, 1905. Amer. Ent. Soc., Trans. 31: 390. °.

Odynerus pulverulenta Viereck, 1908. Amer. Ent. Soc. Trans. 33: 406. °.

Odynerus acuticarinatus Cameron, 1909. Pomona Col. Jour. Ent. 1: 82. 9.

Biology: Rau, 1940. Ent. Soe. Amer., Ann. 33: 592 (nest). —Davis, 1964. U. S. Natl. Mus., Bul.
244: 13 (nest). —Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 93 (nest, parasite).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 85-87, pl. 24, figs. 115-118 (nest, prey,
life history, predator).

erynnis (Lepeletier). S. C. to Fla., west to La., Mass. (rarely after storm). Ecology: Nests in

pre-existing cavities such as borings in wood and abandoned cynipid galls. Parasite:
Megaselia sp.; Toxophora amphitea Whkr., Anthrax a. argyropygus Wied.; Amobia
erythrura (Wulp); Pseudoxenos erynnidis Pierce; Melittobia chalybii Ashm.; Chrysis
inaequidens Dahlb. Prey: Stenomidae spp.; Blastobasidae sp.; Platynota sp., Tortricidae
sp.; Psilocorsis sp., Oecophoridae spp.; Acrobasis sp., Etiella zinckenella (Treitschke),
Phycitinae spp.; Chrysaugidae sp.; Palthis angulalis (Hbn.).

Odynerus erynnis Lepeletier, 1841. Hist. Nat. Ins. Hym. 2: 645. °.

Odynerus (Odynerus) Erinnys (!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 245. d,
9. Emend.

Odynerus errings (!) Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 240.

Biology: Ashmead, 1894. Psyche 7: 76-78 (nest). —Krombein, 1967. Trap-nesting wasps and
bees, pp. 87-90 (nest, prey, life history, associates).
nasidens (Latreille). Southern Ariz., Tex. (Brownsville), Fla. (Plantation Key); Mexico, Central
America, Antilles; adventive in Hawaii, Micronesia. Ecology: Nests in abandoned
mud-dauber cells and makes mud cells behind wooden siding of buildings. Parasite:
Chrysis sp.; Melittobia chalybii Ashm.; Amobia floridensis (Tns.). Prey: Epipaschiidae
sp. Predator: Crematogaster spp.; Macrosiagon sp.
Odynerus nasidens Latreille, 1812. In Humboldt and Bonpland, Voy. aux Reg. Equinox,
Zool., v. 2, p. 112. @.
Odynerus (Odynerus) auratus Saussure, 1858. Rev. Mag. Zool. (2) 10: 166.
Odynerus nasidens var. minor Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 233. d, 2.
Odynerus clavilneatus Cameron, 1912. Timehri 2: 222. 2, (d misdet.).
Odynerus clavilineatus (!) Bohart, 1951. U. S. Dept. Agr., Monog. 2: 892.

Biology: Dow, 1932. Psyche 39: 12 (nest). —Rau, 1933. Jungle bees and wasps of Barro
Colorado Island, pp. 169-170 (nest, prey, parasite). —Freeman and Jayasingh, 1975. Oikos
26: 86-91, 1 fig., 4 tabs. (population dynamics, parasites, predators).

Genus PSEUDODYNERUS Saussure

Pseudodynerus Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 220. Proposed
originally as a division of Odynerus subg. Ancistrocerus Wesmael; validated by Op. 893,
Internatl. Comn. Zool. Nomencl., 1970.

1500 Hymenoptera in America North of Mexico

Type-species: Odynerus luctwosus Saussure. Monotypic.

This is a primarily Neotropical genus of eumenid wasps, only one species being known from
North America.

Revision: Bequaert, 1941. Amer. Mus. Novitates 1106: 1-10 (New World spp.).

Taxonomy: van der Vecht, 1967. Bul. Zool. Nomencl. 24: 31 (request for validation of
Pseudodynerus Sauss., 1855). —Internatl. Comn. Zool. Nomencl., 1970. Bul. Zool. Nomencl.
26: 187 (validation of Pseudodynerus Sauss., type-species Odynerus luctuosus Sauss.).
quadrisectus (Say). N. J. to Fla., west to IIl., Kans., Okla., Tex., Colo. Ecology: Nests in
cavities in wood and makes cell partitions of mud. Prey: Caterpillars.
Odynerus quadrisectus Harris, 1833. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p.
589. Nom. nud. ascribed to Say.
Odynerus quadrisectus Say, 1837. Boston Jour. Nat. Hist. 1: 385. 6, 9.
Odynerus bellone Lepeletier, 1841. Hist. Nat. Ins. Hym., v. 2, p. 660. 6, 2.

Biology: Bequaert, 1925. Amer. Ent. Soc., Trans. 51: 78 (nest). —Rau, 1935. Brooklyn Ent.
Soc., Bul. 30: 111 (nest, prey). —Krombein, 1967. Trap-nesting wasps and bees, pp. 45-46
(nest, prey, life history).

Genus PARANORTONIA Bequaert

Pachymenes subg. Paranortonia Bequaert, 1940. Ent. Soc. Amer., Ann. 33: 100. Bertoni,
1934 (Rev. Soc. Cient. Paraguay 3: 109), proposed Paranortonia as a new genus. His
name is invalid because he did not designate a type-species (Art. 13 (b), Internatl. Code
Zool. Nomencl.).

Type-species: Nortonia tolteca Saussure. Orig. desig.

symmorphus sonorensis Bequaert. Colo., Ariz.
Pachymenes symmorphus sonorensis Bequaert, 1940. Ent. Soc. Amer., Ann. 38: 101. 6, @.

symmorphus symmorphus (Saussure). U. S. west to Tex., Okla., Kans., Iowa.
Odynerus (Odynerus) Symmorphus Saussure, 1855. Etudes sur la famille des Vespides, v.
3, p. 246. 6, 2.
symmorphus toltecus (Saussure). Ariz., Tex.; Mexico (Sonora, Jalisco).
Nortonia Tolteca Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 140. d, 9.

Genus ANCISTROCERUS Wesmael

Ancistrocerus Wesmael, 1836. Acad. Sci. Bruxelles, Bul. 3: 45.
Type-species: Vespa parietum Linnaeus. Desig. by Girard, 1879.
Euancistrocerus Dalla Torre, 1904. In Wytsman, Gen. Ins., fase. 19, p. 36. N. name.

Most species of Ancistrocerus nest in pre-existing cavities, such as borings in twigs, stems and
structural lumber, in abandoned galls, in abandoned mud-dauber cells, and in old burrows of
ground-nesting wasps and bees; a few species build complete mud cells. Only lepidopterous lar-
vae have been recorded as prey of North American species, but a European species is reported
to prey upon coleopterous larvae.

Revision: Bequaert, 1944. Ent. Amer. (n. s.) 23: 225-286 (N. Amer. spp.).

adiabatus adiabatus (Saussure). Transcontinental and widespread in southern Canada and U.
S. Ecology: Nests have been reported from borings in twigs, stems and wood, galls of
other insects, old mud-dauber nests, empty sawfly cocoons, and rubber tubing. Parasite:
Vespacarus tigris Bak. and Cunl.; Anthrax irroratus Say; Amobia distorta (Allen), A.
floridensis (Allen); Pseudoxenos tigridis Pierce; Acroricnus junceus (Cr.), Ephialtes
decumbens (Tow.), E. sp., Calliephialtes notandus Cr., Argothereutes lophyri, n. subsp.;
Melittobia chalybii Ashm.; Monodontomerus dentipes (Dalm.); Chrysis coerulans F., C.
nitidula F. Prey: Psilocorsis sp., Oecophoridae sp.; Griselda radicana Wals.,
Olethreutidae spp. Coleotechnites sp., Eucordylea sp., Gelechiidae spp.; Spilonota
ocellana (D. and §.), Acleris variana Fern., Tortricidae sp.; Coleophora fletcherella
Fern.; Phaloniidae sp. Predator: Philanthus zebratus nitens (Bks.), P. solivagus Say.

Odynerus (Ancistrocerus) adiabatus Saussure, 1852. Etudes sur la famille des Vespides, v.

Tepe l22e oy

Superfamily VESPOIDEA 1501

Odynerus (Ancistrocerus) pertinax Saussure, 1855. Etudes sur la famille des Vespides, v.
3, p- 216. d.

Odynerus (Ancistrocerus) tigris Saussure, 1857. Rev. Mag. Zool. (2) 9: 273. d, 9.

Odynerus (Ancistrocerus) cervus Saussure, 1858. Rev. Mag. Zool. (2) 10: 165.

Odynerus monteregalis Meade-Waldo, 1914. Ann. and Mag. Nat. Hist. (8) 14: 405. N. name
for canadaensis, a nomen nudum.

Odynerus canadaensis Meade-Waldo, 1914. Ann. and Mag. Nat. Hist. (8) 14: 405.
Manuscript name credited to Cameron. Preoce.

Ancistrocerus howardi Cameron, 1909. Pomona Col. Jour. Ent. 1: 78. 2.

Ancistrocerus tenuatus Tucker, 1909. Kans. Acad. Sci., Trans. 22: 286. 9.

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 344-345 (nest, prey). —Champlain,
1922. Psyche 29: 99 (parasite). —Bequaert, 1925. Amer. Ent. Soc., Trans. 51: 106 (nest,
parasite). —Bequaert, 1943. Ent. Amer. (n. s.) 23: 260 (nest). —Boyce, 1946 (1945). Ent.
Soe. Ontario, Ann. Rpt. 76, pp. 35-37 (nest, prey, life history). —Krombein, 1954. Brooklyn
Ent. Soc., Bul. 49: 2-3 (nest, prey). —Coppel, 1961. Ent. News 72: 246-248, 2 figs. (nest).
—Fye, 1965. Canad. Ent. 97: 731-734 (nest, prey, life history, parasite). —Medler, 1965.
Kans. Ent. Soc., Jour. 38: 314-316 (nest, prey, life history, parasite). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 110-115 (nest, prey, life history, parasites). —Evans,
1973. Great Basin Nat. 33: 152-153 (nest, prey, parasites).

adiabatus albolacteus Bequaert. Que. to B. C., Alaska, Wash., Oreg., Idaho, Mont., S. Dak.,
Colo., Utah. Parasite: Stylopidae sp.
Ancistrocerus tigris albolacteus Bequaert, 1943. Ent. Amer. (n. s.) 23: 260. d, 2.
adiabatus cytainus (Cameron). Calif., Oreg., Nev., Colo. in Transition, Upper Sonoran and
Lower Sonoran Zones. Ecology: Nests in Sambucus stems and old mud-dauber nests.
Parasite: Chrysis coerulans F.
Odynerus cytainus Cameron, 1906. Invertebrata Pacifica 1: 149. 2.

Taxonomy: Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 364-372, figs. 1, 9 (chromosome
number).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 93 (nests, parasite).

antilope allegrus Bequaert. Calif., Oreg., Colo.
Ancistrocerus antilope allegrus Bequaert, 1943. Ent. Amer. (n. s.) 23: 276. d, 9.

antilope antilope (Panzer). Transcontinental in southern Canada and U. S. except Fla. and

Ala.; Holarctic, in the Old World throughout Europe and Siberia. Ecology: In North
America it is reported to nest in borings in wood, stems of sumac and elder, and in old
mud-dauber nests. Parasite: Kennethiella trisetosa (Coor.); Amobia distorta (Allen);
Melittobia chalybii Ashm.; Chrysis coerulans F., C. nitidula F., C. spp. Prey:
Psilocorsis sp., Oecophoridae sp.; Anacampsis (?) sp., Gelechiidae spp.; Olethreutidae
sp.; Archips cerasivorana (Fitch), A. fervidana (Clem.), A. spp., Tortricidae spp.;
Tetralopha sp. Epipaschiinae sp., Mineola indigenella (Zell.), Salebria subcaesiella
(Clem.), Nephopteryx sp., Phycitinae spp., Pyralididae spp.; Epizeuxis aemula Hbn.,
Hypeninae sp., Noctuidae spp. Only the typical subspecies of antilope occurs in the Old
World.

Vespa insolens ? Harris, 1780. Expos. Engl. Ins., p. 128, pl. 37, fig. 7. Type destroyed.

Vespa antilope Panzer, 1798. Faunae Ins. German. h. 53, pl. IX, with letterpress. 2. Type
destroyed.

Odynerus pictus Curtis, 1826. Brit. Ent., v. 3, p. 187 b, no. 2.

Odynerus (Ancistrocerus) capra Saussure, 1857. Rev. Mag. Zool. (2) 9: 273.

Ancistrocerus nearcticus Cameron, 1906. Amer. Ent. Soc., Trans. 32: 332. ¢.

Ancistrocerus lecontei Cameron, 1908. Amer. Ent. Soc., Trans. 34: 218. d, 2.

Biology: Rau and Rau, 1918. Wasp studies afield, pp. 345-346 (nest, parasite). —Taylor, 1922.
Psyche 29: 56-58 (nest, life history). —Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 405.
—Buckle, 1929. Canad. Ent. 51: 265-266 (erroneous prey record). —Reinhard, 1929.
Witchery of Wasps, pp. 86-91 (nest, parasite). —Cooper, 1953. Amer. Ent. Soc., Trans. 79:
13-35 (ecology, predation, life history, parasites). —Cooper, 1955. Amer. Ent. Soc., Trans.
80: 119-174, 1 pl., 6 text figs. (relations with mite, Kennethiella). —Medler and Fye, 1956.

1502 Hymenoptera in America North of Mexico

Ent. Soc. Amer., Ann. 49: 97-102, 2 figs. (nest, prey, life history, parasites). —Fye, 1965.
Canad. Ent. 97: 729-731 (nest, prey, life history, parasites; misidentified as c. catskill
(Sauss.)). —Krombein, 1967. Trap-nesting wasps and bees, pp. 90-98, pl. 6, fig. 23 (nest,
prey, life history, parasites).
antilope navajo Bequaert. Ariz., Calif., N. Mex., Colo., Mont. in U. Sonor. Zone. Ecology: Nests
in borings in wood.
Ancistrocerus navajo Bequaert, 1925. Amer. Ent. Soc., Trans. 51: 92. d, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 98-99 (nest, life history).

antilope spenceri Bequaert. B. C.
Ancistrocerus antilope spenceri Bequaert, 1943. Ent. Amer. (n. s.) 23: 277. 6, @.
bustamente bustamente (Saussure). Western U.S. east to western Tex. and Kans., north to
Colo., Utah, Oreg.; Mexico. Ecology: Frequents arid areas; nests in Sambucus stems.
Odynerus (Ancistrocerus) Bustamente Saussure, 1857. Rev. Mag. Zool. (2) 9: 272. 2.
Odynerus pictiventris Cameron, 1906. Amer. Ent. Soc., Trans. 32: 331. @.
Ancistrocerus neocallosus Bequaert, 1943. Ent. Amer. (n. s.) 23: 264. 6, 2.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 92 (nest).

bustamente discopictus Bequaert. Calif., Ariz.
Ancistrocerus neocallosus discopictus Bequaert, 1943. Ent. Amer. (n. s.) 23: 268. d, 2.
campestris (Saussure). U.S. east of Rocky Mts., Ont. Ecology: Nests in borings in wood and
sumac stems, and in abandoned mud-dauber nests. Parasite: Toxophora amphitea Wlkr.;
Amobia (?) sp.; Melittobia chalybii Ashm. Prey: Psilocorsis sp.; Gelechia albisparsella
(Chamb.), Gelechiidae sp. Predator: Macrosiagon cruentum (Germ.).
Odynerus (Ancistrocerus) campestris Saussure, 1852. Etudes sur la famille des Vespides,
Vols pel 2ome.

Biology: Rau and Rau, 1916. Jour. Anim. Behavior 6: 43 (nest, prey). —Snelling, 1963.
Pan-Pacifie Ent. 39: 88 (nest, predator). —Medler, 1966. Ent. Soc. Wash., Proc. 68: 143-144
(nest, life history, parasite). —Krombein, 1967. Trap-nesting wasps and bees, pp. 100-102
(nest, prey, life history, parasites).

catskill albophaleratus (Saussure). Transcontinental in Canadian and Transition Zones in

Alaska, Labrador, Canada and U. S. south to mts. of Ariz., N. Mex., N. C. Ecology:
Nests in borings in wood and sumac, in hollow stems, and in deserted galls. Parasite:
Stylopidae sp.; Melittobia sp.; Chrysis coerulans F., C. nitidula F. Prey: Noctuidae sp.;
Geometridae sp.; Badebecia near urticana Hbn., Exartema punctanum Wals., Hedia
ochroleucana Hbn., Anchylopera sp., Epiblema culminaria Wals., Gypsonoma (?) sp.,
Zeiraphera ratzeburgiana Ratz., Z. fortunana Kft., Z. sp., Griselda radicana Wals.,
Rhopobota naevana geminana Steph., Pseudexentera sp., Epinotia spp., Ptycholoma (?)
sp., Pandemis spp., Choristoneura rosaceana Harr., C. fumiferana (Clem.),
Argyrotaenia velutinana (?) Wlkr., A. spp., Acleris spp., Olethreutidae spp.; Recurvaria
spp., Hucordylea spp., Gelechiidae spp.; Oecophoridae spp.; Salebria fructella Hulst;
Pterophorus delawaricus (Zell.). Predator: Philanthus pulcher D. T., P. solivagus Say.

Odynerus (Ancistrocerus) Albophaleratus Saussure, 1855. Etudes sur la famille des
Vespides, v. 3, p. 217. d, 2.

Ancistrocerus quebecensis Cameron, 1906. Amer. Ent. Soc., Trans. 32: 333. ¢.

Odynerus (Hypodernus (!)) longipilosus Cameron, 1908. Amer. Ent. Soc. Trans. 34: 199. °.

Odynerus allophaleralus (!) Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 240.

Biology: Packard, 1869. Guide Study Ins., pp. 155-156 (nest, prey). —Taylor, 1922. Psyche 29:
60-61 (nest, prey, life history). —Bequaert, 1925. Amer. Ent. Soc., Trans. 51: 104 (nest).
—Bequaert, 1943. Ent. Amer. (n. s.) 23: 253 (nest, parasite). —Fye, 1965. Canad. Ent. 97:
723-729 (nest, prey, life history, parasites). —Krombein, 1967. Trap-nesting wasps and
bees, pp. 104-110, pl. 3, fig. 11 (nest, prey, life history, parasites; a composite account based
on nests of c. albophaleratus, c. catskill, and hybrids between them).

catskill catskill (Saussure). Southern Canada west to Alta., transcontinental in U. S. in
Transition and Austral Zones. Ecology: Nests in pre-existing borings in wood,
Sambucus, and sumac, in abandoned mud-dauber nests, and in deserted burrows of
ground-nesting bees. Parasite: Kennethiella sp., Vespacarus tigris Bak. and Cunl.;

Superfamily VESPOIDEA 1503

Toxophora virgata O. S., Anthrax a. argyropygus Wied., A. irroratus Say; Megaselia
sp.; Amobia (?) sp.; Microdontomerus anthidii (Ashm.); Chrysis coerulans F., C.
nitidula F.; Sphaeropthalma amphion (Fox), S. sp. Prey: Olethreutidae sp.; Archips sp.,
Tortricidae sp. Predator: Philanthus solivagus Say.

Odynerus (Ancistrocerus) Catskill Saussure, 1853. Etudes sur la famille des Vespides, v.
ips LS65d, 02

Odynerus (Ancistrocerus) catskillensis Saussure, 1855. Etudes sur la famille des Vespides,
v. 3, p. 204. Emend.

Odynerus Catskilli(!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 168.

Ancistrocerus sexcingulatus Ashmead, 1901. Psyche 9: 185. ¢.

Ancistrocerus ormsbyensis Cameron, 1905. Invertebrata Pacifica 1: 120. d.

Ancistrocerus gunnisonensis Cameron, 1906. Invertebrata Pacifica 1: 146. ¢.

Ancistrocerus gunnistonensis (!) Cameron, 1908. Amer. Ent. Soc., Trans. 34: 213.

Biology: Myers, 1927. Ent. Monthly Mag. 53: 190-196 (nest, parasite). —Rau, 1935. Brooklyn
Ent. Soc., Bul. 30: 111 (nest). —Rau, 1945. Ent. Soc. Amer., Ann. 38: 88 (nest). —Hobbs,
Numni, and Virostek, 1961. Canad. Ent. 93: 144 (nest). —Medler, 1964. Ent. Soc. Wash.,
Proc. 66: 210 (parasites). —Medler, 1966. Ent. Soc. Wash., Proc. 68: 139-142 (nest, prey, life
history). —Krombein, 1967. Trap-nesting wasps and bees, pp. 104-110, pl. 3, fig. 11 (nest,
prey, life history, parasites; a composite account based on nests of c. albophaleratus, c.
catskill, and hybrids between them). —Parker and Bohart, 1968. Pan-Pacifie Ent. 44: 2,
(nest, parasites).

catskill halophila Viereck. Rocky Mts. west to Pacific Coast. Ecology: Nests in borings in
twigs and Sambucus stems, in old oak galls, and possibly also in abandoned burrows of
ground-nesting bees. Parasite: Chrysis derivata Buyss. :
Ancistrocerus halophila Viereck, 1903 (1902). Acad. Nat. Sci. Phila., Proc. 54: 735. d.
Ancistrocerus trichionotus Cameron, 1905. Invertebrata Pacifica 1: 120. d.
Ancistrocerus fulvitarsis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 205. d, °.
Ancistrocerus tahoensis Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 235. °.

Biology: Bequaert, 1943. Ent. Amer. (n. s.) 23: 255 (nest). —Bohart, 1951. U. S. Dept. Agr.,
Monog. 2: 894 (nests). —Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 92-93 (nests,
parasite).

durangoensis Cameron. Western Okla. and Tex., N. Mex., Ariz., Utah, Colo., Wyo. in Upper
Sonoran Zone; Mexico. Ecology: Nests in borings in wood. Parasite: Chrysis inflata Aar.
Prey: Lepidoptera larvae.
Ancistrocerus durangoensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 216. d.
Ancistrocerus fulvicarpus Cameron, 1908. Amer. Ent. Soc., Trans. 34: 222. °.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 154-155, fig. 3 (sleeping aggregations).
—Krombein, 1967. Trap-nesting wasps and bees, p. 102 (nest, prey, life history, parasite).
lineativentris kamloopsensis Bequaert. B. C., Oreg., Wyo.
Ancistrocerus lineativentris kamloopsensis Bequaert, 1943. Ent. Amer. (n. s.) 23: 280. 6,
Pe
lineativentris lineativentris Cameron. West. U. S. east to S. Dak., west. Kans. mostly in
Transition Zone. Ecology: Nests in Sambucus stems. Parasite: Amobia floridensis
(Tns.), Pimpla spatulata Tow.; Chrysis inflata Aar.
Ancistrocerus lineativentris Cameron, 1906. Invertebrata Pacifica 1: 146. d.

Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 93 (nest, parasites).
lineativentris sinopis Bohart. Colo., west. Tex., Utah, Ariz.
Ancistrocerus lineativentris sinopis Bohart, 1974. Kans. Ent. Soc., Jour. 47: 466. 6, &.
lutonidus Bohart. Mass. to S. C., Ind., Wis., Tex. Ecology: Nests are made of hardened clay,
are attached to twigs of low bushes, and contain 6-20 or more cells arranged somewhat

radially.
Ancistrocerus lutonidus Bohart, 1974. Kans. Ent. Soc., Jour. 47: 465. d, °.

Biology: Packard, 1869. Guide Study Ins., p. 156, pl. 5, fig. 12 (nest). —Riley, 1880. Amer.
Ent. 3: 154. —Strand, 1914. Ent. Mitt. 3: 116-118. —Viereck, 1916. Conn. State Geol. and

1504 Hymenoptera in America North of Mexico

Nat. Hist. Survey, Bul. 22, pl. 4, fig. 1 (nest). Misdet. as birenimaculatus (Sauss.) in
preceding references.
paracallosus Bequaert. S. Dak., Ariz.
Ancistrocerus paracallosus Bequaert, 1943. Ent. Amer. (n. s.) 23: 269. d, @.
parietum (Linnaeus). Que., Ont., Mass. and R. I. south to D. C., Ohio, Mich.; Palaearctic in
Europe and North Africa east to Manchuria, also in the Azores and Madeira. Ecology:
Nests in crevices in mortar and glass tubing in North America; in Europe is reported to
nest in tree stumps, rock crevices, and in burrows in clay banks. Adventive from
Europe. Parasite: Stylopidae sp. Prey: Torticidae sp. in Ont.; in Europe the usual prey is
caterpillars, but there is one record of storing coleopterous larvae, Melasoma populi L.
Vespa parietum Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 572.
Odynerus ? geoffroyanus Spinola, 1808. Insectorum Liguriae 2: 182.
Odynerus flavipes Curtis, 1826. Brit. Ent. 3: 187 b, no. 12.
Odynerus (Ancistrocerus) Ochlerus Saussure, 1852. Etudes sur la famille des Vespides, v.

peel sieoe
Odynerus parietum var. incisoides Verhoeff, 1890. Ent. Nachr. 16: 335. Type probably
destroyed.
Odynerus parietum var. incisus Verhoeff, 1890. Ent. Nachr. 16: 335. Type destroyed.
Odynerus parietum var. parietoides Verhoeff, 1890. Ent. Nachr. 16: 335. Type destroyed.
Ancistrocerus tardinotus Taylor, 1922. Psyche 29: 49. A manuscript name attributed to
Bequaert and validated under the provisions of Art. 12 and Art. 16 (a) (viii) of the Code.

Biology: Taylor, 1922. Psyche 29: 49-55 (nest, prey, life history). —Miller, 1954. Canad. Ent.
86: 197-198, 4 figs. (nest, prey). The foregoing references are to the north American
population; biological references to the Palaearctic population are to be found in the
Palaearctic eumenid catalog by van der Vecht and Fischer, 1972.

parredes (Saussure). Ariz. (Huachuca Mts.); Mexico, Central America to Panama.
Odynerus (Ancistrocerus) Parredes Saussure, 1857. Rev. Mag. Zool. (2) 9: 273. 9.
Odynerus Parredesi (!) Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 180. 2. Emend.
simulator Cameron. Nev., Calif. Ecology: Nests in old Sceliphron nests.
Ancistrocerus simulator Cameron, 1908. Amer. Ent. Soc., Trans. 34: 222. 9.

Taxonomy: Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 364-372, figs. 4, 9 (chromosome
number).

Biology: Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 365 (nest).
spilogaster Cameron. West. U.S. east to Mont., Wyo., Utah, northern Ariz. in Upper Sonoran
and Transition Zones. Ecology: Nests in twigs and Sambucus stems, in old oak galls,
and also makes complete mud cells on rocks. Parasite: Aritranis notata sierrae Tow.;

Chrysis coerulans F., C. pattoni Aar.
Ancistrocerus spilogaster Cameron, 1905. Invertebrata Pacifica 1: 121. d.
Odynerus edwardsii Cameron, 1908. Amer. Ent. Soc., Trans. 34: 207. 2.
Taxonomy: Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 364-372, figs. 2, 3, 9 (chromosome
number).
Biology: Bequaert, 1943. Ent. Amer. (n. s.) 23: 248 (nest). —Parker and Bohart, 1966.
Pan-Pacific Ent. 42: 93 (nests, parasites).
spinolae (Saussure). Conn. to Fla. west to Mich., Ill., Mo. and Tex. Ecology: Nests in borings in
wood and in abandoned mud-dauber nests. Parasite: Anthrax aterrimus (Big.);
Miltogrammini sp.; Kennethiella trisetosa (Coor.). Prey: Lepidoptera larvae.
Odynerus (Ancistrocerus) spinolae Saussure, 1855. Etudes sur la famille des Vespides, v.
SypHZlon ee
Biology: Rau, 1946. Brooklyn Ent. Soc., Bul. 41: 10-11 (nest). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 99-100 (nest, prey, life history, parasites).
tuberculocephalus sutterianus (Saussure). Calif., Nev., Utah, Oreg., B. C. in Upper Sonoran
and Transition Zones. Ecology: Nests in old mud-dauber nests. Parasite: Chrysis

coerulans F., C. inflata Aar.
Odynerus (Ancistrocerus) Sutterianus Saussure, 1875. Smithsn. Inst., Mise. Collect. 254:

Melos Gin Pe

Superfamily VESPOIDEA 1505

Ancistrocerus sutteranus (!) Cameron, 1905. Invertebrata Pacifica 1: 123.

Taxonomy: Goodpasture, 1974. Kans. Ent. Soc., Jour. 47: 364-372, figs. 5, 6, 9 (chromosome
number).

Biology: Bequaert, 1943. Ent. Amer. (n. s.) 23: 285 (nest).
tuberculocephalus tuberculocephalus (Saussure). S. Dak., Wyo., Utah, Colo., Ariz., N. Mex.,

western Tex.; Mexico in temperate regions south to Federal District. Ecology: Nests in
borings in wood and Sambucus, and in abandoned mud-dauber nests. Parasite:
Bombyliidae sp.; Chrysis inflata Aar. Prey: Gelechiidae sp. Predator: Cleridae sp., larva.

Odynerus (Ancistrocerus) tuberculocephalus Saussure, 1852. Etudes sur la famille des
Vespides, v. 1, p. 122. 6, @.

Odynerus tuberculiceps Saussure, 1853. Etudes sur la famille des Vespides, v. 1, (errata).
Emend.

Biology: Rau, 1940. Ent. Soc. Amer., Ann. 33: 593 (nest). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 102-104 (nest, prey, life history, associates). —Parker and Bohart,
1968. Pan-Pacifie Ent. 44: 2 (nest).

unifasciatus seminole Bequaert. Fla.
Ancistrocerus unifasciatus seminole Bequaert, 1943. Ent. Amer. (n. s.) 23: 282. 2.
unifasciatus unifasciatus (Saussure). Southern Canada and eastern U.S. west to Ont., Mich.,
Iowa, Kans. and Tex. Ecology: Nests in abandoned mud-dauber nests.
Odynerus (Ancistrocerus) unifasciatus Saussure, 1852. Etudes sur la famille des Vespides,
ealkye, VALS

Biology: Rau and Rau, 1913. Ent. News 24: 396-397 (nest).
waldenii excavatus Bequaert. B. C., Wash., Oreg., Calif., Mont., Wyo., Colo., Ariz. Ecology:
Builds mud nests on rocks.
Ancistrocerus waldenii excavatus Bequaert, 1943. Ent. Amer. (n. s.) 23: 244. d, 2.

Biology: Bequaert, 1943. Ent. Amer. (n. s.) 23: 246 (nest).
waldenii flavidulus Bequaert. Oreg., Calif. Ecology: Builds mud nests on rocks. Prey: Larvae

of Cnephasia longana (Haw.), Archips argyrospilus Wlkr.
Ancistrocerus waldenii flavidulus Bequaert, 1943. Ent. Amer. (n. s.) 23: 246. d, °.

Biology: Bequaert, 1943. Ent. Amer. (n. s.) 23: 247 (nest). —Richards, 1962. Pan-Pacific Ent.
38: 145-146 (prey).
waldenii waldenii (Viereck). Alaska, Canada and U. S. south to Idaho, Wyo., N. Mex. (high
mountains), Colo., S. Dak., Ill., Mich., Va. Ecology: Builds free mud nests on walls or

rocks, or in cavities in mortar.
Odynerus waldonii (!) Viereck, 1906. Ent. News 17: 304. °.
Odynerus waldenii Viereck, 1906. Ent. News 17: 350. Emend.

Biology: Bequaert, 1943. Ent. Amer. (n. s.) 23: 243 (nest). —Morris, 1959. Canad. Ent. 91: 500
(nest). ;

Genus SYMMORPHUS Wesmael
Genus SYMMORPHUS Subgenus SYMMORPHUS Wesmael

Symmorphus Wesmael, 1836. Acad. Sci. Belg., Bul. 3: 45.
Type-species: Vespa elegans Wesmael. Desig. by Richards, 1935.
Odynerus subg. Protodynerus Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p.
186. N. name for Symmorphus.

Only the typical subgenus occurs in North America. So far as is known, the American species
nest in pre-existing cavities in twigs, stems, logs and structural lumber; a few European species
are also known to nest in abandoned burrows of other wasps or bees in banks or mud walls. Par-
titions between cells are made of mud. Probably most of the North American species prey upon
externally feeding chrysomelid larvae, but canadensis (Sauss.) is unique in that it preys upon
leaf-mining coleopterous and lepidopterous larvae.

Revision: Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 151-157.

1506 Hymenoptera in America North of Mexico

Taxonomy: van der Vecht, 1966. Ent. Ber. 26: 163 (discussion of type-species designation).
albomarginatus (Saussure). Que., Conn., N. Y., Md., Mich., Wis., Wyo., Alta., B. C., Alaska.
Ecology: Nests in borings in wood and sumac stems. Parasite: Chrysis coerulans F.;
Melittobia chalybii Ashm.; Amobia distorta (Allen). Prey: Chrysomela spp., larvae.
Odynerus (Protodynerus) albomarginatus Saussure, 1855. Etudes sur la famille des
Vespides, v. 3, p. 195. @.

Biology: Medler, 1966. Ent. Soc. Wash., Proc. 68: 144-145 (nest, prey, life history).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 120-122 (nest, prey, life history,
associates).

canadensis (Saussure). Transcont. in Canada and U. S. (not known from Fla., Tex., N. Mex.,

Ariz.). Ecology: Nests in borings in wood, twigs, logs, structural lumber. Parasite:
Chrysis cembricola Krom.; Melittobia chalybii Ashm.; Monodontomerus sp.; Pyemotes
ventricosus (Newp.). Prey: Chalepus dorsalis Thunb.; Prionomerus calceatus (Say),
Apion sp.; Antispila nyssaefoliella Clem.; Lithocolletis ostensackenella (Fitch), L. sp.,
Parectopa robiniella Clem., Gracilariidae sp.; Aeaea ostryaeella (Chamb.). Predator:
Trogoderma ornatum Say; Philanthus pulcher D. T.

Odynerus (Protodynerus) canadensis Saussure, 1855. Etudes sur la famille des Vespides,
Voos pal 9Gmo”

Odynerus (Symmorphus) debilis Saussure, 1870. Rev. Mag. Zool. (2) 22: 55. °.

Symmorphus cogitans Cameron, 1906. Amer. Ent. Soc., Trans. 32: 325. 2.

Biology: Reinhard, 1929. Witchery of Wasps, pp. 72-83 (nest, prey, life history, parasite).
—Krombein, 1952. Amer. Ent. Soc., Trans. 78: 91 (nest, prey). —Krombein, 1954. Brooklyn
Ent. Soc., Bul. 49: 3 (nest, prey). —Krombein, 1956. Ent. Soc. Wash., Proc. 58: 155
(provisioning flights, prey). —Weaver and Dorsey, 1965. Ent. Soc. Amer., Ann. 58: 933-934
(prey). —Krombein, 1967. Trap-nesting wasps and bees, pp. 115-120, pl. 4, figs. 14, 15, pl. 5,
figs. 16-18, pl. 6, fig. 19 (nest, prey, life history, associates).

cristatus cristatus (Saussure). Canada and north. U.S. south to Md., Ill. and mountains of

Colo., Utah, Ariz., Calif. Ecology: Nests in borings in wood and sumac. Parasite: Chrysis
coerulans F., C. nitidula F.; Anthrax irroratus Say; Amobia distorta (Allen). Prey:
Chrysomela crotchi Brown, C. scripta (F.) complex, C. spp., Chrysomelinae sp.,
Gonioctenus americanus Brown.

Odynerus (Protodynerus) cristatus Saussure, 1855. Etudes sur la famille des Vespides, v.
3, p. 196. 2.

Odynerus (Protodynerus) Philadelphiae Saussure, 1857. Rev. Mag. Zool. (2) 9: 272. 9°.

Symmorphus hornii Cameron, 1909. Pomona Col. Jour. Ent. 1: 123. 9.

Biology: Fye, 1965. Canad. Ent. 97: 734-735 (nest, life history, prey, parasite; note data
corrections for Fye in Krombein, 1967). —Krombein, 1967. Trap-nesting wasps and bees,
pp. 122-126, pl. 6, figs. 20-22 (nest, prey, life history, parasites). —Evans, 1973. Great Basin
Nat. 33: 151-152 (nest, prey, parasites).

cristatus nevadaensis (Cameron). Wash., Oreg., Nev., Calif. Ecology: Nests in Sambucus
stems. Parasite: Chrysis derivata Buyss.
Nortonia nevadaensis Cameron, 1905. Invertebrata Pacifica 1: 124. o.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest, parasite).

meridionalis Viereck. B. C., Oreg., Calif., Utah, Colo., Nebr., Wyo., N. Mex. Ecology: Nests in
Sambucus stems. Predator: Philanthus zebratus nitens (Bks.).
Symmorphus meridionalis Viereck, 1903. Amer. Ent. Soc., Trans. 29: 69. 3.
Symmorphus trisulcatus Cameron, 1906. Amer. Ent. Soc., Trans. 32: 327. 6.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest).
projectus Bohart. Calif. (sea level to 10,000 feet), Oreg., Wash., Idaho, Wyo., Mont., Ariz.
Ecology: Nests in Sambucus stems.
Symmorphus projectus Bohart, 1950. Biol. Soc. Wash. Proc. 63: 81. d, 9.
Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest).

walshianus (Saussure). U. S. west to Tex., Iowa.
Odynerus (Symmorphus) Walshianus Saussure, 1870. Rev. Mag. Zool. (2) 22: 55. 2.

Superfamily VESPOIDEA 1507

Genus EUMENES Latreille

Genus EUMENES Subgenus EUMENES Latreille

Eumenes Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 360.
Type-species: Vespa coarctata Linnaeus. Desig. by Latreille, 1810.
Alpha Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 128.
Type-species: Vespa coarctata Linnaeus. Desig. by Bequaert, 1926.
Eumenis Kriechbaumer, 1879. Ent. Nachr. 5: 57. Emend.
Eumenidion Schulthess, 1913. Soe. Ent. 28: 2.
Type-species: Vespa coarctata Linnaeus. Orig. desig.

Only the typical subgenus occurs in America north of Mexico.
So far as is known, all species make juglike mud nests, which are usually attached to twigs and
are provisioned with caterpillars.

Revision: Isely, 1917. Ent. Soc. Amer., Ann. 10: 345-363 (N. Amer. spp.). —Bequaert, 1938.
Brooklyn Ent. Soc., Bul. 33: 59-70 (spp. northeastern U.S. and eastern Canada).
—Bequaert, 1944. Nat. Canad. 71: 75-88 (Canad. spp.).

aureus Isely. Okla., western Tex. to southern Calif.; Mexico (Baja California).
Eumenes belfragei aureus Isely, 1917. Ent. Soc. Amer., Ann. 10: 352. 2.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 152-153, figs. 1, 2 (sleeping habits).
bollii bollii Cresson. Western half of U.S. north to Nebr., Colo., Utah, Nev., Calif., Alta., Mo.,
Iowa, Minn.; Mexico (Baja California).
Eumenes bollii Cresson, 1872. Amer. Ent. Soc., Trans. 4: 232. 6, @.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 299-301 (nest).
bollii oregonensis Bequaert. Wash., Oreg., Idaho, Nev.
Eumenes bollii var. oregonensis Bequaert, 1938. Brooklyn Ent. Soc., Bul. 33: 66. 2.
crucifera bolliformis Viereck. Ariz., N. Mex.; Mexico.
Eumenes (Pachymenes) bolliformis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 387. d, °.
Eumenes robustus Isely, 1917. Ent. Soc. Amer., Ann. 10: 360. 2.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 153 (sleeping habits).
crucifera crucifera Provancher. Western U. S. east to Colo., S. Dak. and north to Wyo., B. C.,
Alta.
Eumenes crucifera Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 421. °.
Eumenes (Alpha) marginilineatus Viereck, 1907. Amer. Ent. Soc., Trans. 33: 381. d.
Eumenes emarginilineatus (!) Viereck, 1907. Amer. Ent. Soc., Trans. 33: 388.
Eumenes sternalis Isely, 1917. Ent. Soc. Amer., Ann. 10: 353. d.
Eumenes xanthogaster Isely, 1917. Ent. Soc. Amer., Ann. 10: 353. d.
crucifera flavitinctus Bohart. Calif.
Eumenes crucifera flavitinctus Bohart, 1950. Biol. Soc. Wash., Proc. 63: 79. d, 2.
crucifera nearcticus Bequaert. Transcontinental north to 63 degrees N. Latitude, south to
Oreg., Utah, Colo., Ky., N. J.
Eumenes minuta Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 39. 3. Preoce.
Eumenes (Alpha) Globulosus Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p.
139. 2. Preoce.
Eumenes nearcticus Bequaert, 1944. Nat. Canad. 71: 82. d, 2. N. name for globulosus.
crucifera stricklandi Bequaert. Alta. (Cypress Hills).
Eumenes coloradensis stricklandi Bequaert, 1944. Nat. Canad. 71: 84. °.
fraternus Say. Ont., U. S. west to Tex., Okla., Kans., Nebr., Minn. Parasite: Chrysis conica Br.;
Timulla ferrugata (F.).
Eumenes fraterna Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 344.
Eumenes Fervens Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 40. d, °.

Biology: Isely, 1914. Kans. Univ. Sci. Bul. (2) 8: 301 (nest).

iturbide iturbide Saussure. Colo., N. Mex., Ariz.; Mexico. -
Eumenes (Alpha) Iturbide Saussure, 1857. Rev. Mag. Zool. (2) 9: 271.
Eumenes (Alpha) globulosiformis Viereck, 1908. Amer. Ent. Soc., Trans. 33: 386. d, 9°.

1508 Hymenoptera in America North of Mexico

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 153 (sleeping habits).
iturbide pedalis Fox. B. C., Wash., Idaho, Mont., Wyo., Colo., Utah, Calif.; Mexico (Baja
California).
Eumenes pedalis Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 109. d, 2.
Eumenes crassicornis Isely, 1917. Ent. Soc. Amer., Ann. 10: 362. d.
Eumenes pachygaster Isely, 1917. Ent. Soc. Amer., Ann. 10: 362. 2.
sculleni Bohart. Nev., Utah.
Eumenes sculleni Bohart, 1950. Biol. Soc. Wash., Proc. 63: 78. 3, 2.
smithii americanus Saussure. Colo., Nebr., Kans., Mo., Ark., Okla., Tex.; Mexico (Michoacan,
Guadalajara). Prey: Geometridae larvae.
Eumenes americanus Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 39. @.
Eumenes belfragei Cresson, 1872. Amer. Ent. Soc., Trans. 4: 232. d, 2.
Eumenes brunneus Isely, 1917. Ent. Soc. Amer., Ann. 10: 348. ¢. Preocc.

Biology: Hartman, 1913. Jour. Anim. Behavior 3: 353-360, 7 figs. (nest building, prey, life
history).
smithii smithii Saussure. N. C., Ga., Fla., Ala., Miss.
Eumenes Smithii Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 43. 6, 9.

Biology: Isely, 1917. Ent. Soc. Amer., Ann. 10: 350 (nest).
verticalis coloradensis Cresson. Western U.S. and Canada (except Oreg., Calif.) east to Sask.,
S. Dak., Colo., N. Mex.; Mexico.
Eumenes coloradensis Cresson, 1875. Rpt. Geog. Geol. Expl. and Survey west of 100th
Meridian, v. 5, p. 717. d, 2.
Eumenes (Alpha) cruciferorum Viereck, 1908. Amer. Ent. Soc., Trans. 33: 388. 6, ?.
Eumenes (Alpha) enigmatus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 389. ¢.
Eumenes stenogaster Isely, 1917. Ent. Soc. Amer., Ann. 10: 353. 6, 9.
Eumenes enigmaticus (!) Bequaert, 1938. Brooklyn Ent. Soc., Bul. 33: 69.
verticalis neoboreus Bequaert. Que., Ont., Alta., B. C.
Eumenes verticalis neoboreus Bequaert, 1944. Nat. Canad. 71: 87. 6, 9.
verticalis tricinctus Isely. Alta., Idaho, Utah, Nev., Oreg., Calif., Ariz.; Mexico (Baja
California).
Eumenes tricinctus Isely, 1917. Ent. Soc. Amer., Ann. 10: 361. °.
verticalis verticalis Say. Ont., Man., north. and east. U. S. south to N. C., Mo. and west to N.
Dak., S. Dak.
Eumenes verticalis Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 346.

SUBFAMILY ZETHINAE
TRIBE ZETHINI

There are only four genera in this tribe. Zethus is primarily Neotropical with a few species
entering the Nearctic Region. The other genera are Australian.

Genus ZETHUS Fabricius

The genus is primarily Neotropical in distribution, but it occurs also in the Nearctic, Oriental
and Ethiopian Regions. Very little is known of the biology of the North American species.
Overall, the species range from the more primitive solitary taxa which nest in abandoned
borings of other insects to subsocial species in which several females build a common nest of
vegetable matter and resin, and feed their larvae progressively with lepidopterous larvae.

Revision: Bohart and Stange, 1965. Univ. Calif., Pubs. Ent. 40: 1-208, 354 figs. (New World

Spp.).
Taxonomy: Porter, 1975. Fla. Ent. 58: 303-306, 5 figs. (key to U.S. spp.).

Genus ZETHUS Subgenus ZETHUS Fabricius

Zethus Fabricius, 1804. Systema Piezatorum, pp. xii, 282.
Type-species: Vespa coeruleopennis Fabricius. Desig. by Ashmead, 1902.

Superfamily VESPOIDEA 1509

Didymogastra Perty, 1833. Delect. anim. artic. Brasil, p. 145.
Type-species: Didymogaster fusca Perty. Desig. by Ashmead, 1902.
Lethus (!) Say, 1836. Boston Jour. Nat. Hist. 1: 209. Misspelling.
Heros Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 115.
Type-species: Zethus gigas Spinola. Monotypic.
Wettsteinia Dalla Torre, 1904. In Wytsman, Gen. Ins., fase. 19, p. 13.
Type-species: Labus sichelianus Saussure. Desig. by Bohart and Stange, 1965.
Euzethus Dalla Torre, 1904. In Wytsman, Gen. Ins., fase. 19, p. 14. N. name.
Laboides Zavattari, 1912. Arch. f. Naturgesch. 78, Abt. A, H. 4: 65.
Type-species: Labus sichelianus Saussure. Desig. by Bohart and Stange, 1965.
miscogaster Saussure. Tex. (Hidalgo Co.), Ariz. (Baboquivari Mts.); Mexico to Argentina.
Zethus Miscogaster Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 18. 9.
Zethus Zendalus Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 53. @.
Zethus emarginatus Fox, 1899. Acad. Nat. Sci. Phila., Proe., p. 429. °.
Zethus claripennis Cameron, 1907. Entomologist 40: 63. d.
Zethus punctinodis Cameron, 1907. Entomologist 40: 79. °.
Didymogastra brethesi Zavattari, 1911. Zool. Jahrb. 31: 42. 2.
Zethus pavidus Zavattari, 1912. Arch. f. Naturgesch. 78, Abt. A, 4: 72. °.
Zethus binghami Zavattari, 1913. Arch. f. Naturgesch. 79, Abt. A, 1: 114. d.
montezuma Saussure. Tex. (Hidalgo Co.) south to Colombia.
Zethus montezuma Saussure, 1857. Rev. Zool. (2) 9: 270. °.
spinipes spinipes Say. Mass. to N. C. west to IIl., Kans., Ark.
Lethus (!) spinipes Say, 1837. Boston Jour. Nat. Hist. 1: 387. °.
Eumenes substricta Haldeman, 1844. Acad. Nat. Sci. Phila., Proc. 2: 54.
Eumenes pensilvanica Haldeman, 1853. Acad. Nat. Sci. Phila., Proc. 6: 365.
Zethus pensylvanica Saussure, 1875. Smithsn. Inst., Mise. Collect. 254: 29. Emend.
Zethus Pennsylvanica Dalla Torre, 1894. Cat. Hym., v. 9, p. 14. Emend.
spinipes variegatus Saussure. Md. to Fla., Ala., La., Tex., Okla. Prey: Rhyacionia frustrana
(Comst.).
Zethus variegatus Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 13. °.
Zethus bicolor Saussure, 1852. Etudes sur la famille des Vespides, v. 1, p. 17. 6.

Biology: Lashomb and Steinhauer, 1975. Ent. Soc. Wash., Proc. 77: 164 (prey capture).
Genus ZETHUS Subgenus ZETHUSCULUS Saussure

Zethusculus Saussure, 1855. Etudes sur la famille des Vespides, v. 3, p. 118.
Type-species: Zethus jurinei Saussure. Desig. by Ashmead, 1902.
slossonae Fox. Southern Fla. Ecology: Nests in twigs.
Zethus slossonae Fox, 1892. Ent. News 2: 29. d, °.

Biology: Bohart and Stange, 1965. Univ. Calif., Pubs. Ent. 40: 16 (nest).
Genus ZETHUS Subgenus ZETHOIDES Fox

Zethoides Fox, 1899. Acad. Nat. Sci. Phila., Proc., p. 436.
Type-species: Zethoides smithii Fox. Monotypic.
Baeoprymna Cameron, 1912. Timehri 2: 225.
Type-species: Baeoprymna rufoornata Cameron. Monotypic.
Protozethus Bertoni, 1925. Soc. Cient. Paraguay, Rev. 2: 75.
Type-species: Zethus olmecus Saussure. Orig. desig.
aztecus Saussure. Tex. (Hidalgo Co.) south to Guatemala and El Salvador.
Zethus aztecus Saussure, 1857. Rev. Zool. (2) 9: 270. d.
Zethus campestris Zavattari, 1913. Arch. f. Naturgesch. 79, Abt. A, h 1: 108. 2.
guerreroi arizonensis Bohart. Ariz., N. Mex.; Mexico (Sonora, Sinaloa). Typical guerrero
Zavattari occurs in Mexico and E] Salvador.
Zethus (Zethusculus) arizonensis Bohart, 1950. Biol. Soc. Wash., Proc. 63: 77. 3, &.

UNPLACED TAXA OF EUMENIDAE

Odynerus bimaculatus Provancher, 1895. Nat. Canad. 22: 157. °. Type in Laval Univ., Que.

1510 Hymenoptera in America North of Mexico

Odynerus bisstrigatus Spinola, 1808. Insectorum Liguriae, v. 1, p. 180. Amer. sept. Described
originally as bis-strigatus; location of type unknown.

Odynerus (2) bradleyi Cameron, 1909. Pomona Col. Jour. Ent. 1: 81 2. Colo. (Durango). Type
depository unknown.

Odynerus crotchii Cameron, 1908. Amer. Ent. Soe., Trans. 34: 197. 6, 2. Tex. (Lee Co.).
Lectotype in British Museum (Nat. Hist.); possibly a Leptochilus.

Odynerus (Odynerus) nortonianus Saussure, 1870. Rev. Mag. Zool. (2) 22: 105. 2. Conn.

Odynerus pulchrinervis Cameron, 1906. Amer. Ent. Soc., Trans. 32: 328. 2. N. Mex. Type
depository unknown, the type probably lost.

Odynerus robustus Provancher, 1895. Nat. Canad. 22: 157. 2. Calif. (Los Angeles). Type in
Laval Univ., Que.

Odynerus scudderi Cameron, 1908. Amer. Ent. Soc., Trans. 34: 196. 2. Southwest. U. S. Type
depository unknown, the type probably lost.

Odynerus tricolor Provancher, 1895. Nat. Canad. 22: 158. 3. Calif. (Los Angeles). Type in
Laval Univ., Que.

NoMINA NUDA IN EUMENIDAE

Ancistrocerus behrensi Tucker, 1909. Kans. Acad. Sci., Trans. 22: 286.

Leionotus scrophulariae Robertson, 1928. Flowers and Insects, p. 11.

Leionotus ziziae Robertson, 1928. Flowers and Insects, p. 12.

Odynerus daedalus Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p. 589.

Family VESPIDAE

The wasps belonging to this family are all social species, the colonies consisting of one or more
fertile queens, workers which are frequently numerous and usually infertile, and, at times, males

and new queens.

Taxonomy: Lewis, 1897. Amer. Ent. Soc., Trans. 24: 169-192, 1 pl. (N. Amer. spp.) —Bohart
and Bechtel, 1957. Calif. Ins. Survey, Bul. 4: 73-102, 8 pls., 34 figs., 7 maps (Calif. spp.).

Biology: Rau, 1929. Canad. Ent. 61: 219-221 (cocoon spinning). —Jeanne, 1975. Quart. Rev.
Biol. 50: 267-287, 9 figs. (adaptiveness of nest architecture).

Morphology: van der Vecht, 1968. K. Nederland. Akad. van Wetensch., Proc., Ser. C, 71:
411-422, 5 figs. (terminal gastral sternite, female, worker).

SUBFAMILY POLISTINAE

These are commonly called paper wasps in North America. The nests consist of a single comb
which is not enclosed in a paper envelope. In most of the Temperate Zone species the nests are
annual, each being founded by one or more overwintering females. One of the females becomes
dominant and is the only one to lay eggs.

The normal larval food consists of dismembered caterpillars. Some species of Polistes exert
considerable predator pressure on economically important insects such as the tobacco hornworm
and cotton bollworm. In an attempt to utilize the wasps as a biological control agent, shelters are
sometimes placed in cultivated fields to afford nesting sites to founding females.

Taxonomy: Yoshikawa, 1962. Jour. Biol. 13: 19-43, 1 map (list of world spp., distribution).
Biology: Yoshikawa, 1962. Jour. Biol. 13: 45-64 (evolution of sociality).

Genus POLISTES Latreille

Polistes Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 363.
Type-species: Vespa gallica Linnaeus. Desig. by Blanchard, 1840.

Richard’s (1973) reclassification of the subgenera of Polistes, listed below in Taxonomy, was
received too late to assign the North American species to subgenera in the format used el-
sewhere in the catalog for subgenera. Our species are assigned to the following subgenera.

Polistes subg. Fuscopolistes Richards, 1973, pp. 95, 100 (type-species, Vespa fuscata

Superfamily VESPOIDEA 1511

Fabricius, orig. desig.)-- apachus, bellicosus, carolina, dorsalis, flavus, fuscatus, metricus and
perplexus.

Polistes subg. Palisotius Richards, 1973, pp. 95, 100 (type-species, Polistes major Beauvois,
orig. desig.)--major.

Polistes subg. Onerarius Richards, 1973, pp. 94, 101 (type-species, Vespa carnifex Fabricius,
orig. desig.)--carnifex.

Polistes subg. Polistarchus Richards, 1973, pp. 94, 101 (type-species, Vespa canadensis Lin-
naeus, orig. desig.)--annularis, canadensis, exclamans and instabilis.

Polistes subg. Epicnemius Richards, 19738, pp. 96, 102 (type-species, Polistes bicolor Lepeleti-
er, orig. desig.)--pacificus.

The typical subgenus does not occur in North America.

In the extensive series of papers listed below, Rau consistently used two misidentifications:
pallipes for the taxon now recognized as metricus, and variatus for the taxon now recognized as
typical fuscatus; he also used rubiginosus for carolina, a senior synonym.

Revision: Bequaert, 1937. Arch. Inst. Biol. Veg., Rio de Janeiro 3: 171-205 (New World taxa
of Gallicus and Bicolor species groups). —Bequaert, 1940. N. Y. Ent. Soc., Jour. 48: 1-31
(N. Amer. taxa). —Bequaert, 1943. Ent. Venezolana, Bol. 2: 107-124 (New World vars. of
canadensis (L.)). —Snelling, 1955. Amer. Mus. Novitates 1701: 1-9 (subspp. of major
Beauv. and exclamans Vier.).

Taxonomy: Enteman, 1904. Carnegie Inst. Wash., Pub. 19: 1-88, 6 pls., 27 figs. (coloration).
—Rau, 1942. Ent. Soc. Amer., Ann. 35: 335-338 (behavioral characters of carolina, f.
fuscatus, metricus). —Richards, 1973. Rev. Ent. Brasil. 17: 85-1038, 12 figs. (classification
of subgenera). —Snelling, 1974. Ent. Soc. Wash., Proc. 76: 476-479 (status of some N.
Amer. taxa).

Biology: Rau, 1928. Biol. Bul. 54: 503-519 (honey-gathering by annularis, f. fuscatus,
metricus). —Rau, 1928. Psyche 35: 147-150, 4 figs. (modification of nests by carolina,
annularis, metricus). —Rau, 1928. Brooklyn Ent. Soc., Bul. 23: 230-235 (hibernation,
colony founding by annularis, metricus). —Duncan, 1928. Pan-Pacific Ent. 5: 90 (building
material). —Rau, 1928. Psyche 35: 151-152 (reconstruction of destroyed nests by f.
fuscatus, metricus). —Rau, 1929. Ecology 10: 191-200, 4 figs. (nesting habitat of
annularis, carolina, f. fuscatus, metricus). —Rau, 1930. Compar. Psychol., Jour. 10:
267-286 (interspecific animosity and tolerance in workers of f. fuscatus, metricus,
annularis). —Rau, 1930. Ent. Soc. Amer., Ann. 23: 461-466 (hibernation behavior of
carolina, f. fuscatus, metricus, annularis). —Rau, 1931. Ecology 12: 690-693 (use of water
by annularis, metricus, f. fuscatus, carolina). —Rau, 1931. Ent. Soc. Amer., Ann. 24:
515-518 (hibernation behavior of annularis, metricus). —Rau, 1939. Compar. Psychol.,
Jour. 27: 259-269 (inter- and intraspecific animosity and tolerance of female annularis, f.
fuscatus, metricus). —Rau, 1939. Ecology 20: 439-442 (population and caste studies of
metricus, f. fuscatus, annularis). —Rau, 1941. Ent. Soc. Amer., Ann. 34: 355-366
(parasites of carolina, annularis, metricus, f. fuscatus). —Rau, 1941. Ent. Soc. Amer.,
Ann. 34: 580-584 (swarming of annularis, f. fuscatus, metricus). —Rau, 1946. Ent. Soe.
Amer., Ann. 39: 11-27 (nest and cell size of f. fuscatus, metricus). —Rabb and Lawson,
1957. Econ. Ent., Jour. 50: 778-784, 2 figs. (predation on tobacco hornworm in N. C. by e.
exclamans, f. fuscatus). —Rabb, 1960. Ent. Soc. Amer., Ann. 53: 111-121 (nest founding,
life history, adult and larval food, prey, foraging, parasites in N. C. in f. fuscatus,
annularis, e. exclamans, metricus, d. dorsalis, carolina). —West, 1967. Science 157:
1584-1585 (female dominancy hierarchies in canadensis, fuscatus). —Nelson, 1968. Ent.
Soc. Amer., Ann. 61: 1528-1539, 3 figs. (parasites, symbionts). —Eberhard, 1969. Mich.
Univ., Mus. Zool., Mise. Pub. 140: 1-101, 23 figs. (social biology of f. fuscatus, canadensis
erythrocephalus Latr., c. infuscatus Lep.). —Kirkton, 1970. Tall Timbers Conf., Proce. 2:
243-245 (population management in Ark.). —Gillaspy, 1971. Ent. Soc. Amer., Ann. 64:
1357-1361, 4 figs. (population management in Tex., Calif.). —Gillaspy, 1973. Amer. Midland
Nat. 90: 1-12, 5 figs. (behavior of metricus, annularis, major, bellicosus, exclamans,
apachus, carolina in artificial nesting sites).

annularis (Linnaeus). Conn. to Fla., west to S. Dak. and central Tex. Ecology: Nests
suspended from twigs and branches of shrubs and trees in exposed situations, and

1512 Hymenoptera in America North of Mexico

occasionally in sheltered sites on buildings. Parasite: Xenos pallidus Brues; Sarcophaga
sp.; Apanteles carpatus (Say); Pachysomoides fulvus (Cr.), P. stupidus (Cr.); Elasmus
polistis Burks. Prey: Halisidota tessellaris (Sm.), Hyphantria cunea (Dru.); Anisota
virginiensis (Dru.), A. senatoria (Sm.), A. stigma (F.), Citheronia regalis (F.);
Geometridae spp.; Limacodidae sp.; Hemerocampa leucostigma (Sm.); Datana spp.,
Heterocampa manteo (Dbldy.), H. spp., Nadata gibbosa (Grt.), Peridea angulosa (A. and
S.), Schizura leptinoides (Grt.), S. spp.; Basilarchia sp.; Psilocorsis sp.; Acronicta
afflicta Grt., A. retardata (Wlkr.), A. spp. Autographa sp., Catocala sp., Heliothis zea
(Boddie), Hypsoropha hormos Hbn., Phosphila turbulente (Hbn.); Tropaea luna (L.);
Ampeloeca myron (Cram.), Cressonia juglandis (A. and S.), Protoparce sexta (Johan.);
prey is usually obtained in wooded areas. Predator: Dicymolomia pegasalis (Wlkr.),
Chalcoela iphitalis Wlkr.

Vespa annularis Linnaeus, 1763. Centuria Ins. Rar., p. 30.

Vespa cincta Drury, 1773. Illus. Nat. Hist., Index to pt. 1 published with pt. 2.

Biology: Fox, 1896. Ent. News 7: 57 (nest). —Brimley, 1908. Ent. News 19: 107 (male
hibernation). —Rau and Rau, 1918. Wasp Studies Afield, pp. 283-290, fig. 52 (hibernation,
colony founding). —Rau, 1930. Canad. Ent. 62: 81-83 (hibernation mortality). —Rau, 1930.
Canad. Ent. 62: 119-120 (behavior on nest). —Rau, 1931. Brooklyn Ent. Soc., Bul. 26:
116-118, fig. 5 (nest). —Rau, 1940. Ent. Soc. Amer., Ann. 33: 617-620 (cooperative nest
founding). —Rau, 1942. Ent. Soc. Amer., Ann. 35: 94-96 (temperature inducing
hibernation). —Balduf, 1961. Ent. News 72: 259-260 (autumnal swarming). —Hermann and
Dirks, 1974. Ga. Ent. Soc., Jour. 9: 1-8, 3 figs. (smearing on nest by sternal glands).
—Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc., Jour. 9: 203-204 (hibernation, spring
mating). —Hermann and Dirks, 1975. Psyche 82: 97-108 (spring nesting behavior).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): 48-49, figs. 16 I-P (male
genitalia). —Hunt and Hermann, 1971. Ga. Ent. Soc., Jour. 5: 210-216, 2 figs. (poison
apparatus).

apachus Saussure. Western Kans. and Tex., to southern Colo., introduced into Calif.; Mexico
(Nuevo Leon, San Luis Potosi, Coahuila, Sonora, Durango). Ecology: Nests suspended
from branches, usually at tops of low trees. Parasite: Pachysomoides fulvus Cr.;
Sarcophaga sp.; Xenos peckii Kby. Predator: Chalcoela iphitalis Wlkr. This is commonly
called the Apache wasp in California where it is a pest in fig trees.
Polistes apachus Saussure, 1857. Soc. Ent. France, Ann. (3) 5: 314.
Polistes texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 246. 6, °.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 522, fig. 15 (nest). —Simmons, Fisher and
Tyler, 1948. Ent. Soc. Amer., Ann. 41: 450-455, 1 pl. (nest, life history). —Snelling, 1952.
Pan-Pacific Ent. 28: 177 (hibernation).

canadensis (Linnaeus). Southern Ariz. to Peru, Bolivia and Paraguay. Ecology: Nests in trees,
culverts and under eaves of buildings.
Vespa canadensis Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 574.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 524, fig. 11 (nest).
carnifex carnifex (Fabricius). South. Tex., Ariz. (Nogales); Mexico to Argentina. Ecology:

Nests under eaves, in culverts and in shrubby trees. Other subspecies occur in South
America.

Vespa carnifex Fabricius, 1775. Syst. Ent., p. 365. 9.

Polistes onerata Lepeletier, 1836. Hist. Nat. Ins., Hym. 1: 524. ¢.

Polistes valida Say, 1837. Boston Jour. Nat. Hist. 1: 389. 2, d.

Polistes transverso-strigata Spinola, 1851. Accad. Sci. Torino, Mem. (2) 13: 78. 2.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 521, figs. 3, 4 (nest). —Corn, 1978 (1972).
Psyche 79: 150-157, 2 figs. (nest, behavior).
carolina (Linnaeus). Eastern U. S., Pa. to Fla., west to Kans. and central Tex. Ecology: Nests
in very sheltered situations, such as in hollow trees or in sidings of houses. Parasite:
Xenos nigrescens Brues, X. rubiginosi (Pierce); Pachysomoides fulvus (Cr.). Predator:
Dicymolomia pegasalis (Wlkr.); Chalcoela iphitalis Wlkr. P. carolina (L.) has been

Superfamily VESPOIDEA 1513

confused with perplexus Cr. in the past. Some of the parasite-predator records and
biological references listed here may actually refer to the latter species.

Vespa carolina Linnaeus, 1767. Syst. Nat., Ed. 12, v. 1 (pt. 2), p. 948. d.

Polistes rubiginosa Lepeletier, 1836. Hist. Nat. Ins., Hym. v. 1, p. 524. 6.

Biology: Girault, 1907. Canad. Ent. 39: 355-356 (as predator). —Rau and Rau, 1918. Wasp
Studies Afield, pp. 280-283 (prey forage, hibernation). —Rau, 1929. Ent. News 40: 7-13
(behavior female, male at end of season). —Rau, 1931. Brooklyn Ent. Soc., Bul. 26: 115-116
(nest). —Rau, 1931. Psyche 38: 129-144, 6 figs. (nesting habits, pleometrosis). —Wilson,
1933. Fla. Entomologist 17: 12 (as predator). —O’Byrne, 1938. Ent. News 49: 288-289 (nest
site).

comanchus comanchus Saussure. Southwestern Tex.; Mexico (Coahuila, Durango).
Polistes comanchus Saussure, 1857. Soc. Ent. France, Ann. (3) 5: 314.
comanchus navajoe Cresson. Western Tex. to Ariz.; Mexico (Sonora, Chihuahua, Durango,
Sinaloa). Ecology: Nests in cavities having very small openings.
Polistes navajoe Cresson, 1868. Amer. Ent. Soc., Trans. 1: 383. °.
dorsalis californicus Bohart. Calif., Ariz.; Mexico (Baja California, Sonora, Sinaloa). Parasite:
Xenos peckii Kby.
Polistes hunteri californicus Bohart, 1949. Pan-Pacifie Ent. 25: 101. d, 9.
dorsalis dorsalis (Fabricius). Va. to Fla., west to Colo., Okla., Tex., N. Mex. south to Honduras.
Ecology: Nests in shrubbery and beneath eaves. Parasite: Xenos peckii Kby.
Vespa dorsalis Fabricius, 1775. Systema Ent., p. 367. °.
Polistes fuscatus hunteri Bequaert, 1940. N. Y. Ent. Soc., Jour. 48: 23. 2, d.

Taxonomy: van der Vecht, 1970. K. Nederland. Akad. van Wetensch., Proc., Ser. C, 71: 23
(identity of Fabrician type).

Biology: Spieth, 1948. N. Y. Ent. Soc., Jour. 56: 155-169 (nest, life history). —Hermann,
Gerling and Dirks, 1974. Ga. Ent. Soc., Jour. 9: 203-204 (hibernation).

exclamans arizonensis Snelling. Ariz., Calif.; Mexico (Sonora).
Polistes exlamans arizonensis Snelling, 1954. Kans. Ent. Soc., Jour. 27: 152. 2, d.

Biology: Hopkins, 1955. Jour. Econ. Ent. 48: 161-163 —Fye, 1972. Envir. Ent. 1: 55-57 (colony
manipulation).
exclamans bahamensis Bequaert and Salt. La. to Fla.; Bahamas.
Polistes bahamensis Bequaert and Salt, 1931. Ent. Soc. Amer., Ann. 34: 794. 2°, 8, d.
Polistes exclamans lowisianus Bequaert, 1940. N. Y. Ent. Soe., Jour. 48: 14. 2.
exclamans exclamans Viereck. N. J. to Fla., west to Nebr., Colo., and Tex.; Mexico (Nuevo
Leon, Coahuila). Ecology: Nests are usually in sheltered sites, frequently under eaves or
roofs. Parasite: Sarcophaga polistensis Hall, S. sp.; Pachysomoides stupidus (Cr.), P.
fulvus (Cr.); Elasmus polistis Burks; Monodontomerus minor (Ratz.). Prey: Arctiidae
sp.; Anisota senatoria (Sm.); Hesperiidae sp.; Heterocampa sp.; Alabama argillacea
(Hbn.), Autographa spp., Feltia subterranea (F.), Heliothis zea (Boddie), Laphygma
frugiperda (Sm.), Plathypena scabra (F.), Prodenia prob. ornithogalli Guen.; Colias
philodice eurytheme Bdv.; Loxostege similalis (Guen.); Protoparce sexta (Johan.);
usually obtained from herbaceous vegetation in open fields. Predator: Dicymolomia
pegasalis (Wlkr.), Chalcoela iphitalis Wlkr.
Polistes exclamans Viereck, 1906. Amer. Ent. Soc., Trans. 32: 201. 2.

Taxonomy: Snelling, 1955. Amer. Mus. Novitates 1701: 5-6 (synonymy). —Eickwort, 1969.
Evolution 23: 391-405 (variation between sexes and castes).

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 525-526, figs. 7-9 (nest). —Hodapp and Bickley,
1959. Ent. Soc. Wash., Proc. 61: 73 (autumnal swarming). —West, 1968. Psyche 75: 118-123
(distribution, nest founding). —Eickwort, 1969. Insectes Sociaux 16: 67-72 (caste separation
female, worker). —Lin, 1972. Ent. Soe. Wash., Proc. 74: 148-155 (male territoriality).
—Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc., Jour. 9: 203-204 (hibernation).
—Hermann and Dirks, 1974. Ga. Ent. Soc., Jour. 9: 1-8 (smearing on nest by sternal
glands). —Hermann, Barron and Dalton, 1975. Ent. News 86: 173-178 (nest founding

behavior in spring).

1514 Hymenoptera in America North of Mexico

Morphology: Crouch and Smith, 1958. Tex. Jour. Sci. 10: 38-59, 20 figs. (sting and associated
glands).
flavus Cresson. Southern Calif., Nev., Utah, Ariz., N. Mex., western Tex.; Mexico (Sonora,
Chihuahua, Coahuila). Parasite: Xenos peckii Kby.
Polistes flavus Cresson, 1868. Amer. Ent. Soc., Trans. 1: 383. °.
fuscatus aurifer Saussure. B. C. to Mont., south to Calif. and Nev.; Mexico (Baja California).
Ecology: Nests in sheltered situations.Adventive in Hawaii, Johnston, Niihau and
Society Islands. Parasite: Gordius sp.; Xenos peckii Kby.
Polistes aurifer Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 78. °.
Polistes anaheimensis Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 423. d,
OF
Polistes fuscatus var. utahensis Hayward, 1933. Utah Acad. Sci., Proc. 10: 142. d, 9.
Polistes fuscatus connectens Bequaert, 1940. N. Y. Ent. Soc., Jour. 48: 21. 2, ¢.
Polistes fuscatus montanus Bequaert, 1940. N. Y. Ent. Soc., Jour. 48: 26. 2, 3.

Taxonomy: Snelling, 1954. Kans. Ent. Soc., Jour. 27: 154 (synonymy).

Biology: Duncan, 1928. Pan-Pacifie Ent. 5: 90 (nest materials). —Bohart, 1942. Pan-Pacific
Ent. 18: 30 (feeding, hibernation). —Snelling, 1952. Pan-Pacific Ent. 28: 177 (hibernation).
—Snelling, 1954. Kans. Ent. Soc., Jour. 27: 154 (nest site).

fuscatus bellicosus Cresson. S. C. to Fla., west to Tex.
Polistes bellicosus: Cresson, 1872. Amer. Ent. Soc., Trans. 4: 247. 6, @.

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 294-295 (drinking habits).
fuscatus centralis Hayward. Western Tex., Colo., N. Mex., Ariz., Utah, Calif.; Mexico (Baja
California, Sonora, Chihuahua, Coahuila). Ecology: Nests in sheltered situations.
Polistes fuscatus var. centralis Hayward, 1933. Utah Acad. Sci., Proc. 10: 143. d, @.

fuscatus fuscatus (Fabricius). U. S. east of Rocky Mts. except in extreme southeast and Mich.

where it is replaced by f. bellicosus Cr. and f. lawrentianus Begq. respectively. Ecology:
Nests in sheltered situations, frequently beneath eaves and roofs, or in cavities near
ground. Parasite: Xenos peckii Kby.; Pachysomoides fulvus (Cr.); Elasmus polistis
Burks; Dasymutilla castor (Bl.). Prey: Estgimene acrea (Dru.), Hyphantria cunea
(Dru.); Geometridae spp.; Hesperiidae sp.; Everes comyntes (Godt.); Dasylophia anguina
(A. and S.), Heterocampa manteo (Dbldy.); Euptoieta sp., pupa; Acronicta sp.,
Autographa spp., Feltia subterranea (F.), Heliothis zea (Boddie), H. virescens (F.),
Hypsoropha hormos Hbn., Laphygma frugiperda (Sm.), Peridroma margaritosa (Haw.),
Plathypena scabra (F.), Prodenia prob. ornithogalli Guen., Schinia sp.; Colias philodice
eurytheme Bdv., Phoebis sp.; Loxostege sp.; Protoparce sexta (Johan.); Orthoptera spp.;
usually obtained from herbaceous vegetation in open fields. Predator: Dicymolomia
pegasalis (Wlkr.), Chalcoela iphitalis Wlkr.

Vespa fuscata Fabricius, 1793. Ent. System., v. 2, p. 260.

Vespa nestor Fabricius, 1798. Sup. Ent. System., p. 262.

Polistes pallipes Lepeletier, 1836. Hist. Nat. Ins. Hym., v. 1, p. 5380. °.

Polistes exilis Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 85.

Polistes variatus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 247. d, °.

Polistes pallidipes Dalla Torre, 1894. Cat. Hym., v. 9, p. 133. Emend.

Biology: Girault, 1911. Wis. Nat. Hist. Soc., Bul. 9: 49-63 (observations on female). —Davis,
1919. Brooklyn Ent. Soe., Bul. 14: 122-123 (nest). —Bequaert, 1923. Brooklyn Ent. Soc.,
Bul. 18: 73-80 (vestigial pleometrosis). —Davis, 1924. Staten Island Inst. Arts and Science
2: 137-138 (nest). —Bugbee, 1936. Ent. Soc. Amer., Ann. 29: 614-616 (nest). —Gaul, 1940. N.
Y. Ent. Soc., Jour. 48: 391-393 (artificial brood rearing). —Beall, 1942. Canad. Field-Nat.
56: 64-67 (autumnal swarming). —Eberhard, 1969. Mich. Univ., Mus. Zool., Mise. Pub. 140:
1-101 (social biology). —Gibo, 1972. N. Y. Ent. Soc., Jour. 80: 105-108 (hibernation sites,
temperature tolerance). —Hermann and Dirks, 1974. Ga. Ent. Soe., Jour. 9: 1-8 (smearing
on nest by sternal glands). —Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc., Jour. 9:
203-204 (hibernation). —Gibo, 1974. Canad. Ent. 106: 101-106 (colony founding in lab.). Rau
published many notes on this taxon as variatus; they are cited under the generic heading.

Superfamily VESPOIDEA 1515

Morphology: Marshall and Dernehl, 1905. Ztschr. f. Wiss. Biol. 80: 122-154, 2 pls.
(embryology). —Marshall, 1907. Ztschr. f. Wiss. Biol. 86: 173-213, 3 pls. (embryology).
fuscatus laurentianus Bequaert. Que., Ont., Mich.
Polistes fuscatus laurentianus Bequaert, 1942. Canad. Ent. 74: 159. d, °.
instabilis Saussure. South. 'Tex.; Mexico to Costa Rica. Ecology: Nests in trees, in culverts,
under bridges and overhanging rocks. Predator: Chalcoela iphitalis Wlkr.
Polistes instabilis Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 91, pl. 10, fig.
Zplemllefigrele Or

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 524-525, figs. 5, 6, 13 (nest).

kaibabensis Hayward. Ariz. (Grand Canyon), southern Utah.
Polistes canadensis var. kaibabensis Hayward, 1932. Utah Acad. Sci., Proc. 9: 89. 6, @.
major castaneicolor Bequaert. Southern N. Mex. and Ariz.; Mexico (Baja California, Sonora,
Chihuahua, Jalisco), Colombia.
Polistes major var. castaneicolor Bequaert, 1936. Ent. News 47: 12. 2, 6.
major major Beauvois. Southern Fla., Tex., N. Mex. and Ariz.; Mexico south to Brazil; Antilles.
Ecology: Nests in shrubbery, in culverts and beneath eaves. Parasite: Elasmus polistis
Burks. Predator: Chalcoela iphitalis Wlkr.
Polistes Major Beauvois, 1818. Ins. Afr., Amer., p. 206.
Polistes major bakeri Bequaert, 1940. N. Y. Ent. Soc., Jour. 48: 15. 3, °.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 522, fig. 12 (nest).
major palmarum Bequaert. Southern Calif.; Mexico (Baja California).
Polistes major var. palmarum Bequaert, 1936. Ent. News 47: 11. 2.
metricus Say. Eastern U.S. from N. Y. to Fla., west to Nebr., Okla. and Tex. Ecology: Nests
in shrubbery. Parasite: Xenos peckii Kby.; Sarcophaga polistensis Hall, S. sp.;
Sphecophaga vesparum burra (Cr.); Pachysomoides fulvus (Cr.). Predator:
Dicymolomia pegasalis (Wlkr.), Chalcoela iphitalis Wlkr.
Polistes metrica Say, 1831. Indiana: School Press, New Harmony, p. 15.

Taxonomy: Rau, 1929. Psyche 36: 34-36 (color variation).

Biology: Turner, 1912. Psyche 19: 184-190 (orphan nest, feeding, homing, hunting). —Pellett,
1916. Iowa Acad. Sci., Proce. 23: 275-284 (life history, behavior in lab.). —Rau and Rau,
1918. Wasp Studies Afield, pp. 244-280, 291-294 (homing experiments with female, worker;
nest sites). —Rau, 1928. Psyche 35: 153-156 (trophallaxis). —Rau, 1929. Canad. Ent. 61:
25-30 (feeding experiments). —Rau, 1929. Ent. News 40: 226-232, 256-259 (orphan nests).
—Rau, 1930. Canad. Ent. 62: 143-147, 2 figs. (life history, nest destruction). —Rau, 1931.
Brooklyn Ent. Soc., Bul. 26: 111-113 (nest, nest site). —Rau, 1935. Ent. News 46: 25-27
(female duties). —Madden and Chamberlain, 1938. Jour. Econ. Ent. 31: 705 (as predator).
—Hermann, Gerling and Dirks, 1974. Ga. Ent. Soc., Jour. 9: 203-204 (hibernation). In these
papers Rau cited metricus as pallipes.

pacificus pacificus Fabricius. South. Tex. to South America. Other subspp. occur in Central

and South America.
Polistes pacifica Fabricius, 1805. Systema Piezatorum, p. 274. °.

Taxonomy: Bequaert, 1937. Inst. Biol. Veget. Arch. 3: 196.
perplexus Cresson. Md. to Ga., west to southern IIl., Kans., Okla. and Tex. P. perplexus Cr. has
been confused with carolina (L.) in the past. Some of the parasite-predator records and
biological references listed under the latter species may actually refer to perplewus.
Polistes perplexus Cresson, 1870. Amer. Ent. Soe., Trans. 4: 245. d.
Polistes generosus Cresson, 1870. Amer. Ent. Soc., Trans. 4: 246. ¢.

UNPLACED TAXON OF POLISTES LATREILLE
-

nigripennis (DeGeer). Pa.
Vespa nigripennis Degeer, 1773. Mem. Serv. Hist. des Ins., v. 3, p. 582, pl. 29, fig. 10.

Bequaert suggested that this was possibly the same as rubiginosus (carolina).

1516 Hymenoptera in America North of Mexico

SUBFAMILY POLYBIINAE
Revision: Bequaert, 1933. Ent. Amer. (n. s.) 13: 87-150, 3 pls. (Nearctic taxa).
Genus MISCHOCYTTARUS Saussure

The nests are similar to those of Polistes, but average somewhat smaller. They consist of a
single free comb of hexagonal paper cells and are attached by a short pedicel in North American
forms to a natural or man-made support usually some distance from the ground. Each colony is
generally started by a single fertile female or queen.

Revision: Richards, 1945. Roy. Ent. Soc., London, Trans. 95: 295-462, 4 pls., 119 figs. (New
World spp.).

Biology: Jeanne, 1972. Mus. Comp. Zool., Bul. 144: 63-150, 46 figs., 4 pls. (biology, social
behavior of M. drewseni Sauss.).

Genus MISCHOCYTTARUS Subgenus MISCHOCYTTARUS Saussure

Mischocythorus (!) Saussure, 1853. Etudes sur la famille des Vespides, v. 1, Introduct. p.
VIII. Nom. nud.
Mischocyttarus Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 19.
Type-species: Zethus labiatus Fabricius. Desig. by Ashmead, 1902.

The species occur in Mexico, Central America, and South America.

Genus MISCHOCYTTARUS Subgenus KAPPA Saussure

Kappa Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 200.
Type-species: Polybia injucunda Saussure. Desig. by Bequaert, 1933.

cubensis cubensis (Saussure). Ga., Fla., Ala; Bahamas, Cuba.

Polybia cubensis Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 202. 6, 9.

Polybia phthisica Cresson, 1865. Ent. Soc. Phila., Proc. 4: 167. d, 2.
cubensis mexicanus (Saussure). Tex. (Brownsville); Mexico, Central America.

Polybia mexicana Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 203. 2.
flavitarsis centralis Bequaert. Utah, Colo., Nebr., N. Mex., western Tex.

Mischocyttarus flavitarsis var. centralis Bequaert, 1933. Ent. Amer. (n. s.) 13: 129. 2, 6.
flavitarsis flavitarsis (Saussure). Calif., Oreg., Utah, Colo., Nebr., Ariz.

Polybia flavitarsis Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 199. °.

Biology: Smith, 1944. Pan-Pacific Ent. 20: 80 (hibernation site). —Snelling, 1953. Kans. Ent.

Soce., Jour. 26: 143-145 (hibernation, nest size and site).
flavitarsis idahoensis Bequaert. B. C., Wash., Oreg., Idaho, Mont., Nev., Utah, Colo.

Mischocyttarus flavitarsis var. idahoensis Bequaert, 1933. Ent. Amer. (n. s.) 13: 183. 2, d.
flavitarsis kaibabensis Bequaert. Ariz. (Grand Canyon, Kayenta).

Mischocyttarus flavitarsis var. kaibabensis Bequaert, 1933. Ent. Amer. (n. s.) 13: 133. 9.
flavitarsis navajo Bequaert. Ariz.; Mexico (Chihuahua).

Mischocyttarus flavitarsis var. navajo Bequaert, 1933. Ent. Amer. (n. s.) 13: 181. 2, ¢.

Genus BRACHYGASTRA Perty

Brachygastra Perty, 1833. Delect. Anim. Articul. Brasil, p. 145.
Type-species: Brachygastra analis Perty Desig. by Bequaert, 1933 (=Polistes
lecheguana Latreille).

Nectarina Swainson and Shuckard, 1840. On the history and natural arrangement of
insects, p. 183. N. name for Brachygastra.

Melissaia Shuckard, 1841. Jn White, Ann. and Mag. Nat. Hist. 7: 320. N. name for
Nectarina.

Caba Thering, 1904. Rev. Mus. Paulista 6: 103. N. name for Nectarina.

The genus is essentially Neotropical but one species extends its range into southern United
States. The perennial paper nests are attached to branches of low trees and are begun by small

Superfamily VESPOIDEA 1517

swarms of fertile queens and workers. An average nest is subglobular, is enclosed in a single
sheet of papery material, is about 1 foot in diameter, contains about 50,000 cells arranged in
layers, has several entrances, and has spiral ramps connecting the tiers of cells. The adults col-
lect and store honey but do not cap the cells.

Revision: Buysson, 1905. Soc. Ent. France, Ann. 74: 537-566, 6 pls. —Naumann, 1968. Kans.
Univ. Sci. Bul. 47: 929-1003, 69 figs. (New World spp.).
mellifica (Say). Southern Tex. and Ariz. to Costa Rica and Panama (?).
Polistes mellifica Say, 1837. Boston Jour. Nat. Hist. 1 (pt. 4): 390. d, @.
Nectarina mellifera Dalla Torre, 1904. Gen. Ins., fase. 19, p. 86. Emend.
Chartergus aztecus Cameron, 1906. Invertebrata Pacifica 1: 154. 2. Preoce.
Chartergus arizonaensis Cameron, 1907. Invertebrata Pacifica 1: 181. 2.
Chartergus centralis Cameron, 1907. Invertebrata Pacifica 1: 181. 9.
Nectarina cameroni Meade-Waldo, 1911. Ann. and Mag. Nat. Hist. (8) 7: 112. N. name.

Biology: Barber, 1905. Ent. Soc. Wash., Proc. 7: 25 (nest). —Schwarz, 1929. Nat. Hist. 29:
421-426, 5 figs. (nest, honey production).

- Genus POLYBIA Lepeletier

Polybia Lepeletier, 1836. Hist. Nat. Ins. Hym., v. 1, p. 533.
Type-species: Polistes liliacea Fabricius Desig. by Ashmead, 1902.
Myrapetra White, 1841. Ann. and Mag. Nat. Hist. 7: 320.
Type-species: Myrapetra scutellaris White. Monotypic.
Myraptera Curtis, 1844. Linn. Soe. London, Proc., p. 188. Emend.
Alpha Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 167.
Type-species: Polybia bifasciata Saussure. Desig. by Bequaert, 1944.
Eupolybia Dalla Torre, 1904. In Wytsman, Gen. Ins., fase. 19, p. 76.
Type-species: Polistes liliacea Fabricius. Present desig. by Richards.

The species of Polybia build phragmocyttarous nests which consist of one to many combs at-
tached to a branch, leaf or rock, surrounded by an envelope of carton or of mud containing an
exit hole at the side or bottom of the comb. A number of species enlarge the nest by adding suc-
cessive combs, each covered by the envelope, so that access to the upper combs is through the
exit-holes of earlier stages of the nest. Some of the taxa store large quantities of honey, or sex-
ual castes of ants or termites, some of which are fed to larvae and some of which maintain the
adults during hibernation. Larvae may also be fed on partially masticated insects.

Taxonomy: Richards, 1951. Jn Richards and Richards, Royal Ent. Soc. London, Trans. 102:
129-150, figs. 11-16 (taxa allied to occidentalis (Oliv.) with key including the two taxa
occurring in America north of Mexico).

diguetana Buysson. Ariz. (Nogales); Mexico, Central America, Peru. Ecology: Buysson noted
that the nest was like that of typical occidentalis which consists of as many as 6 combs
surrounded by a carton envelope.
Polybia occidentalis var. diguetana Buysson, 1905. Soc. Ent. France, Bul. p. 9. 2, %, d.
occidentalis nigratella Buysson. Ariz. (Nogales); Mexico, Guatemala, Honduras. Typical
occidentalis (Oliv.) oceurs in South America.
Polybia occidentalis var. nigratella Buysson, 1905. Soc. Ent. France, Bul., p. 9. 2, 2.
Polybia occidentalis nausicaa Richards, 1951. In Richards and Richards, Royal Ent. Soe.
London, Trans. 102: 140. 2, d. N. syn. (O. W. Richards).

SUBFAMILY VESPINAE

This is morphologically the most specialized subfamily of the social wasps. Component species
are commonly called hornets (those nesting above ground in North America) and yellow jackets
(those nesting usually subterraneously). The nests consist of several to many combs of hex-
agonal cells composed of paper; cells constructed early in the year may be used for several lar-
vae in succession. The combs are usually enclosed in a paper envelope. The nests are annual, new
queens and males being produced late in the summer or early in the fall; the newly fertilized
queens overwinter and begin new nests in the spring. There are relatively few precise records

1518 | Hymenoptera in America North of Mexico

of the insects used to feed vespine larvae; apparently dismembered and masticated adult Dip-
tera and honeybees are commonly used; the wasps may also obtain bits of flesh from fresh and
decaying carcasses. Adult vespines feed on liquid foods, primarily nectar or honey; some species
are known to prey upon adult honeybees, which they kill and extract nectar from the crop.

Revision: Bequaert, 1932. Ent. Amer. (n. s.) 12: 71-138, 6 figs. (N. Amer. taxa).

Taxonomy: McFarland, 1888. Amer. Ent. Soc., Trans. 15: 297 (key). —Bequaert, 1930.
Brooklyn Ent. Soc., Bul. 25: 59-70 (generic and subgeneric classification). —Bequaert, 1935.
Brooklyn Ent. Soc., Bul. 30: 119-124 (check list, correction, additions). —Duncan, 1939.
Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1): 85-97 (genera). —Bequaert, 1941.
Brooklyn Ent. Soc., Bul. 36: 111-117 (corrections, additions).

Biology: Duncan, 1939. Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1): 98-176 (composite
summary for American spp.).

Genus VESPA Linnaeus

Vespa Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 343.
Type-species: Vespa crabro Linnaeus. Desig. by Lamarck, 1801.
Macrovespa Dalla Torre, 1904. Jn Wytsman, Gen. Ins., fase. 19, p. 64.
Type-species: Vespa crabro Linnaeus. Desig. by Bequaert, 1930.

The giant hornets are conspicuous members of the wasp fauna in the Palaearctic and Oriental
Regions; one taxon is adventive in North America. The species of Vespa are quite diverse in
nesting habits; some construct aerial nests in trees, others nest in sheltered sites above ground,
such as in hollow trees, and still others are subterranean. The nests, especially in exposed situa-
tions, are usually enclosed in a paper envelope.

crabro germana Christ. Ont., Mass., R. I., Conn., N. Y., N. J., Pa., Del., Md., D. C., Va., N. C., S.
C., Ga., Ala., W. Va., Ohio, Ind., Ky., Tenn., Wis., N. Dak., S. Dak.; central and western
Europe. Ecology: Nests in very sheltered sites above ground, frequently in hollow trees,
attics, porches, sheds, and rarely in underground cavities; exposed nests have a
complete paper envelope, but in quite sheltered sites the envelope may be only partial
or completely lacking. Adventive from Europe in mid-1800’s. The official common name
is the giant hornet; in some rural areas of the mid-Atlantic states it is known as the
hybrid bee. Typical crabro L. and other subspecies occur in the Palaearctic Region.

Vespa crabro germana Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u.
Ameisengeschl., p. 215.
Vespa crabro vulgata Birula, 1925. Arch. f. Naturgesch. 90, Abt. A, H. 12, p. 100. 2, %, 6.

Taxonomy: Bequaert, 1931. Konowia 10: 101-109 (color forms of crabro L. with key).

Biology: Beutenmuller, 1898. N. Y. Ent. Soc., Jour. 6: 199, pls. 9-10 (nest). —Felt, 1915. N. Y.
State Mus., Bul. 180: 71-73, pl. 2 (nest). —Cory, 1931. Jour. Econ. Ent. 24: 50-52 (nest,
injury to lilac, apples). —Ikan et al., 1969. Jour. Insect Physiol. 15: 1703-1712 (queen
pheromones). —Islay and Schaudinschky, 1973. Acoust. Soc. Amer., Jour. 53: 640-649
(acoustical communication).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Coll. 99 (14): 49-50, pl. 17, figs. I-Q, pl. 18,
figs. A-D (male genitalia).

Genus VESPULA Thomson

These smaller social vespines are restricted to the Holarctic Region.
Revision: Miller, 1961. Canad. Ent., Sup. 22, v. 93: 1-52, 84 figs., 15 maps.

Taxonomy: Sladen, 1918. Ottawa Nat. 32: 71-72 (key to Canadian spp.). —Miller, 1958 (1956).
Tenth Internat]. Cong. Ent., Proc. 1: 257-264 (nomenclature and distribution). —Fluno,
1973. Ent. Soc. Wash., Proc. 75: 80-83, 1 fig. (differentiation of subgenera by attraction to

chemical).

Biology: Taylor, 1939. Ent. Soc. Amer., Ann. 32: 304-315 (origin and evolution of social
parasitism). —Gaul, 1941. Psyche 48: 16-19 (artificial housing of colonies). —Gaul, 1942.

Superfamily VESPOIDEA 1519

Brooklyn Ent. Soc., Bul. 37: 57-61 (brood rearing). —Morse and Gary, 1961. Bee World 42:
179-181 (predation on honeybees). —Akre, Hill and MacDonald, 1973. Jour.Econ. Ent. 66:
803-805, 4 figs. (artificial housing of colonies).

Genus VESPULA Subgenus VESPULA Thomson

Vespula Thomson, 1869. Opuse. Ent., v. 1, p. 79.
Type-species: Vespa austriaca Panzer. Desig. by Ashmead, 1902.
Pseudovespa Schmiedeknecht, 1881. Ent. Nachr. 7: 314.
Type-species: Vespa austriaca Panzer. Monotypic.
Paravespula Bluethgen, 1938. Konowia 16: 271.
Type-species: Vespa vulgaris Linnaeus. Orig. desig.
Allovespula Bluethgen, 1943. Stettin. Ent. Ztg. 104: 149.
Type-species: Vespa rufa Linnaeus. Orig. desig.

Nests are usually underground or close to it in hollow logs or stumps. Combs, except for a
small conical area around the main suspensorium, are usually essentially plane. Members of this
subgenus are commonly called yellow jackets.

acadica (Sladen). Transcontinental in Canadian Zone from Newfoundland to Yukon Terr. and
Alaska, Maine to Minn., west of 100th meridian south to N. Mex. and Calif. Ecology:
Location of nests variable, sometimes aerial, subterranean or in rotten logs.
Vespa rufa var. americana Buysson, 1905. Soe. Ent. France, Ann. 63: 499. d. Preoce.
Vespa acadica Sladen, 1918. Ottawa Nat. 32: 72. d, 2, 9.
Vespula rufa sladeni Bequaert, 1931. Ent. Amer. (n. s.) 12: 102. d, 2, 8.

Biology: Sladen, 1918. Ottawa Nat. 32: 72 (nest site). —MacDonald, Akre and Hill, 1975.
Kans. Ent. Soc., Jour. 48: 114-121 (nest location and structure).
atropilosa (Sladen). Canada and U. S. west of 100th meridian mostly in Canadian and
Transition Zones; Mexico (Baja California). Ecology: Most nests are subterranean and
usually in rodent burrows. Parasite: Sphecophaga vesparum burra (Cr.); Triphleba
lugubris (Meig.).
Vespa atropilosa Sladen, 1918. Ottawa Nat. 22: 72. 2, ©.

Biology: MacDonald, Akre and Hill, 1974. Melanderia 18: 1-93, 13 figs. (comparative biology
and behavior). —MacDonald, Akre and Hill, 1975. Kans. Ent. Soc., Jour. 48: 53-63 (nest
associates). —MacDonald, Akre and Hill, 1975. Kans. Ent. Soc., Jour. 48: 114-121, 2 figs.
(nest location and structure).

austriaca (Panzer). Que., Ont., Man., Sask., Alta., B. C., N. W. T., Alaska, Maine, INo Gen IN lo dle
Colo., N. Mex., Utah, Calif., Oreg., Wash., Idaho; Holarctic. Host: Inquiline in nests of
rufa (L.) in Europe; North American host(s) unknown. The worker caste is absent.
Vespa austriaca Panzer, 1799. Faunae Ins. German., h. 68, pl. 2. 3.
Vespa borealis Smith, 1843. Zoologist 1: 170. 2. Preoce.
Vespa arborea Smith, 1849. Zoologist 7, App., p. lx. N. name.
Vespa infernalis Saussure, 1853. Etudes sur la famille des Vespides, v. 2, p. 189. 9.
Vespa tripunctata Packard, 1870. Chicago Acad. Sci., rans: 20s6er
consobrina (Saussure). Transcontinental in Canada and northern U. S. in Canadian and
Transition Zones, W. Va., N. C., Ga., Colo., N. Mex., Ariz., Calif. Ecology: Nests are
usually subterranean, but occasionally are in shrubs just above the ground surface.
Vespa consobrina Saussure, 1864. Etudes sur la famille des Vespides, v. 2, p. 141. &.
Vespa scelesta McFarland, 1888. Amer. Ent. Soc., Trans: 15: 298: 6, 9, %-
Vespa sulcata Howard, 1901. Insect Book, pl. 6, fig. 18. 8.

Biology: Dow, 1930. Boston Soc. Nat. Hist., Bul. 56: 12, 1 fig. (nest).
germanica (Fabricius). N. Y., N. J., Pa., Del., Md.; Europe. Ecology: Nests in sheltered
situations, frequently in structures. Adventive from Europe, apparently in mid-20th
century. References to taxonomy, biology and morphology of the Palaearctic population
will be found in Guiglia, 1973 (1972), Les Guepes Sociales d’Europe Occidentale et
Septentrionale, vol. 6 of Faune de l'Europe et du Bassin Mediterraneen.
Vespa Germanica Fabricius, 1793. Ent. System., v. 2, p. 256.

1520 Hymenoptera in America North of Mexico

Taxonomy: Menke and Snelling, 1975. U. S. Dept. Agr., Coop. Ins. Rpt. 25: 193-200, 33 figs.
(characters to distinguish from native North American spp.).
intermedia (Buysson). Newfoundland, Que., Man., Sask., Alta., N. W. T., Yukon Terr., Alaska in
Hudsonian Zone.
Vespa rufa var. intermedia Buysson, 1905. Soc. Ent. France, Ann. 73: 591. d, 2.

maculifrons (Buysson). Southern Canada and U. S. east of 100th meridian in Transition and

Austral Zones. Ecology: Nests are usually subterranean, occasionally in decayed stumps
or logs; nest usually contains 4-8 combs, rarely only a single comb. It may live as an
inquiline in nests of vulgaris (L.) and interbreed with it. Parasite: Vespula squamosa
(Dru.). V. maculifrons was frequently misidentified as germanica in publications prior
to 1931.

Vespa maculifrons Buysson, 1905. Soe. Ent. France, Ann. 63: 608.

Vespa communis var. flavida Sladen, 1918. Ottawa Nat. 32: 71. 2.

Biology: Couper, 1870. Canad. Ent. 2: 49-53 (nest). —Marlatt, 1891. Ent. Soc. Wash., Proc. 2:
80-83 (nest). —Rau and Rau, 1918. Wasp Studies Afield, pp. 295-297, fig. 53 (foraging,
nest). —Beamer, 1925. Ent. News 36: 309-310 (hibernation female). —Rau, 1930. Ent.
News 41: 185-190, pl. 19 (life history, nest). —Rau, 1931. Brooklyn Ent. Soc., Bul. 26: 85-89,
pl. 4 (nest, colonization). —Bromley, 1931. N. Y. Ent. Soc., Jour. 39: 129 (foraging,
predation). —Gaul, 1948. Brooklyn Ent. Soc., Bul. 43: 37-41 (interspecific tolerance).
—Haviland, 1962. Ent. Soc. Wash., Proc. 64: 181-183 (colony size). —Kurezewski, 1968. N.
Y. Ent. Soc., Jour. 76: 84-86 (predation). —Balduf, 1968. Ent. Soc. Wash., Proc. 70: 336-338,
1 fig. (inquilinism and interbreeding with vulgaris). —Simon and Benton, 1968. Ent. Soc.
Amer., Ann. 61: 542 (winter activity in aerial nest). —Green, Heckman, Benton and Coon,
1970. Ent. Soc. Amer., Ann. 63: 1197-1198, 2 figs. (exposed aerial nest). —MacDonald and
Matthews, 1975. Science 190: 1003-1004, 1 fig. (parasite).

pensylvanica (Saussure). Canada and western U. S. west of 100th meridian in Canadian and
Transition Zones; Mexico (Michoacan, Mexico). Ecology: Nests are subterranean.
Parasite: Bareogonalos canadensis (Harr.); Sphecophaga vesparum burra (Cr.);
Triphleba lugubris (Meig.).
Vespa pensylvanica Saussure, 1857. Stettin. Ent. Ztg. 18: 117. °@.
Vespa occidentalis Cresson, 1874. Amer. Ent. Soc., Trans. 5: 100. 2, %. Preoce.

Biology: Snow, 1882. Psyche 3: 339 (predation). —Metzmain, 1903. Entomologist 36: 137
(female hibernation). —Duncan, 1939. Stanford Univ., Pubs., Univ. Ser. Biol. Sei. 8 (1):
98-173 (nest, life history). —Smith, 1956. Ent. News 67: 141-146, 1 pl. (nest). —Chapman,
1963. Ecology 44: 766-767 (predation). —MacDonald, Akre and Hill, 1974. Melanderia 18:
1-93, 13 figs. (comparative biology and behavior).

Morphology: Duncan, 1939. Stanford Univ., Pubs., Univ. Ser. Biol. Sci. 8 (1): 13-84, pls.
I-XXVII (female, male).

squamosa (Drury). Ont., N. Y. to Wis. and Iowa, south to Fla. and Tex., most common in

Austral Zones; Mexico, Guatemala. Host: Vespula maculifrons (Buyss.), V. vidua
(Sauss.); the sgwamosa queen is a temporary, apparently facultative social parasite.

Vespa squamosa Drury, 1773. Illus. Nat. Hist., Index to pt. 1 published with pt. 2.

Vespa lineata Fabricius, 1775. Systema Ent., p. 365.

Vespa conchacea Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u. Ameisengeschl., p.
259.

Vespa cuneata Fabricius, 1804. Systema Piezatorum, p. 258.

Vespa cruciata Lepeletier, 1836. Hist. Nat. Ins. Hym., v. 1, p. 513. Emend. of cuneata.

Vespa bistriata MacFarland, 1888. Amer. Ent. Soc., Trans. 15: 298. ?. Preoce.

Vespa macfarlandi Lewis, 1897. Amer. Ent. Soc., Trans. 24: 172. 2. N. name.

Vespula squamosa var. (or subsp.) michoacana Bequaert, 1941. Ent. News 52: 249. 8.

Taxonomy: Turner, 1908. Psyche 15: 1-3, 1 pl. (coloration worker, male).

Biology: Taylor, 1939. Ent. Soe. Amer., Ann. 32: 310-313 (temporary parasitism by female in
nest of vidwa). —Gaul, 1947. Brooklyn Ent. Soc., Bul. 42: 87-96 (behavior, nest). —Gaul,
1948. Brooklyn Ent. Soc., Bul. 43: 37-41 (interspecific tolerance). —Tissot and Robinson,
1954. Fla. Ent. 37: 73-92 (aerial and subterranean nests). —MacDonald and Matthews,
1975. Science 190: 1003-1004, 1 fig. (host, nest).

Superfamily VESPOIDEA 1521

sulphurea (Saussure). Oreg., Calif., Nev., Ariz.; Mexico (Baja California). Ecology: Nests are
subterranean.
Vesna sulphurea Saussure, 1854. Etudes sur la famille des Vespides 2: 137. °.

Biology: Bequaert, 1931. Ent. Amer. (n. s.) 12: 112 (nest).
vidua (Saussure). N. S., N. B., Ont., in northeastern U.S. from Maine to N. Dak., south to Va.
and Iowa, south in Appalachian Mts. to N. C., Ga., mostly in Transition and U. Austral
Zones. Ecology: Nests are subterranean. Parasite: Sphecophaga vesparum burra (Cr.);
Vespula squamosa (Dru.), the queen of which is a temporary social parasite.
Vespa vidua Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 136.

Biology: Taylor, 1939. Ent. Soc. Amer., Ann. 32: 310-313 (nest, parasitism by squamosa).
vulgaris (Linnaeus). Alaska, Canada, U.S. south to Calif., Ariz., N. Mex., S. Dak., Iowa, Ind.,

Ill., Ohio, N. C., mostly in Canadian and Transition Zones; Mexico (Michoacan, Mexico);
widely distributed in Palaearctic Region; Ecology: Nests are usually subterranean or in
stumps with paper envelope present or absent; aerial nests are close to ground and
always with envelope. adventive in New Zealand. Parasite: Sphecophaga vesparum
burra (Cr.). May interbreed with maculifrons (Buyss.).

Vespa vulgaris Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 572. 9.

Vespa communis Saussure, 1857. Stettin. Ent. Ztg. 18: 117. °.

Vespa alascensis Packard, 1870. Chicago Acad. Sci., Trans. 2: 27. °.

Vespa westwoodii Shipp, 1893. Psyche 6: 450.

Biology: Bequaert, 1931. Ent. Amer. (n. s.) 12: 92-93 (nest). —Balduf, 1968. Ent. Soc. Wash.,
Proc. 70: 332-336, 1 fig. (nest, life history).

Genus VESPULA Subgenus DOLICHOVESPULA Rohwer

Dolichovespula Rohwer, 1916. Conn. State Geol. and Nat. Hist. Survey, Bul. 22: 642.
Type-species: Vespa maculata Linnaeus. Orig. desig.
Pseudovespula Bischoff, 1931. Gesell. Naturf. Freunde Sitzber. (1930), p. 346.
Type-species: Vespa norwegica var. adulterina Buysson. Orig. desig.
Boreovespula Bluethgen, 1943. Stettin. Ent. Ztg. 104: 149.
Type-species: Vespa norwegica Fabricius. Orig. desig.
Metavespula Bluethgen, 1943. Stettin. Ent. Ztg. 104: 149.
Type-species: Vespa silvestris Scopoli. Orig. desig.

Nests are usually aerial, suspended at some height or in bushes near the ground. Combs are
usually turned up at the margins so as to be concave above.

albida (Sladen). Labrador, Newfoundland, Que., Maine, Man., Alta., B. C., N. W. T., Yukon
Terr., Alaska, chiefly in Hudsonian Zone. Ecology: Nests are small and subterranean.
Vespa marginata Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 265. 2. Preoce.
Vespa albida Sladen, 1918. Ottawa Nat. 32: 71. d, 8.

Biology: Sladen, 1919. Rpt. Canad. Arctic Exped. 1913-18, v. 3, p. 26g (nest).
arctica Rohwer. Newfoundland to Yukon Terr., Alaska, in lower U. S. chiefly in Canadian
Zone. Host: Vespula arenaria (F.). The worker caste is absent.
Vespa borealis Lewis, 1897. Amer. Ent. Soc., Trans. 24: 171. d, 2. Preoce.
Vespula arctica Rohwer, 1916. Conn. State Geol. and Nat. Hist. Survey, Bul. 22: 642. N.
name.

Biology: Wheeler and Taylor, 1921. Psyche 28: 135-144, 3 figs. (permanent social parasitism
of arctica in nests of arenaria). —Wheeler, 1939. Ent. Soe. Amer., Ann. 32: 305-310 (social
parasitism in arenaria nests).

arenaria (Fabricius). In Canada from Newfoundland to Yukon Terr., Alaska, in lower U. S.
transcontinental chiefly in Canadian and Transition ones. Ecology: Nests are usually
aerial but placed close to ground in shrubbery, occasionally under eaves or roofs, rarely
the nests may be attached to tree roots and only partly exposed. Parasite: Sphecophaga
vesparum burra (Cr.); Vespula arctica Roh.

Vespa arenaria Fabricius, 1775. Systema Ent., p. 365.
Vespa borealis Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 264. Preoce.

1522 Hymenoptera in America North of Mexico

Vespa diabolica Saussure, 1854. Etudes sur la famille des Vespides, v. 2, p. 1S Sauer aor
Vespa fernaldi Lewis, 1897. Amer. Ent. Soc., Trans. 24: 171. 9, &.

Biology: Wheeler and Taylor, 1921. Psyche 28: 135-144, 3 figs. (nest, parasite). —Hungerford,
1930. Ent. News 41: 329-330, 1 pl. Mest). —Taylor, 1939. Ent. Soc. Amer., Ann. 32: 305-310
(nest, parasite). —Gaul, 1941. N. Y. Ent. Soc., Jour. 49: 367-369 (taste sensitivity of adults,
larvae). —Gaul, 1941. Psyche 48: 16-19 (colony housing, interspecific tolerance). —Gaul,
1942. Brooklyn Ent. Soc., Bul. 37: 57-61 (brood rearing, mating). —Gaul, 1948. Brooklyn
Ent. Soc., Bul. 43: 73-79 (distribution of labor). —Sailer, 1950. Kans. Ent. Soc., Jour. 23:
134-137, 3 figs. (nest temperature). —Gaul, 1952. N. Y. Ent. Soc., Jour. 60: 17-20 (flight in
stormy weather). —Gaul, 1952. Brooklyn Ent. Soc., Bul. 47: 79-92 (temperature regulation
in nest). —Gibo, 1972. N. Y. Ent. Soc., Jour. 80: 105-108 (hibernation sites, temperature
tolerance). —Gibo, Yarascavitch and Dew, 1974. Canad. Ent. 106: 503-507, 1 fig. (colony
thermoregulation). —Gibo, Dew and Hajduk, 1974. Canad. Ent. 106: 873-879, 2 figs.
(relation of colony biomass to calorie production).

maculata (Linnaeus). In Canada from Nova Scotia to Yukon Terr., Alaska, in lower U. S. from

Maine to Fla., west to 100th meridian in Canadian, Transition, U. and L. Austral Zones,
west of 100th meridian chiefly in Canadian, Transition and U. Sonoran Zones. Ecology:
Nests are always aerial and are usually suspended from limbs or beneath eaves and
roofs. Parasite: Sphecophaga vesparum burra (Cr.). The official common name is the
bald-faced hornet; sometimes it is also called the white-faced hornet.

Vespa maculata Linnaeus, 1763. Centuria Ins. Rar., p. 30.

Vespa maculata americana Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u.
Ameisengeschl., p. 239.

Biology: Couper, 1870. Canad. Ent. 2: 49-53 (mest). —Fyles, 1903. 33rd Ann. Rpt., Ent. Soe.
Ontario, pp. 69-70, figs. 47-48 (nest). —Howard, 1915. Ent. Soc. Wash., Proc. 17: 148 (Mest
color). —Rau and Rau, 1918. Wasp Studies Afield, pp. 297-298 (predation). —Davis, 1919.
Brooklyn Ent. Soc., Bul. 14: 119-123, 1 pl. (nest). —parker, 1928. Ent. Soc. Wash., Proc. 30:
14 (succession of brood in one cell). —Rau, 1929. Ent. Soc. Amer., Ann. 22: 659-675, 10 figs.
(esting habits). —Bromley, 1931. N. Y. Ent. Soc., Jour. 39: 126-128 (predation). —Betz,
1932. Quart. Rev. Biol. 8: 197-209, 4 figs. (colony size). —Rau, 1934. Brooklyn Ent. Soc.,
Bul. 19: 170 (hibernating female). —Rau, 1934. Brooklyn Ent. Soc., Bul. 19: 171 (predation).
—Balduf, 1936. Canad. Ent. 68: 138-139 (colony size). —Gaul, 1941. Psyche 48: 16-19
(colony housing, interspecific tolerance). —Gaul, 1952. N. Y. Ent. Soc., Jour. 60: 21-24
(metabolic cycles and flight). —Balduf, 1954. Ent. Soc. Amer., Ann. 47: 445-458, 3 pls. (nest
structure and growth, foraging, life history, population size). —Gibo, 1972. N. Y. Ent. Soc.,
Jour. 80: 105-108 (hibernation sites, temperature tolerance). —Howell, 1973. Ent. News 84:
141-142 (predation on Vespula maculifrons (Buyss.)). —Gibo, Yarascavitch and Dew, 1974.
Canad. Ent. 106: 503-507, 1 fig. (colony thermoregulation). —Gibo, Dew and Hajduk, 1974.
Canad. Ent. 106: 873-879, 2 figs. (relation of colony biomass to calorie production).

Morphology: Bequaert, 1932. Ent. Amer. (n. s.) 12: 73-75, fig. 1 (male genitalia). —Snodgrass,
1941. Smithsn. Inst., Mise. Collect. 99 (14): 49, pl. 17, figs. A-H (male genitalia). —Hermann
and Krispyn, 1975. Ga. Ent. Soc., Jour. 10: 307-313, 8 figs. (venom apparatus).

norvegicoides (Sladen). In Canada from Newfoundland to Yukon Terr., Alaska, in lower U. S.
transcontinental chiefly in Canadian Zone. Ecology: Nests in low shrubbery.
Vespa norvegicoides Sladen, 1918. Ottawa Nat. 32: 71. d, 2.

Biology: Bequaert, 1932. Ent. Amer. (n. s.) 12: 119 (nest).

1523

Superfamily POMPILOIDEA

By KARL V. KROMBEIN

This superfamily includes two families, the very large and abundant Pompilidae and the very
small and rare Rhopalosomatidae.

Members of the Pompilidae are commonly called spider wasps because they prey exclusively
on spiders. Only a single larger spider is stored per cell, rather than a number of smaller spiders
as in the spider-preying sphecoid genera such as Trypoxylon, Trypargilum and Sceliphron. The
majority of species, including the more primitive forms, capture the spider before preparing a
burrow in the soil. Some Pepsinae make multicellular nests in pre-existing cavities in twigs or in
the ground, and some build individual mud cells which may be joined in a series; in both types
the cell is constructed before the spider is captured. Some genera, such as Pepsis, may use the
burrow of the prey spider as a nesting site. The peculiar genus Minagenia is a spider ec-
toparasite.

The Rhopalosomatidae are ectoparasites of nymphal crickets.

Family POMPILIDAE

The entire pompilid fauna of America north of Mexico has been adequately monographed in
recent years by Bradley, Evans, Hurd and Townes. References will be found under the headings
Pepsinae, Pepsis, Aporini, Pompilinae, Pompilini, and Ceropalinae.

Taxonomy: Ashmead, 1900-1902. Canad. Ent. 32: 145-155, 185-188, 295-296; 34: 79-88, 131-137
(keys to world genera). —Banks, 1912 (1911). N. Y. Ent. Soce., Jour. 19: 219-237 (eastern
species). —Banks, 1919. Mus. Compar. Zool., Bul. 63: 229-248 (western species). —Brimley,
1936. Elisha Mitchell Sci. Soc., Jour. 52: 107-131 (N. C. species). —Pate, 1946. Amer. Ent.
Soc., Trans. 72: 65-137 (generic names and type-species). —Dreisbach, 1948 (1946). Mich.
Acad. Sci., Arts, and Letters, Papers 32: 239-247 (key to Mich. genera). —Dreisbach, 1949.
Mich. Acad. Sci., Arts, and Letters, Papers 33: 63-71 (American genera). —Evans, 1959.
Ent. Soc. Amer., Ann. 52: 430-444, 51 figs. (larva). Except for Evans’ paper on larvae, the
foregoing references are obsolete for identification purposes; they are superseded by the
revisionary papers cited under several of the following subfamily, tribal and generic
headings.

Biology: Evans, 1953. Syst. Zool. 2: 155-172 (comparative ethology). —Evans and Yoshimoto,
1962. Ent. Soc. Amer., Misc. Pub. 3: 65-119 (ecology, nesting behavior and prey of
northeastern spp.). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 24-33

(prey selection factors).
SUBFAMILY PEPSINAE
Revision: Townes, 1957. U. S. Natl. Mus., Bul. 209: 1-220, 129 figs.

1524 Hymenoptera in America North of Mexico

TRIBE PEPSINI
Taxonomy: Dreisbach, 1952. N. Y. Ent. Soc., Jour. 60: 119-125 (key to New World genera).
Genus CHIRODAMUS Haliday

Chirodamus Haliday, 1837. Linn. Soc. London, Trans. (3) 17: 326.
Type-species: Chirodamus kingui Haliday. Monotypic.
Calopompilus Ashmead, 1900. Canad. Ent. 32: 188.
Type-species: Pompilus maculipennis Smith. Orig. desig.
Dinocnemis Banks, 1925. Mus. Compar. Zool., Bul. 67: 336.
Type-species: Pompilus fortis Cresson. Desig. by Bradley, 1944.
Onochares Banks, 1933. Psyche 40: 9.
Type-species: Onochares brazoria Banks (=Priocnemis Heiligbrodtii Cresson).
Trichocurgus Haupt, 1937. Ztschr. f. Naturw. 91: 127, 134.
Type-species: Pompilus monachus Smith. Orig. desig.
Chrysocurgus Haupt, 1937. Ztschr. f. Naturw. 91: 127, 134.
Type-species: Sphex nitida Fabricius. Orig. desig.
Derochilus Banks, 1941. Canad. Ent. 73: 119, 120.
Type-species: Pompilus validus Cresson. Orig. desig.
Reedimia Banks, 1946. Mus. Compar. Zool., Bul. 96: 482.
Type-species: Agenia hirsutula Spinola. Orig. desig.
Anacyphonyx Banks, 1946. Mus. Compar. Zool., Bul. 96: 520.
Type-species: Anacyphonyzx fidelis Banks. Orig. desig.
albopilosus (Cresson). N. Y. to Ga., W. Va. in Transition Zone. Ecology: Occurs in rich moist
woods from early to late summer.
Pompilus (Agenia) albopilosus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 125. ¢.
Priocnemis fortella Banks, 1915. Canad. Ent. 47: 401. 2.
Pseudagenia najacra Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 203. 3.

Taxonomy: Bradley, 1944. Canad. Ent. 76: 182 (synonymy of fortella, male description).

deceptus (Banks). Tex.
Priocnemis decepta Banks, 1926. Canad. Ent. 58: 201. 9.

feroculis (Banks). Va., Tex.
Pseudagenia feroculis Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 232. 6.

fortis (Cresson). N. Y. to S. C. in Transition Zone. Ecology: Occurs in woods.
Pompilus (Priocnemis) fortis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 113. .
Pompilus (Agenia) nigropilosus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 124. d.
Pseudagenia mariva Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 202. ¢d.

Taxonomy: Bradley, 1944. Canad. Ent. 76: 131-132 (male description).
heiligbrodtii (Cresson). Tex.
Priocnemis Heiligbrodtii Cresson, 1872. Amer. Ent. Soc., Trans. 4: 204. 2.
Agenia Belfragei Cresson, 1872. Amer. Ent. Soc., Trans. 4: 205. ¢.
Onochares brazoria Banks, 1933. Psyche 40: 9. 2.
maculipennis (Smith). Kans., Mo., Ark., Tex., Miss., Ala., Ga., N. C.
Pompilus maculipennis Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 157. @.
pyrrhomelas (Walker). B. C., Wash., Oreg., Calif., Idaho, Nev., Utah, Ariz., N. Mex., commonest
in Canad. Zone; Mexico. Prey: Antrodiaetus pacificus Simon.
Pompilus pyrrhomelas Walker, 1866. In Lord, Naturalist in Vancouver Isl. and B. C., v. 2,
p. 341. 9.
Cryptocheilus rugosus Banks, 1917. Mus. Compar. Zool., Bul. 61: 101. 9.
Cryptocheilus inaequalis Banks, 1917. Mus. Compar. Zool., Bul. 61: 102. 3.

Taxonomy: Bradley, 1944. Canad. Ent. 76: 132 (male description).

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 170 (prey).

validus (Cresson). N. C., 8S. C., Ga., Fla., Ala., in Lower Austral zone.
Pompilus (Priocnemis) validus Cresson, 1867, Amer. Ent. Soc., Trans. 1: 116. 2.

Superfamily POMPILOIDEA 1525
Genus PEPSIS Fabricius

The species, commonly called tarantula hawks, are most frequently encountered at the flowers
of various woody and suffrutescent plants of which the milkweed genus Asclepias appears to be
the preferred floral host. So far as is known, all the species are predaceous on spiders of the
family Theraphosidae (= Aviculariidae), most commonly on species of Bothriocyrtum,
Aphonopelma and Mygale. The wasps may prepare a burrow before hunting for prey, or they
may utilize the burrow of the tarantula itself as a nesting site.

Pepsis occurs only in the New World, and the majority of species are Neotropical.

Revision: Lucas, 1895 (1894). Berlin. Ent. Ztschr. 39: 449-839, pls. XXII-XXXIII (New World

spp.). —Fox, 1898. Ent. Soc. Wash., Proc. 4: 140-148 (U. S. spp.). —Hurd, 1952. Amer. Mus.
Nat. Hist., Bul. 98: 257-334, 49 figs., 2 tables (Nearctic spp., list of spp. in genus).

Taxonomy: Banks, 1921. Ent. Soc. Amer., Ann. 14: 22-23 (key to U.S. spp.).

Biology: Williams, 1956. Ent. Soc. Amer., Ann. 49: 447-466, 24 figs. (life history, prey, nesting
behavior). E

Morphology: Snodgrass, 1910. U. S. Natl. Mus., Proc. 39: figs. 61, 69 (lateral view of thorax
and base of abdomen, base of forewing).

Genus PEPSIS Subgenus PEPSIS Fabricius

Pepsis Fabricius, 1804. Systema Piezatorum, pp. 207-208.
Type-species: Sphex stellata Fabricius. Desig. by Latreille, 1810.
chrysothemis chrysothemis Lucas. Tex. to Okla. w. into Calif.; Mexico (Sonora, Baja
California). Prey: Aphonopelma sp.
Pepsis chrysothemis Lucas, 1894. Berlin. Ent. Ztschr. 39: 731, 739. d.
Pepsis cinnabarina Lucas, 1894. Berlin. Ent. Ztschr. 39: 782, 804. °.
Pepsis circularis Fox, 1898. Ent. Soc. Wash., Proc. 4: 142, 144. d.

Taxonomy: Hurd, 1952. Amer. Mus. Nat. Hist., Bul. 98: 312 (identity of circularis).

Biology: Williams, 1956. Ent. Soc. Amer., Ann. 49: 456 (prey, life history).
chrysothemis lucasii Fox. Tex.; Mexico (Tamaulipas).
Pepsis Lucasii Fox, 1898. Ent. Soc. Wash., Proce. 4: 141, 145. °.

mexicana Lucas. Tex. w. into south. Calif., s. to Colombia. Prey: Aphonopelma sp.
Pepsis mexicana Lucas, 1894. Berlin. Ent. Ztschr. 39: 560, 561, 566. d, 2.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 431-482, fig. 41 (larva).

Biology: Williams, 1956. Ent. Soc. Amer., Ann. 49: 456, 458 (prey, life history).
pallidolimbata pallidolimbata Lucas. Tex. to Kans. w. into south. Calif.; Mexico (Coahuila,
Chihuahua, Baja California).
Pepsis pallidolimbata Lucas, 1894. Berlin. Ent. Ztschr. 39: 733, 745. °.
Pepsis bequaerti Salman, 1928. Pan-Pacifie Ent. 5: 23. d.
pallidolimbata smithi Hurd. Calif. (west-central flanks of Sierra Nevada and Mt. Diablo-Mt.

Hamilton ranges). Prey: Aphonopelma sp.
Pepsis pallidolimbata smithi Hurd, 1948. Calif. Univ., Pubs., Ent. 8: 126, 142. 6, 2.

Biology: Williams, 1956. Ent Soc. Amer., Ann. 49: 456 (prey, life history).
thisbe Lucas. Tex. to Nebr. w. into Calif.; Mexico (Coahuila, Chihuahua, Sonora, Baja
California). Prey: Aphonopelma sp., Avicularia californica Bks.
Pepsis thisbe Lucas, 1894. Berlin. Ent. Ztschr. 39: HO2 Od, MAL, Oe oe
Pepsis sayi Banks, 1926. Canad. Ent. 58: 202. d (2 misdet.).
Pepsis sherillae Hurd, 1948. Calif. Univ., Pubs., Ent. 8: 124, 146. 6.

Taxonomy: Williams, 1956. Ent. Soc. Amer., Ann. 49: 457, figs. 10-13 (egg, larva). —Evans,
1959. Ent. Soc. Amer., Ann. 52: 431, figs. 1-7, 40 (larva).

Biology: Cockerell, 1916. Canad. Ent. 48: 55 (prey). —Williams, 1956. Ent. Soc. Amer., Ann.
49: 452-456, figs. 2, 9-14, 19, 21, 23-26 (prey, hunting behavior, life history).

1526 Hymenoptera in America North of Mexico

Genus PEPSIS Subgenus STENOPEPSIS Banks

Pepsis subg. Stenopepsis Banks, 1945. Bol. Ent. Venezolana 4: 82.
Type-species: Pepsis hymenaea Moesary. Orig. desig.
venusta Smith. South. Ariz. to Brazil.
Pepsis venusta Smith, 1855. Cat. Hym. Brit., Mus., v. 3, p. 196. d.

Genus PEPSIS Subgenus DINOPEPSIS Banks

Pepsis subg. Dinopepsis Banks, 1945. Bol. Ent. Venezolana 4: 83.
Type-species: Pepsis grossa Fabricius. Orig. desig.
formosa formosa (Say). Mo., Tex. to Kans. w. into Ariz. and Nev.; Mexico (Chihuahua). Prey:
Aphonopelma chalcodes Chamb., Dugesiella sp.
Pompilus formosus Say, 1832. West. Quart. Rptr. 2: 76. 9.
Pepsis nephele Lucas, 1894. Berlin. Ent. Ztschr. 39: 732, 739. 9.
Pepsis pseudoformosa Cockerell, 1898. Davenport Acad. Nat. Sci., Proc. 7: 146. d.

Biology: Buckley, 1862. Ent. Soc. Phila., Proc. 1: 138-139 (prey, misdet. as Mygale).
—Lincecum, 1867. Amer. Nat. 1: 137-141, 2 figs. (prey, misdet. as Mygale). —Cazier and
Mortenson, 1964. Ent. Soc. Amer., Ann. 57: 533-541, 8 figs. (prey, nesting behavior).

formosa pattoni Banks. N. Mex., Ariz., Calif.; Mexico (Sonora, Baja California).
Pepsis pattoni Banks, 1941. Mus. Compar. Zool., Bul. 94: 181. 3, 9.

Genus PEPSIS Subgenus GIGANTOPEPSIS Lucas

Pepsis subg. Gigantopepsis Lucas, 1919. Arch. f Naturgesch., div. A 83: 10, 41.
Type-species: Pepsis gigantea Lucas. Orig. desig.
aquila Lucas. South. Ariz. and N. Mex.; Mexico (Durango).
Pepsis aquila Lucas, 1894. Berlin. Ent. Ztschr. 39: 797. 3.
arizonica Banks. Tex., Ariz., Calif.; Mexico (Chihuahua, Durango).
Pepsis arizonica Banks, 1921. Ent. Soc. Amer., Ann. 14: 21. d.
Pepsis hirsuta Salman, 1933. Pan-Pacific Ent. 9: 9. °.
marginata Palisot de Beauvois. South. Fla.; West Indies. Prey: Cyrtopholis portoricensis
Chamb.
Pepsis marginata Palisot de Beauvois, 1809. Ins. Afr., Amer., p. 94, pl. 2, figs. 2,3..d, °.
Pepsis heros Dahlbom, 1843. Hym. Europaea, v. 1, p. 122. °.

Biology: Petrunkevitch, 1926. Jour. Expt. Zool. 45: 367-394, 2 pls. (prey, hunting behavior,
nest, life history). —Petrunkevich, 1952. Sci. Amer. (for Aug.), pp. 20-23 (prey, hunting
behavior, nest, life history).

Genus PEPSIS Subgenus UNASSIGNED

angustimarginata Viereck. Tex. to Kans. w. to south. Calif.; Mexico (Sonora, Chihuahua).
Pepsis angustimarginata Viereck, 1908 (1907). Amer. Ent. Soc., Trans. 33: 398. °.
Pepsis sayi Banks, 1926. Canad. Ent. 58: 202. 2 (d misdet.).
azteca Cameron. Tex. south to Panama.
Pepsis azteca Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 215. °.
cerberus Lucas. Kans., Tex., Ariz.; Mexico (Chihuahua, Baja California).
Pepsis cerberus Lucas, 1894. Berlin. Ent. Ztschr. 39: 780, 790. d.
Pepsis inermis Fox, 1898. Ent. Soc. Wash., Proc. 4: 141, 146. °.
Pepsis novitia Banks, 1921. Ent. Soc. Amer., Ann. 14: 21. ¢. Apparently a hybrid between
cerberus and elegans Lep. with most characters as in cerberus.
Taxonomy: Hurd, 1952. Amer. Mus. Nat. Hist., Bul. 98: 295 (status of novitia).
elegans Lepeletier. Pa. to Kans., s. to Fla. and Tex. Ecology: Nests in rodent burrow. Prey:
Huge spider.
Pepsis elegans Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 489. d.
Pepsis dubitata Cresson, 1867. Amer. Ent. Soc., Trans. 1: 144. d, 2.

Biology: Rau and Rau, 1918. Wasp studies afield, pp. 67-71 (prey transport, nesting site).

Superfamily POMPILOIDEA 1527

Morphology: Salman, 1929. Amer. Ent. Soc., Trans. 55: 119-153, pls. VI-IX.
mildei Stal. Tex. to Kans. w. into Calif.; Mexico (Coahuila, Baja California). Ecology: Nests in
prey burrow. Prey: Bothriocyrtum californicum (Camb.), Aphonopelma sp.
Pepsis mildei Stal, 1857. Ofvers. Kongl. Vetensk. Akad., Forh. 14: 64.
Pepsis hesperiae Patton, 1894. Ent. Soe. Wash., Proc. 3: 46. ¢.
Pepsis Boguei Fox, 1898. Ent. Soc. Wash., Proc. 4: 141, 142, 146. 6, 9.

Biology: Passmore, 1933. Natl. Geog. Mag., 64: 203, 205 (prey capture, nest). —Passmore,
1936. Nat. Mag. 27: 155-159 (prey capture, nest; misdet. as formosa). —Hurd, 1952. Amer.
Mus. Nat. Hist., Bul. 98: 268 (prey). —Williams, 1956. Ent. Soc. Amer., Ann. 49: 458, 460,
figs. 4-8, 15-19 (prey capture, nest, life history).

saphirus Palisot de Beauvois. South. Fla.; West Indies.
Pepsis saphirus Palisot de Beauvois, 1805. Ins. Afr., Amer., p. 39, pl. 1, fig. 4. 9.

s Genus HEMIPEPSIS Dahlbom

Hemipepsis Dahlbom, 1844. Hym. Europaea, v. 1, p. 123.
Type-species: Hemipepsis capensis Dahlbom. Desig. by Ashmead, 1900.
Pallosoma Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 492.
Type-species: Pallosoma barbara Lepeletier. Desig. by Ashmead, 1900.
Tetraodontonyx Ashmead, 1900. Canad. Ent. 32: 187.
Type-species: Tetraodontonyx rufipes Ashmead. Orig. desig. (=Pompilus heros
Guerin).
Cryptocheilus subg. Tetracryptocheilus Zavattari, 1907. R. Univ. Torino, Mus. Zool.
Compar. Anat., Bol. 22 (555): 4.
Type-species: Cryptocheilus (Tetracryptocheilus) ascensi Zavattari. Orig. desig.
Tetracryptochilus Schulz, 1911. Zool. Ann. 4: 122. Emend.
Trichonyx Haupt, 1929. Rev. Zool. Bot. Africaines 17: 195.
Type-species: Hemipepsis unguicularis Kohl. Orig. desig.
Pachynimia Haupt, 1929. Rev. Zool. Bot. Africaines 17: 197, 202.
Type-species: Priocnemis tinctor Saussure. Orig. desig.
Hemipepsis subg. Xenopepsis Arnold, 1932. Transvaal Mus., Ann. 14: 291, 323, 367.
Type-species: Hemipepsis (Xenopepsis) commixta Arnold. Orig. desig.
Hemipepsis subg. Moropepsis Banks, 1934. Amer. Acad. Arts and Sci., Proc. 69: 6, 8.
Type-species: Hemipepsis (Moropepsis) croesus Banks. Monotypic.
Hovagenia Banks, 1941. Acad. Nat. Sci. Phila., Proc. 92: 343.
Type-species: Hovagenia saussurei Banks. Orig. desig.
mexicana (Cresson). Tex. (Davis Mts.), s. to Panama.
Mygnimia mexicana Cresson, 1867. Amer. Ent. Soc., Trans. 1: 143. 9.
toussainti (Banks). Ariz., N. Mex.; Mexico (Durango, Michoacan, Mexico); Haiti.
Mygnimia toussainti Banks, 1928. Studies on Cuban Insects, v. 1, p. 5. d.
ustulata ochroptera Stal. Cent. and south. Calif., Ariz., Nev., in U. Austr. and L. Austr. Zones.
Prey: Brachythele longitarsis Simon, Aphonopelma sp.
Hemipepsis ochroptera Stal, 1857. Ofvers. Svenska Vetensk. Akad., Forh. 14: 64.
Mygnimia hesperina Banks, 1917. Mus. Compar. Zool., Bul. 61: 102. d, °.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 432-433, fig. 42 (larva).
Biology: Williams, 1956. Ent. Soc. Amer., Ann. 49: 460-462, figs. 20, 22 (prey capture, nest,
life history).
ustulata ustulata Dahlbom. Tex., Okla., Kans., Colo., N. Mex., Utah, Ariz. in L. Austr. and U.
Austr. Zones, less common in Transit. Zone; Mexico (Baja California, Durango,
Michoacan, Hidalgo, Mexico).
Hemipepsis ustulata Dahlbom, 1843. Hym. Europaea, v. 1, p. 123. 2.
Mygnimia cressoni Banks, 1926. Canad. Ent. 58: 203. d, °.

Genus PRIOCNESSUS Banks

Priocnessus Banks, 1925. Mus. Compar. Zool., Bul. 77: 337.
Type-species: Salius neotropicalis Cameron. Desig. by Pate, 1946.

1528 Hymenoptera in America North of Mexico

Cressochilus Banks, 1941. Canad. Ent. 73: 119, 120.
Type-species: Pompilus nuperus Cresson. Orig. desig.
Amerocnemis Banks, 1945. Bol. Ent. Venezolana 4: 93.
Type-species: ‘Amerocnemis bequaerti Banks. Orig. desig.
apache (Banks). Tex., Ariz. in L. Austr. Zone.
Priocnemis (Priocnessus) apache Banks, 1933. Psyche 40: 11. °.
coloradensis (Banks). Colo., Tex.
Cryptocheilus coloradensis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 121. 9.
dakota (Cresson). Conn. and N. Y. to Ga., W. Va., Kans., “Dakota”.
Pompilus (Agenia) Dakota Cresson, 1867. Amer. Ent. Soc., Trans. 1: 124. ¢.
Pompilus (Agenia) Dakota pallidicornis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 124. 3.
Preoce.
Priocnemis (Priocnessus) kiowa Banks, 1933. Psyche 40: 12. °.

Taxonomy: Bradley, 1944. Canad. Ent. 76: 151-152 (female, male description, synonymy).
nebulosus (Dahlbom). Ont., U.S. to 100° W. in Transit., U. Austr., and L. Austr. zones.

Ecology: Occurs in open woods. Prey: Agelenopsis potteri (Blackw.), A. emertont Chamb.
and Ivie, A. pensylvanica (Koch), A. naevia (Walck.), A. sp.

Priocnemis nebulosus Dahlbom, 1843. Hym. Europaea, v. 1, p. 96. @.

Pompilus (Agenia) pulchrinus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 126. ¢.

Priocnemis subconicus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 556. 2.

Priocnemis leibyi Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 203. 3.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 43 (prey records). —Kurezewski, 1961.
Brooklyn Ent. Soc., Bul. 56: 23 (prey record). —Evans and Yoshimoto, 1962. Ent. Soc.
Amer., Misc. Pub. 3: 100-101 (ecology, prey records). —Kurezewski and Kurczewski, 1968.
Kans. Ent. Soc., Jour. 41: 3 (prey records).

nigricans Townes. Ariz.
Priocnessus nigricans Townes, 1957. U. S. Natl. Mus., Bul. 209: 47. °.
nuperus (Cresson). N. Y. to Ga., Kans., Tex. in U. and L. Austral zones.
Pompilus (Priocnemis) nuperus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 118. 2.

Taxonomy: Bradley, 1944. Canad. Ent. 76: 150-151 (male description).
Genus ENTYPUS Dahlbom

Entypus Dahlbom, 1843. Hym. Europaea, v. 1, p. 35.
Type-species: Entypus ochrocerus Dahlbom. Monotypic.
Priocnemioides Radoszkowski, 1888. Soc. Imp. Nat. Moscou, Bul. (n. s.) 2: 482.
Type-species: Pompilus fulvicornis Cresson. Desig. by Banks, 1944.
Prionocnemoides Dalla Torre, 1897. Cat. Hym., v. 8, p. 211. Emend.
Priocnemoides (!) Ashmead, 1900. Canad. Ent. 32: 187.
Cheilotus Bradley, 1946. Soc. Cubana Hist. Nat., Mem. 18: 124.
Type-species: Pompilus ignipennis Cresson. Orig. desig.

Taxonomy: Day, 1974. Ent. News 85: 92-94 (generic synonymy).
angusticeps (Townes), n. comb. Tex.; Mexico (Sinaloa). Prey: Lycosa antelucana Mont.
Priocnemioides angusticeps Townes, 1957. U. S. Natl. Mus., Bul. 209: 60, pl. 2, fig. 17. d,
oF
aratus (Townes), n. comb. Idaho, Utah, Ariz., N. Mex., Tex., Kans., in U. and L. Sonoran
Faunas; Mexico (Durango). Prey: Lycosa carolinensis Walck.
Priocnemioides aratus Townes, 1957. U. S. Natl. Mus., Bul. 209: 51. 2, 6.
austrinus austrinus (Banks), n. comb. Tex., Kans., Colo.
Cryptocheilus austrinus Banks, 1917. Mus. Compar. Zool., Bul. 61: 102. 9.
austrinus fuscatus (Townes), n. comb. Ala., Kans.
Priocnemioides austrinus fuscatus Townes, 1957. U. S. Natl. Mus., Bul. 209: 53. 2.
fulvicornis (Cresson). Atlantic to 100° W. in U. Austr. and L. Austr. Zones. Prey: Lycosa
avida Walck.
Pompilus (Priocnemis) fulvicornis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 112. ¢, 2.

Superfamily POMPILOIDEA 1529

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 59 (prey record).
magnus (Cresson), n. comb. Gulf and Atlantic States, Tex. to N. Y.
Pompilus (Priocnemis) magnus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 111. 2.
texanus atripennis (Townes), n. comb. La. (Opelousas).
Priocnemioides texanus atripennis Townes, 1957. U.S. Natl. Mus., Bul. 209: 56. d.
texanus texanus (Cresson), n. comb. Kans. to Tex., w. to Calif.; Mexico (Coahuila).
Priocnemis texanus Cresson, 1872. Amer. Ent. Soe., Trans. 4: 204. d, °.
unifasciatus californicus (Townes), n. comb. Calif.; Mexico (Baja California). Prey: Lycosa
pacifica Bks.
Priocnemioides unifasciatus californicus Townes, 1957. U. S. Natl. Mus., Bul. 209: 65, pl.
2p See

Biology: Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 393 (prey).
unifasciatus cressoni (Banks), n. comb. Kans. and Utah s. to Tex. and Ariz., s. to Guatemala.

Prey: Lycosa antelucana Mont.
Cryptocheilus cressoni Banks, 1929. Psyche 36: 326. 6, °.

Biology: Hurd and Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey record).
unifasciatus unifasciatus (Say), n. comb. Conn., N. Y. to Fla., w. to Wis., Ill, Kans. and Tex. in
U. Austral Zone. Ecology: Occurs in open woods and overgrown fields. Prey: Lycosa
rabida Walck., L. riparia Hentz.
Pompilus unifasciatus Say, 1828. American Entomology 3: 92. °.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 63 (ecology, prey record). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 100 (prey record).

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99, no. 14: 46-47, pl. 16,. figs. H-Q
(male genitalia).

Genus CRYPTOCHEILUS Panzer

Salius Fabricius, 1804. Systema Piezatorum, p. 124. Preoce.
Type-species: Sphex sexpunctata Fabricius. Desig. by Guerin, 1849.
Cryptocheilus Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 120.
Type-species: Sphex annulata Fabricius. Desig. by Westwood, 1840.
Cryptochilus Rafinesque, 1815. Analyse de la Nature., p. 125. Emend.
Adonta Billberg, 1820. Enum. Ins., p. 101. N. name for Salius.
Stenoclavelia Arnold, 1932. Transvaal. Mus., Ann. 15: 44.
Type-species: Stenoclavelia mirablis Arnold. Orig. desig.
Chilochares Banks, 1941. Canad. Ent. 73: 119, 120.
Type-species: Cryptocheilus birkmanni Banks. Orig. desig.
Adirostes Banks, 1946. Mus. Compar. Zool., Bul. 96: 465.
Type-species: Adirostes tolteca Banks. Orig. desig.
attenuatum Banks. Tenn., La., Iowa, Kans., Tex., Colo.; Mexico (Morelos and Jalisco). Prey:
Lycosa antelucana Mont., L. avida Walck., L. near helluo Walck., L. sp.; all prey records
are of juvenile spiders.
Cryptocheilus attenuwatum Banks, 1933. Psyche 40: 8. d.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 76 (prey record). —Kurczewski and
Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 3 (prey records).
hesperus (Banks). Oreg., Calif., Nev., Utah.
Priocnemis hesperus Banks, 1915. Canad. Ent. 47: 401. °.
Cryptocheilus atratus Banks, 1919. Mus. Compar. Zool., Bul. 63: 247. d, 2.
idoneum birkmanni Banks. 100° W. to Rocky Mts. in U. Austr. and L. Austr. Zones; also
Ariz., N. Mex., and south. Calif.; Mexico (Baja California). Prey: Lycosa tigana Gertsch
and Wallace, adult.
Cryptocheilus birkmanni Banks, 1926. Canad. Ent. 58: 202 6, °@.

Biology: Evans, 1959. Kans. Ent. Soc., Jour. 32: 75 (prey record).
idoneum idoneum Banks. Va., N. C., Ga., Fla., Minn. Prey: Lycosa impavida Walck., juvenile.
Psammochares tenuicornis Banks, 1910. Psyche 17: 249. ¢. Preoce.

1530 Hymenoptera in America North of Mexico

Cryptocheilus idoneus Banks, 1910. Psyche 17: 250. 9.
Psammochares gracilicornis Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 225. N. name.

Biology: Kurezewski, 1963. Fla. Ent. 46: 209 (prey transport).
pallidipenne (Banks). Oreg., Calif., Ariz., and N. Mex. in L. Sonor. Zone.
Priocnemoides (!) pallidipennis Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 236. 2.
severini Banks. 100°W. to Rocky Mts. in U. Austr. and L. Austr. Zones; also N. Mex., Ariz.,
and south. Calif.; Mexico (Durango, Chihuahua, Nuevo Leon).
Cryptocheilus severini Banks, 1926. Canad. Ent. 58: 202. d.
Cryptocheilus arizonicus Banks, 1933. Psyche 40: 7. 2 (6 misdet.).
terminatum subopacum (Cresson). Atlantic to 100° W. in U. Austr. Zone.
Pompilus (Priocnemis) subopacus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 114. 6, °.
terminatum terminatum (Say). 100° W. to Rocky Mts. in U. Austr. Zone; Mexico (Zacatecas
and Teotihuacan). Prey: Lycosa sp.
Pompilus terminatus Say, 1828. American Entomology v. 3, p. 92.
Cryptocheilus carinatus Banks, 1920. Canad. Ent. 58: 202. 3.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 480 (prey, nest).
Genus PRIOCNEMIS Schiodte

So far as known all species are ground-nesting. Many species apparently do not dig a burrow
from the ground surface, but construct a cell or cells off the side of a mammal burrow or a
pre-existing crevice in the ground.

Genus PRIOCNEMIS Subgenus SPHICTOSTETHUS Kohl

Sphictostethus Kohl, 1885. Zool.-Bot. Gesell. Wien, Verh. 34: 37, 47.
Type-species: Pompilus gravesii Haliday. Orig. desig.

Haploneura Kohl, 1885. Zool.-Bot. Gesell. Wien, Verh. 34: 37, 47. Preoce.
Type-species: Haploneura apogona Kohl. Orig. desig.

Haploneurion Kohl, 1885. Ent. Nachr. 11: 163. N. name.

Anapriocnemis Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 25-26.
Type-species: Pompilus flavipes Guerin. Orig. desig.

pretiosa Banks. Ariz., N. Mex. at 5,000 to 9,100 ft.; Mexico. Prey: Lycosa sp., juvenile.
Priocnemis pretiosa Banks, 1933. Psyche 40: 18. 3.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 83 (prey record).
Genus PRIOCNEMIS Subgenus PRIOCNEMISSUS Haupt

Priocnemissus Haupt, 1949. Beitr. Tax. Zool. 1: 75.
Type-species: Priocnemis coriaceus Dahlbom, Orig. desig. Mispelled coriarius in
orig. descr.

The North American species are all vernal.

minorata Banks. N.S., Que., Ont., U. S. east of 100th Meridian in Alleghanian and Carolinian
Faunas, and in Transition Fauna in Pacific Northwest. Ecology: Occurs in woods and
appears to dig its own burrow. Prey: Coras juvenilis (Keys.), C. sp., Wadotes calcaratus
(Keys.), W. hybridus (Em.), W. sp.; Dolomedes tenebrosus Hentz; Arctosa rubicunda
(Keys.), Lycosa gulosa Walek., L. sp., Trochosa pratensis (Em.); Agroeca ornata Bks.,
Clubiona canadensis Em., C. obesa Hentz., C: spiralis Em.; Aysha gracilis (Hentz);
Amaurobius bennetti (Blackw.).

Priocnemis minorata Banks, 1912. Canad. Ent. 44: 197. °.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 434, figs. 8-13, 44 (larva).

Biology: Yoshimoto, 1954. Brooklyn Ent. Soc., Bul. 49: 130-138, 2 figs. (prey, nesting
behavior). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 99 (prey, nest).
—Kurezewski, 1963 (1962). Brooklyn Ent. Soc., Bul. 57: 86-87 (prey transport, nest).
—Kurczewski and Kurczewski, 1972. Kans. Ent. Soc., Jour. 45: 182-184 (prey records).

Superfamily POMPILOIDEA 1531

nigripes (Cresson). N. C., Tenn., Ark., Mo., Nebr., Kans.
Pompilus (Priocnemis) nigripes Cresson, 1865. Ent. Soc. Phila., Proe. 4: 454. @.
Priocnemis gomelza Brimley, 1934. Ent. News 45: 43. @.
oregona Banks. B. C., Wash., Oreg., Calif., Idaho, Nev., Utah, Ariz. Prey: Brachythele sp.;
Actinoxia versicolor Sim.; Atypoides riversi P.-Camb.
Pompilus comparatus Walker, 1866. Jn Lord, Naturalist in Vancouver Isl. and B. C., v. 2,
p. 341. 2. Preoce.
Priocnemis oregona Banks, 1933. Psyche 40: 11. N. name.

Biology: Hurd and Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey record). —Wasbauer
and Powell, 1962. Kans. Ent. Soc., Jour. 35: 394 (prey records).

Genus PRIOCNEMIS Subgenus PRIOCNEMIS Schiodte

Priocnemis Schiodte, 1837. Kroyer’s Naturhist. Tidsskr. 1: 324.
Type-species: Sphex exaltata Fabricius. Desig. by Westwood, 1840.
Prionocnemus Burmeister, 1872. Stettin. Ent. Ztg. 33: 235. Emend.
Prionocnemis Kirby, 1884. Zool. Rec. 20, Ins., p. 131. Emend. Preoce.
Myrmecosalius Ashmead, 1903. Ent. Soc. Wash., Proce. 5: 307.
Type-species: Myrmecosalius nigriceps Ashmead. Monotypic.
abbreviata Townes. Tex. (Ft. Davis).
Priocnemis (Priocnemis) abbreviatus Townes, 1957. U. S. Natl. Mus., Bul. 209: 102. 2.

aequalis (Banks). Transcont. in Canad. Zone and cooler parts of Transit. Zone.
Ageniella aequalis Banks, 1919. Mus. Compar. Zool., Bul. 63: 248. d.

cornica (Say). N. B., Ont., Man., transcontinental in U. S.; Mexico (Puebla, Durango). Ecology:

Occurs in open country, usually in sandy soil, but also in heavier soil. Prey: Drassylus
rufulus Bks.; Clubiona abbotti Koch, C. tibialis Em., C. sp., Trachelas tranquillus
(Hentz); Aysha gracilis (Hentz), A. sp.; Oxyopes salticus Hentz; Habronattus decorus
Black., Evarcha hoyi (Peckh.), Zygoballus nervosus Peckh., Pellenes viridipes (Hentz),
P. borealis (Bks.), P. spp., Salticus scenicus (Clerck); Arctosa littoralis Hentz, Lycosa
avida Walck., L. helluo Walck., L. spp., Pardosa milvina Hentz, P. moesta Bks., P.
saxatilis (Hentz), P. groenlandica (Thor.), P. spp., Pirata arenicola Em., P. sedentaria
Montg., P. sp., Trochosa avara Keys., Sosippus floridanus Sim., Lycosidae Sp.

Pompilus (Miscus) cornicus Say, 1836. Boston Jour. Nat. Hist. 1: 305. 2 (6 misdet.?).

Pompilus (Miscus) conicus Leconte, 1859. Writings of Thomas Say on Entomology, v. 2, p.
746. Emend.

Pompilus (Priocnemis) pomilius Cresson, 1867. Amer. Ent. Soc., Trans. 1: 116. 2.

Agenia atrata Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 263. d, ?.

Salius pompilus (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 237.

Priocnemis pompilus (!) Banks, 1919. Mus. Compar. Zool., Bul. 68: 245.

Ageniella eximia Banks, 1919. Canad. Ent. 51: 83. 6.

Ageniella aludra Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 201. d.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 434, fig. 38 (larva). —Townes, 1963. Ent.
Soc. Wash., Proc. 65: 115 (synonymy of atrata).

Biology: Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 53-55 (prey capture). —Rau
and Rau, 1918. Wasp studies afield, pp. 71-77, fig 14 (prey, nest). —Evans and Yoshimoto,
1955. Kans. Ent. Soc., Jour. 28: 17 (prey). —Townes, 1959. U. S. Natl. Mus., Bul. 209:
200-201 (prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 96-97 (prey,
nesting behavior). —Kurezewski, 1961. Brooklyn Ent. Soc., Bul. 56: 23 (prey).
—Kurezewski, 1963. Fla. Ent. 46: 210 (prey transport). —Kurezewski, 1963. Brooklyn Ent.
Soc., Bul. 57: 87-88 (prey, nest). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour.
41: 3-4 (prey records). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 368
(prey records). —Kurezewski and Kurezewski, 1972. Kans. Ent. Soc., Jour. 45: 184-185
(prey records).

germana (Cresson). Atlantic to 100° W. in Transit. and U. Austr. Zones, Colo., Ariz. Ecology:
Occurs in woods. Prey: Agelenopsis utahana (Chamb. and Ivie), Coras sp., prob.

1532 Hymenoptera in America North of Mexico

juvenilis (Keys.), Wadotes hybridus (Em.), W. sp.; Clubiona spiralis Em.; Aysha gracilis
(Hentz); Maevia vittata (Hentz); Amaurobius bennetti (Blackw.), A. sp.
Pompilus (Priocnemis) germanus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 116. °.
Pompilus (Agenia) iridipennis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 127. ¢.

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 98 (hunting behavior,
prey transport, nest ?). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 4
(prey records). —Kurezewski and Kurczewski, 1972. Kans. Ent. Soc., Jour. 45: 185-186
(prey records).

hestia (Banks). N. H. to Va., W. Va. Ecology: Occurs in woods. Prey: Agroeca sp.
Ageniella hestia Banks, 1915. Canad. Ent. 47: 400. 6.
Ageniella crassicornis Banks, 1917. Mus. Compar. Zool., Bul. 61: 108. 3.

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 98 (prey hunting,
transport, nest ?).
minuscula (Banks). N. C., 8. C., Tex., Kans., Ill.
Ageniella minuscula Banks, 1917. Mus. Compar. Zool., Bul. 61: 110. 3.
nigriceps (Ashmead). Iowa, Kans., Tex.
Myrmecosalius nigriceps Ashmead, 1903. Ent. Soe. Wash., Proc. 5: 308. ?.
notha alaskensis Townes. Alaska, N. W. T. Prey: Paraphidippus marginatus (Walck.).
Priocnemis (Priocnemis) notha alaskensis Townes, 1957. U. S. Natl. Mus., Bul. 209: 106.
Ones
Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 106 (prey).
notha navajo Banks. B. C., Oreg., Calif., Ariz., N. Mex.; Transit. and Canad. Zones; Mexico.
Priocnemis navajo Banks, 1933. Psyche 40: 15. 9.
notha notha (cresson). P. E. I., N. B., Que., Ont., Man., in U. S. from Atlantic to Rocky Mts. in
Canad., Transit., and U. Austr. Zones. Ecology: Occurs in open country. Prey: Trochosa
pratensis Em., Schizocosa crassipes (Walck.), S. sp., Clubiona sp.
Pompilus (Priocnemis) nothus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 118. 9.
Cryptocheilus paeneparcus Viereck, 1906. Amer. Ent. Soc., Trans. 32: 202. @.

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 97-98 (prey transport,
nest). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 4-5 (prey transport).
—Kurczewski and Kurczewski, 1972. Kans. Ent. Soc., Jour. 45: 186 (prey).

notha occidentis Banks. Oreg., Wash., Calif.; in Transit. and Canad. Zones.
Priocnemis occidentis Banks, 1944. Mus. Compar. Zool., Bul. 94: 172. 9.
scitula relicta Banks. Que., Ont. to N. C., W. Va., Ohio, Wis. Ecology: Occurs in bottomland
woods. Prey: Habrocestum pulex (Hentz), Maevia vittata (Hentz); Agelenopsis sp.;
Agroeca sp., Clubiona kastoni Gertsch, C. spiralis Em., C. spp.; Xysticus sp.;
Amaurobius bennetti (Blackw.).
Priocnemis relicta Banks, 1912. Canad. Ent. 44: 198. °.
Ageniella tenella Banks, 1915. Canad. Ent. 47: 400. ¢.

Biology: Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 62 (prey transport). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 98-99 (prey transport, nest). —Kurezewski
and Kurezewski, 1972. Kans. Ent. Soc., Jour. 45: 186-187 (prey).

scitula scitula (Cresson). Atlantic to 100° W. in U. Austr. Zone. Ecology: Bottomland woods.
Pompilus (Priocnemis) scitulus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 118. 9.
Agenia perfecta Provancher, 1882. Nat. Canad. 13: 44. ¢.

Taxonomy: Townes, 1963. Ent. Soc. Wash., Proc. 65: 115 (synonymy of perfecta).

Genus CALICURGUS Lepeletier,

Calicurgus Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 397.
Type-species: Pompilus fasciatellus Spinola. Desig. by Kohl, 1884 © Sphex
hyalinatus Fabricius).
Caliadurgus Pate, 1946. Amer. Ent. Soc., Trans. 72: 78.
Type-species: Sphex hyalinata Fabricius. Orig. desig.

Superfamily POMPILOIDEA 1533

hyalinatus alienatus (Smith). Atlantic to 100° W. in Transit. and U. Austr. Zones. Ecology:

Occurs along stream and river bottoms; nests in sand and heavier soil. Prey: Araneus
marmoreus Clerck., A. patagiatus Clerck., A. spp., Araniella displicata (Hentz),
Neoscona spp., Wixia ectypa (Walck.), Eustala anastera (Walck.), Acanthepeira stellata
(Walck.), Araneinae sp. C. hyalinata hyalinata (Fabricius) occurs in Europe.

Pompilus fascipennis Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app. p. 332. °.
Preoce.

Pompilus alienatus Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 159. N. name.

Pompilus (Agenia) calcaratus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 128. d.

Salius fasciipennis Dalla Torre, 1897. Cat. Hym., v. 8, p. 223. Emend.

Biology: Evans and Yoshimoto, 1955. Kans. Ent. Soc., Jour. 28: 17 (prey). —Townes, 1957. U.
S. Natl. Mus., Bul. 209: 113 (prey). —Krombein, 1958. Biol. Soc. Wash., Proc. 71: 21 (prey
transport). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 52-53 (prey). —Krombein, 1961.
Brooklyn Ent. Soc., Bul. 56: 62 (prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer.,
Misc. Pub. 3: 101 (prey transport, nest). —Kurezewski and Kurczewski, 1968. Kans. Ent.
Soc., Jour. 41: 5 (prey and transport). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soc.,
Jour. 41: 368 (prey and capture). —Kurezewski and Kurcezewski, 1972. Kans. Ent. Soc.,
Jour. 45: 187 (prey and transport).

hyalinatus borealis (Banks). Transcont. in Canad. Zone.
Priocnemis alienatus borealis Banks, 1933. Psyche 40: 10. °.
hyalinatus excoctus Townes. N. Mex, Ariz.; Mexico (Morelos, Durango, Jalisco).
Calicurgus hyalinatus excoctus Townes, 1957. U.S. Natl. Mus., Bul. 209: 115. ?, d.
hyalinatus rupex (Cresson). Md., N. C., Ga., Kans., Tex., mostly in Austroriparian Fauna;
Mexico (Puebla).
Pompilus (Priocnemis) rupex Cresson, 1869. Boston Soe. Nat. Hist., Proc. 12: 372. 9.
Pompilus (Agenia) accolens Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 374. ¢.
Salius ruspex (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 239.

Genus DIPOGON Fox

Our species are most commonly found in wooded areas. The North American species for which
the nesting habits are known all use pre-existing cavities in wood as a nesting site. The nest con-
sists of a linear series of cells usually separated by complex partitions of debris such as bits of
wood or leaf, dead insects, caterpillar frass with an outer layer of compacted soil. Errant spiders
are used as prey.

Revision: Evans, 1974. Amer. Ent. Soc., Trans. 100: 29-51, 23 figs. (spp. of southwestern U.
S., Mexico, Central America).

Genus DIPOGON Subgenus DEUTERAGENIA Sustera

Agenia Schiodte, 1837. Kroyer’s Naturhist. Tidsskr. 1: 821. Preoce.
Type-species: Sphex variegata Linnaeus. Desig. by Westwood, 1840.
Pogonius Dahlbom. 1845. Hym. Europaea, v. 1, p. 453. Preoce.
Type-species: Sphea variegata Linnaeus. Desig. by Pate, 1946.
Deuteragenia Sustera, 1913. Zool.-Bot. Gesell. Wien, Verh. 16: 191. N. name for Agenia.
Dipogon subg. Adipogon Banks, 1944. Mus. Compar. Zool., Bul. 94: 181.
Type-species: Pompilus pulchripennis Cresson. Orig. desig.
calipterus calipterus (Say). Mass. to N. C., Ind., Ill. Prey: Amaurobius sp.
Pompilus calipterus Say, 1836. Boston Jour. Nat. Hist. 1: 302. °.
Deuteragenia pilosa Banks, 1933. Psyche 40: 16. °.
Dipogon femur-aureus Dreisbach, 1953. Amer. Midland Nat. 49: 832. °.

Biology: Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 394 (prey).

calipterus duplicatus Townes. Fla., Ga.
Dipogon (Deuteragenia) calipterus duplicatus Townes, 1957. U. S. Natl. Mus., Bul. 209:

125. 9.

1534 Hymenoptera in America North of Mexico

calipterus nubifer (Cresson). Calif. s. to Panama. Ecology: Nests in hollow stems. Prey:
Gnaphosa sp.; Misumena sp.; Trachelas pacificus Chamb. and Ivie.
Pompilus (Agenia) nubifer Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 374. 9.
Pseudagenia isthmica Cameron, 1891. Biol. Cent.-Amer., Hym. 2: 165, pl. 10, fig. 11. @.
Pseudagenia isthmia(!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 204.

Taxonomy: Wasbauer, 1960. Pan-Pacifie Ent. 36: 174, figs. 4-6. ¢. —Evans, 1974. Amer. Ent.
Soc., Trans. 100: 36, fig. 19. 2, 3.

Biology: Williams, 1966. Amer. Midland Nat. 24: 33-47 (prey, nest, life history). —Parker and
Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest).
geronimo Evans. Southeastern Ariz.
Dipogon (Deuteragenia) geronimo Evans, 1974. Amer. Ent. Soce., Trans. 100: 37, figs. 4, 7,
pel Seon Gr
hurdi Evans. Southeastern Ariz.; Mexico (Durango). Prey: Iciws sp.
Dipogon (Deuteragenia) hurdi Evans, 1974. Amer. Ent. Soc., Trans. 100: 41, figs. 5, 21. @,
3.

Biology: Evans, 1974. Amer. Ent. Soc., Trans. 100: 41 (prey).

iracundus Townes. Ariz.; Mexico (Durango). Ecology: Nests in borings in wood.
Dipogon (Deuteragenia) iracundus Townes, 1957. U.S. Natl. Mus., Bul. 209: 127. 2.

Taxonomy: Evans, 1974. Amer. Ent. Soc., Trans. 100: 40-41. 2, 6.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 168-169 (nest, life history).
melanocephalus (Cameron). Tex. (Hidalgo Co.); Mexico (Veracruz, Oaxaca, Morelos).
Pseudagenia melanocephala Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 172, pl. 10,
fig 18) 9%

Taxonomy: Evans, 1974. Amer. Ent. Soc., Trans. 100: 43-45, figs. 3, 8, 15, 22. 2, 3.
papago anomalus Dreisbach. Canad., Transit. and U. Austr. Zone from Atlantic to 100° W.
Ecology: Nests in borings in wood. Prey: Paraphidippus aurantius. (Luc.), Phidippus
sp.; Sergiolus variegatus (Hentz), Haplodrassus hiemalis (Em.), Poecilochroa capulata
(Walck.); Clubiona canadensis Em.
Dipogon anomalus Dreisbach, 1953. Amer. Midland Nat. 49: 834, figs. 4, 8. ¢.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 436 (larva).

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 122 (prey, nest). —Evans and Yoshimoto,
1962. Ent. Soc. Amer., Mise. Pub. 3: 104-105 (prey transport, nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 167-168 (prey, nest, life history). —Kurezewski and
Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 368 (prey).

papago floridanus Townes. Fla.
Dipogon (Deuteragenia) papago floridanus Townes, 1957. U. 8. Natl. Mus., Bul. 209: 122.
25 6)
papago papago (Banks). B. C., Ariz., Tex.; Mexico (Morelos).
Deuteragenia papago Banks, 1943. Psyche 40: 17. °.
pulchripennis (Cresson). Atlantic to 100° W., mostly in Canad. and Transit. Zones; also Ariz.
(9,000 ft. in Santa Catalina Mts.); Mexico (Durango). Prey: Xysticus sp.; Amaurobius
bennetti (Blackw.); Phidippus audax (Hentz).
Pompilus (Agenia) pulchripennis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 123. 6, @.

Biology: Kurezewski and Kurezewski, 1972. Kans. Ent. Soc., Jour. 45: 188 (prey).
sayi nigrior Townes. Transit. Fauna, B. C., Oreg., Calif., Mont., Colo., N. Mex. Prey: Xysticus

sp.
Dipogon (Deuteragenia) sayi nigrior Townes, 1957. U.S. Natl. Mus., Bul. 209: 130. °.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 131 (prey).
sayi sayi Banks. Atlantic to 100° W. in Transit. and U. Austr. Zones. Ecology: Nests in
borings in wood. Parasite: Anthrax irroratus Say; Melittobia chalybii Ashm.,
Tetrastichus johnsoni Ashm.; Ephuta p. pauxilla Brad. Prey: Xysticus bicuspis Keys.,

Superfamily POMPILOIDEA 1535

X. elegans Keys., X. ferox (Hentz), X. pellax Camb., X. triguttatus Keys., X. fraternus
Bks., X. funestus Keys., X. canadensis Gertsch, X. discursans Keys., X. obscurans Coll.,
X. punctatus (Keys.), X. lutulentus Gertsch, X. spp., Miswmena vatia (Clerck),
Misumenoides formosipes (Walck.), Tmarus angulatus (Walck.), Coriarachne versicolor
Keys., C. utahensis Gertsch; Agelenopsis utahana (Chamb. and Ivie); Phidippus
whitmanii Peckh.; Nodocion melanie Levi, Poecilochroa capulata (Walck.), P. montana
Em.; Amaurobius bennetti(Blackw.), A. sp.

Dipogon sayi Banks, 1941. Canad. Ent. 73: 122. d, 9.

Deuteragenia fascipennis Haupt, 1959. Nova Acta Leopoldina 21, no 141: 32, fig. 16. 2.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 436, figs. 14-19, 45 (larva). —Evans, 1974.
Amer. Ent. Soc., Trans. 100: 30 (synonymy).

Biology: Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 144 (prey, nest; misdet. as
calipterus). —Townes, 1957. U.S. Natl. Mus., Bul. 209: 130 (nest). —Medler and Koerber,
1957. Ent. Soc. Amer., Ann. 50: 621-625, 6 figs. (nest, prey, life history, parasite).
—Krombein, 1958. Ent. Soe. Wash., Proc. 60: 52 (prey). —Evans and Yoshimoto, 1962. Ent.
Soc. Amer., Misc. Pub. 3: 102-104 (nest, prey, life history, parasites). —Fye, 1965. Canad.
Ent. 97: 735-736 (nest, prey, life history). —Krombein, 1967. Trap-nesting wasps and bees,
pp. 161-167, pl. 10, figs. 46 (?), 47 (2), 48 (nest, prey, life history, parasites). —Kurczewski
and Kurezewski, 1972. Kans. Ent. Soc., Jour. 45: 188 (prey).

sericeus Banks. Oreg., Calif.
Dipogon sericea Banks, 1944. Mus. Compar. Zool., Bul. 94: 180. 2.
thoracicus Townes. N. Mex., Ariz.; Mexico (Sinaloa, Durango, Morelos). Prey: Xysticus spp.,
juv.
Dipogon (Deuteragenia) thoracicus Townes, 1957. U. S. Natl. Mus., Bul. 209: 126. ?.

Biology: Evans, 1974. Amer. Ent. Soc., Trans. 100: 40 (prey).

Genus DIPOGON Subgenus DIPOGON Fox

Dipogon Fox, 1897. Acad. Nat. Sci. Phila., Proc., p. 241.
Type-species: Dipogon populator Fox. Orig. desig.
Agriogenia Banks, 1919. Canad. Ent. 51: 83.
Type-species: Pompilus brevis Cresson. Orig. desig.
brevis brevis (Cresson). Carol. Fauna, Mass. to N. C. Prey: Phidippus purpuratus Keys.,
Pellenes hoyi (Peckh.), P. sp.
Pompilus (Agenia) brevis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 123. 6.

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 105 (prey transport).
—Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 5 (prey).
brevis ochreus Townes. Md., S. C. Prey: Phidippus (?) sp., juv.
Dipogon (Dipogon) brevis ochreus Townes, 1957. U.S. Natl. Mus., Bul. 209: 137. 9.
Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 137 (prey).
brevis recalvus Townes. Alleghan. Fauna, Que. and Ont. to Va. and Wis.
Dipogon (Dipogon) brevis recalvus Townes, 1957. U. S. Natl. Mus., Bul. 209: 134. 2, 6.
diablo Wasbauer. Calif. (Contra Costa Co.).
Dipogon (Dipogon) diablo Wasbauer, 1960. Pan-Pacific Ent. 36: 172, figs. 1-3. 2, d.
graenicheri atratus Townes. N. J., N. C.
Dipogon (Dipogon) graenicheri atratus Townes, 1957. U. S. Natl. Mus., Bul. 209: 139, pl. 2,
fig Zoe:
graenicheri graenicheri Banks. Fla., La.
Dipogon graenicheri Banks, 1939. Canad. Ent. 71: 230. °.
leechi Wasbauer. Calif. (Marin Co.); Mexico (Baja California).
Dipogon (Dipogon) leechi Wasbauer, 1960. Pan-Pacifie Ent. SOs iene
paludis Townes. N. J. (Atsion).
Dipogon (Dipogon) paludis Townes, 1957. U. S. Natl. Mus., Bul. 209: 133. 2.
parkeri Wasbauer. Nev. (Washoe Co.). Ecology: Nests in borings in Sambucus stems.
Dipogon (Dipogon) parkeri Wasbauer, 1966. Biol. Soe. Wash., Proc. 79: 17. 2, d.

15386 Hymenoptera in America North of Mexico

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest).

pygmaeus Townes. Ala. (Prattsville).
Dipogon (Dipogon) pygmaeus Townes, 1957. U.S. Natl. Mus., Bul. 209: 132. 9.
texanus texanus Banks. Tex. (Brownsville). Another subspecies occurs in Mexico.
Dipogon texanus Banks, 1944. Mus. Compar. Zool., Bul. 94: 179. °.

Genus DIPOGON Subgenus WINNEMANELLA Krombein

Dipogon subg. Winnemanella Krombein, 1962. Biol. Soc. Wash., Proc. 75: 7.
Type-species: Dipogon (Winnemanella) fulleri Krombein. Orig. desig.
fulleri Krombein. Md., S. C. Prey: Icius hartii Em.
Dipogon (Winnemanella) fulleri Krombein, 1962. Biol. Soc. Wash., Proce. 75: 8. 2.

Biology: Krombein, 1962. Biol. Soc. Wash., Proc. 75: 8 (prey).
TRIBE AUPLOPODINI

Members of this tribe usually amputate one or more of the spider’s legs, presumably to
facilitate prey transport, although there are records of some wasps feeding on body fluids exud-
ing from the severed leg stumps. Phanagenia and Auplopus are mud-daubers, constructing mud
cells in sheltered situations. A few species of Ageniella have been recorded as nesting in the
ground, either in pre-existing crevices or in burrows which they dig themselves. The nesting
habits of Priocnemella are unknown, but the lack of a pygidial area in the female suggests that
it is not a mud-dauber.

Genus PHANAGENIA Banks

Phanagenia Banks, 1933. Psyche 40: 18.
Type-species: Phanagenia osceola Banks. Orig. desig. — Pompilus bombycinus
Cresson).

Only the type-species occurs in the New World. At least two other species occur in Africa.

bombycina (Cresson). Atlantic to 100° W. in U. Austr. and L. Austr. Zones, also N. Mex.
Ecology: Builds mud cells beneath loose bark and under stones. Parasite: Ceropales
robinsonii Cr.; Ephuta scrupea (Say ). Prey: Lycosa avida Walck., L. gulosa Walck.,
Lycosidae sp.; Maevia vittata (Hentz); Agelenopsis sp.
Pompilus (Agenia) bombycinus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 125. d, °.
Ageniella annecta Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 233. 9.
Phanagenia osceola Banks, 1933. Psyche 40: 18. °.

Biology: Walsh and Riley, 1869. Amer. Ent. 1: 131-132, 136, 163 (nest, parasite). —Peckham
and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 164-165 (nest, prey, life history).
—Peckham and Peckham, 1905. Wasps social and solitary, pp. 244-247 (nest, life history,
prey). —Savin, 1924. Nat. Hist. 24: 520-522 (nest, prey). —Schuster, 1951. N. Y. Ent. Soc.,
Jour. 59: 34 (parasite). —Townes, 1957. U.S. Natl. Mus., Bul. 209: 143 (nest, prey).
—Kurezewski, 1961. Brooklyn Ent. Soc., Bul. 56: 23 (prey transport). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 108 (prey transport). —Kurezewski and
Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 5 (prey).

Genus AUPLOPUS Spinola

Auplopus Spinola, 1841. Soe. Ent. France, Ann. 10: 108.
Type-species: Pompilus femoratus Fabricius. Monotypic.
Aoplopus Agassiz, 1846. Nomencl. Zool., Index Univ., pp. 27, 41. Emend.
Pilpomus Costa, 1859. Fauna Regno Napoli, Imenotteri Aculeati, Pompilidea, p. 3.
Type-species: Sphex carbonarius Scopoli. Desig. by Pate, 1946.
Pseudagenia Kohl, 1885. Zool.-Bot. Gesell. Wien, Verh. 34: 38, 42.
Type-species: Sphex carbonarius Scopoli. Orig. desig.
Tumagenia Banks, 1934. Amer. Acad. Arts and Sci., Proc. 69: 39, 67.
Type-species: Tumagenia iris Banks. Monotypice.
Calagenia Banks, 1934. Amer. Acad. Arts and Sci., Proc. 69: 40, 72.

Superfamily POMPILOIDEA 1537

Type-species: Calagenia hermosa Banks. Orig. desig.
Lophagenia Banks, 1934. Amer. Acad. Arts and Sci., Proe. 69: 40, 74.
Type-species: Pseudagenia erigone Bingham. Orig. desig.

The genera Stenagenia Saussure, 1892, and Schizagenia Cameron, 1910, are questionable
synonyms.

Members of this genus make mud cell nests under stones, under logs, and in other protected
places.

adjunctus (Banks). Md. to Fla., west to Tex. Ecology: Reared from mud cell; occurs in damp
bottomland woods. Prey: Chiracanthium inclusum (Hentz); Phidippus rimator (Walck.).
Pseudagenia mellipes var. adjuncta Banks, 1911. Ent. Soc. Wash., Proc. 13: 238.
Pseudagenia marionae Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 48: 202. d.

Taxonomy: Banks, 1912 (1911). N. Y. Ent. Soe. Jour. 19: 233. @.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 151 (nest). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 6 (prey).
architectus architectus (Say). Transit. to Austrorip. Zones from Atlantic to Rocky Mts., and

west of Rocky Mts. in Colo., N. Mex. and Ariz.; Mexico (Durango, Zacatecas, Puebla).
Ecology: Builds mud cells under loose bark, under stones, and in abandoned burrows of
ground-nesting bees. Parasite: Sphaeropthalma pensylvanica scaeva (Bl.). Prey:
Trachelas tranquillus (Hentz), Clubiona sp.; Phidippus princeps (Peckh.), P. audax
(Hentz), P. sp. near whitmannii Peckh., Icius similis Bks.; Misumenops oblongus
(Keys.).

Pompilus architectus Say, 1836. Boston Jour. Nat. Hist. 1: 303. 2.

Taxonomy: Walsh, 1869. Amer. Ent. 1: 163.

Biology: Say, 1836. Boston Jour. Nat. Hist. 1: 303 (nest). —Walsh and Riley, 1869. Amer.
Ent. 1: 132, 163 (nest). —Rau and Rau, 1918. Wasp studies afield, pp. 83-84, fig. 16 (nest).
—Krombein, 1955. Brooklyn Ent. Soe., Bul. 50: 14-15 (prey transport). —Townes, 1957. U.
S. Natl. Mus., Bul. 209: 164-165 (nest, prey). —Kurezewski, 1961. Brooklyn Ent. Soe., Bul.
56: 24 (prey transport). —Krombein, 1961. Brooklyn Ent. Soe., Bul. 56: 62-63 (prey).
—Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 108-109 (nest, prey).
—Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 6 (prey transport).

architectus metallicus (Banks). West of Rocky Mts., from B. C. to north Mexico. Parasite:
Sphaeropthalma difficilis (Bak.). Prey: Thiodina sp., Phidippus formosus Peckh. and
Peckh.; Trachelas pacificus Chamb. and Ivie.
Pseudagenia metallica Banks, 1910. N. Y. Ent. Soe., Jour. 18: 125. ?.

Biology: Hurd and Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 109 (prey).
caerulescens caerulescens (Dahlbom). Md. to S. C., Kans., Tex.
Agenia caerulescens Dahlbom, 1843. Hym. Europaea, v. 1, p. 93. °.
Pseudagenia coerulescens (!) Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 2382. 6, @.
caerulescens floridanus (Banks). Austrorip. Fauna, Fla. to S. C.
Pseudagenia floridana Banks, 1921. Ent. Soc. Amer., Ann. 14: 21. °.
caerulescens subcorticalis (Walsh). Atlantic to 100° W. in Transit., U. Austr., and L. Austr.
Zones, though replaced in parts of its range by the subspecies antennalis and
caerulescens; Mexico (Guayamas). Ecology: Makes mud cells beneath loose bark and in
borings in wood; occurs in woods. Prey: Trachelas tranquillus (Hentz), Clubiona obesa
Hentz, C. sp.; Phidippus audax (Hentz); Anyphaena pectorosa Koch.
Agenia subcorticalis Walsh, 1869. Amer. Ent. 1: 162. d, °.
Pseudagenia antennalis Banks, 1910. Psyche 17: 251. °.
Pseudagenia ariella Banks, 1941. Canad. Ent. 73: 122. °.

Biology: Walsh and Riley, 1869. Amer. Ent. 1: 131-182 (nest). —Evans and Yoshimoto, 1955.
Kans. Ent. Soc., Jour. 28: 17 (prey). —Townes, 1957. U.S. Natl. Mus., Bul. 209: 159 (nest).
—Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 109-110 (prey transport).
—Medler, 1964. Ent. News 75: 190-191 (nest, prey, life history). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 170-171, pl. 9, figs. 42-44 (nest, prey, life history).

1538 Hymenoptera in America North of Mexico

flavicoxae (Banks). Ariz. (Palmerlee).
Pseudagenia mexicana var. flavicoxae Banks, 1911. Ent. Soe. Wash., Proc. 13: 238. 3, °.
Pseudagenia mexicana var. flavicora(!) Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 233.
inermis Townes. Tex.; Mexico (Morelos). Prey: Lycosa sp.
Auplopus inermis Townes, 1957. U.S. Natl. Mus., Bul. 209: 148. °.

Biology: Evans, 1964. Kans. Ent. Soc., Jour. 37: 306 (prey transport).
mellipes mellipes (Say). N. J. to Fla., west to Kans. and Tex. Ecology: Builds mud cells in cells

of abandoned nests of Polistes, Sceliphron, and Trypargilum, in borings in wood,
beneath loose bark, under exposed roots, and in abandoned borings of ground-nesting
bees; occurs in woods. Parasite: Sphaeropthalma pensylvanica scaeva (Bl.). Prey:
Pisaurina undata Hentz, P. mira (Walck.); Herpyllus vasifer (Walck.); Philodromus sp.;
Marpissa undata (DeG.), Phidippus audax (Hentz), P. sp.

Pompilus mellipes Say, 1836. Boston Jour. Nat. Hist. 1: 304. 9.

Agenia fulvipes Dahlbom, 1843. Hym. Europaea, v. 1, p. 92. 9.

Pseudagenia mellipes var. interior Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 233. ¢.

Biology: Walsh, 1869. Amer. Ent. 1: 132 (nest). —Rau and Rau, 1916. Jour. Anim. Behavior 6:
42-43 (nest). —Rau and Rau, 1918. Wasp studies afield, pp. 86-89, figs. 18-20 (nest). —Rau,
1926. St. Louis Acad. Sci., Trans. 25: 196-197 (nest, prey). —Rau, 1928. St. Louis Acad. Sci.,
Trans. 25: 342-358 (nest, prey). —Krombein, 1952. Ent. Soc. Wash., Proc. 54: 176-177
(nesting behavior, prey). —Krombein, 1955. Brooklyn Ent. Soc., Bul. 50: 15 (prey).
—Townes, 1957. U. S. Natl. Mus., Bul. 209: 153 (nest). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 171-178, pl. 9, fig. 45 (nest, life history).

mellipes meridianus Townes. Fla. (Paradise Key).
Auplopus mellipes meridianus Townes, 1957. U. S. Natl. Mus., Bul. 209: 154. 2.
mellipes variitarsatus (Dalla Torre). Atlantic to 100° W. in Canad. and Transit. Zones.
Ecology: Makes mud nests in and under decaying logs and stumps, and in borings in
wood; occurs in woods.
Pompilus (Agenia) varipes Cresson, 1867. Amer. Ent. Soc., Trans. 1: 126. 2. Preoce.
Agenia variitarsata Dalla Torre, 1897. Cat. Hym., v. 8, p. 210. N. name.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 155 (nest). —Evans and Yoshimoto, 1962.
Ent. Soe. Amer., Mise. Pub. 3: 110 (nest). —Medler, 1964. Ent. News 75: 191 (nest).
mexicanus (Cresson). South. Ariz.; Mexico (Veracruz).
Pompilus (Agenia) mexicanus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 130. °.
mollis Townes. South. Tex.
Auplopus mollis Townes, 1957. U. S. Natl. Mus., Bul. 209: 157. 2.
nigrellus (Banks). Most of south. Canad. and U. S., but not known from northwest except
Oreg., or from southeast; Mexico (Baja California). Ecology: Occurs in or at edge of
woods; builds mud cells beneath stones and in boring in stem. Prey: Clubiona abboti
Koch, Trachelas tranquillus (Hentz), T. sp.; Phidippus spp., Maevia inclemens (Walck.);
Anyphaena fraterna Bks., Aysha gracilis (Hentz).
Pseudagenia nigrella Banks, 1912 (1911). N. Y. Ent. Soe. Jour. 19: 232. 2.
Pseudagenia nanella Banks, 1912. Canad. Ent. 44: 198. °.

Biology: Krombein, 1954. Brooklyn Ent. Soc., Bul. 49: 4 (prey). —Krombein, 1955. Brooklyn
Ent. Soc., Bul. 50: 15 (prey). —Townes, 1957. U. S. Natl. Mus., Bul. 209: 167 (nest).
—Evans and Yoshimoto, 1962. Ent. Soe. Amer., Mise. Pub. 3: 109 (prey transport).
—Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest). —Kurezewski and Kurczewski,
1968. Kans. Ent. Soc., Jour. 41: 6 (prey transport). —Kurezewski and Kurczewski, 1968.
Kans. Ent. Soc., Jour. 41: 369 (prey).

variolarum Townes. Tex. (Chisos Mts.).
Auplopus variolarum Townes, 1957. U. S. Natl. Mus., Bul. 209: 149. °.

Genus AGENIELLA Banks

Genus AGENIELLA Subgenus LEUCOPHRUS Townes
Ageniella subg. Leucophrus Townes, 1951. U.S. Dept. Agr., Monog. 2: 917.

Superfamily POMPILOIDEA 1529

Type-species: Priocnemis semitincta Banks. Orig. desig.
fulgifrons (Cresson). U. Austr. and L. Austr. Zones from Atlantic to 100° W. Prey: Phidippus
audax Hentz, P. sp., Paraphidippus sp. near marginatus (Walck.).
Pompilus (Priocnemis) fulgifrons Cresson, 1867. Amer. Ent. Soc., Trans. 1: 114. °.
Pompilus (Agenia) agilis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 125. d.
Salius fulgidifrons Dalla Torre, 1897. Cat. Hym., v. 8, p. 225. Emend.

Biology: Evans, 1959. Kans. Ent. Soe., Jour. 32: 75 (prey). —Kurezewski and Kurcezewski,

1968. Kans. Ent. Soc., Jour. 41: 6-7 (prey).

incita (Banks). Kans., Tex.
Cryptocheilus incitus Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 234. 9.

reynoldsi (Banks). Colo., Kans., Tex.
Priocnemis reynoldsi Banks, 1933. Psyche 40: 12. 9.

semitincta (Banks). Transcont. in U. Austr. and L. Austr. Zones. Prey: Agelenopsis potteri

Blackw., A. pennsylvanica (Koch), A. spp.

Priocnemis semitincta Banks, 1912. Canad. Ent. 44: 197. °.
Ageniella festina Banks, 1917. Mus. Compar. Zool., Bul. 61: 109. d.
Ageniella fraternella Banks, 1917. Mus. Compar. Zool., Bul. 61:.109. ¢.
Priophanes holonis Banks, 1944. Mus. Compar. Zool., Bul. 94: 174. °.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 171 (prey). —Kurezewski, 1961. Brooklyn
Ent. Soc., Bul. 56: 24 (prey transport). —Kurezewski and Kurcezewski, 1968. Kans. Ent.

Soc., Jour. 41: 7 (prey).
Genus AGENIELLA Subgenus NEMAGENIA Banks

Ageniella subg. Nemagenia Banks, 1944. Mus. Compar. Zool., Bul. 94: 179.
Type-species: Pompilus (Agenia) longulus Cresson. Orig. desig.
longula (Cresson). Calif., “Dakota”, Kans., Mo., La., Tex.; British Guiana; Bolivia. Prey:
Pardosa sp., Lycosa mccooki Montgomery, L. sp.
Pompilus (Agenia) longulus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 129. 6.
Agenia longa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 205. “2” = ¢.
Priocnemis directa Banks, 1912. Canad. Ent. 44: 197. 9.
Priophanes otiosa Banks, 1946. Mus. Compar. Zool., Bul. 96: 442. 2.

Biology: Townes, 1951. U. S. Dept. Agr., Monog. 2: 918 (prey). —Evans, 1959. Kans. Ent.
Soc., Jour. 32: 75 (prey).

Genus AGENIELLA Subgenus PRIOPHANES Banks

Priophanes Banks, 1944. Psyche 50: 82.
Type-species: Priocnemis facetus Cresson. Orig. desig.
agenioides (Fox). Atlantic to 100° W. in U. Austr. and L. Austr. Zones. Prey: Xysticus sp.;
Maevia inclemens (Walck.), M. vittata (Hentz), Salticidae sp.
Priocnemis agenioides Fox, 1893. N. Y. Ent. Soc., Jour. 1: 54. 2.
Pseudagenia virginica Banks, 1910. Psyche 17: 251. 6.
Ageniella subra Brimley, 1934. Ent. News 45: 42. ¢.
Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 187 (prey). —Kurezewski and Kurczewski,
1968. Kans. Ent. Soc., Jour. 41: 8 (prey transport).
arcuata (Banks). Md. to Fla., Alta., Colo., Kans., Tex. to south. Calif.; Mexico (Nayarit,
Durango, Zacatecas, Teotihuacan, Veracruz, Jalisco). Prey: Oxyopes salticus Hentz;

Misnmenops celer Hentz, Sassacus papenhoei Peckh.
Cryptocheilus arcuatus Banks, 1910. N. Y. Ent. Soe., Jour. 18: 120. °.
Pseudagenia birkmanni Banks, 1910. N. Y. Ent. Soe., Jour. 18: 124. d.

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 107 (prey transport).
—Evans, 1964. Kans. Ent. Soe., Jour. 37: 306 (prey transport).

arizonica arizonica (Banks). Ariz. (Tempe).
Priocnemis arizonica Banks, 1933. Psyche 40: 14. 2.

1540 Hymenoptera in America North of Mexico

arizonica concolor Townes. Kans. (Manhattan).
Ageniella (Priophanes) arizonica concolor Townes, 1957. U. S. Natl. Mus., Bul. 209: 179. 2,
Ge

carolae Wasbauer. Calif. (Riverside Co.). On flowers of Asclepias erosa.

Ageniella (Priophanes) carolae Wasbauer, 1959. Kans. Ent. Soc., Jour. 32: 109, fig. 4. 2.
faceta faceta (Cresson). Md. to Ga., Tex. Prey: Pardosa sp., Schizocosa sp.

Priocnemis facetus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 205. 9.

Cryptocheilus pallescens Banks, 1910. N. Y. Ent. Soe., Jour. 18: 121. 9.

Biology: Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 8 (prey).

faceta ventralis Townes. Fla.; Mexico (Nuevo Leon).
Ageniella (Priophanes) faceta ventralis Townes, 1957. U. S. Natl. Mus., Bul. 209: 183. ?,
é.

fuscipennis Townes. Calif. Prey: Oxyopes salticus Hentz; Xysticus sp.
Ageniella (Priophanes) fuscipennis Townes, 1957. U.S. Natl. Mus., Bul. 209: 189. 9.

Taxonomy: Wasbauer, 1959. Kans. Ent. Soc., Jour. 32: 109-111, figs. 1-3. ¢.

Biology: Hurd and Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey). —Wasbauer and
Powell, 1962. Kans. Ent. Soc., Jour. 35: 395 (prey).
placita placita (Banks). Ala., La., Tex.
Cryptocheilus placitus Banks, 1910. N. Y. Ent. Soe., Jour. 18: 122. °.
Pseudagenia apicipennis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 128. 3.
placita sonorensis Townes. Calif.; Mexico (Nogales).
Ageniella (Priophanes) placita sonorensis Townes, 1957. U. S. Natl. Mus., Bul. 209: 185. °.
rufescens (Banks). Kans., N. Mex., Ariz., Wyo.
Priocnemis rufescens Banks, 1939. Canad. Ent. 71: 229. 9.

Genus AGENIELLA Subgenus AGENIELLA Banks

Ageniella Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 222.
Type-species: Pompilus (Agenia) acceptus Cresson. Orig. desig.
accepta (Cresson). Transcont. in L. Austr. Zone; Mexico (Coahuila). Prey: Lycosa sp.
Pompilus (Agenia) acceptus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 130. °.
Pseudagenia texana Banks, 1910. N. Y. Ent. Soc., Jour. 18: 125. ¢.
Ageniella adara Brimley, 1934. Ent. News 45: 41. 6.

Biology: Dow, 1930. Psyche 37: 182 (prey).
blaisdelli (Fox). South. B. C. to south. Calif., Idaho, Utah in Transit. and U. Austr. Zones.
Ecology: Nests in compacted sand along river. Prey: Pardosa sp., Tarentula sp., prob.
kochi (Keys.).
Pseudagenia Blaisdelli Fox, 1892. Ent. News 3: 171. 2.
Ageniella praestans Banks, 1914. N. Y. Ent. Soc., Jour. 22: 305. 6.

Biology: Wasbauer and Leech, 1973. Pan-Pacifie Ent. 49: 182 (prey, nest).
conflicta Banks. Alta., U. S., Transcont., mostly in U. Austr. Zone; Guatemala. Prey: Trochosa
gosiuta Chamb., T. avara (Keys.), T. sp., Arctosa littoralis Hentz, Pardosa valens
Barnes, Lycosa spp.
Pompilus (Agenia) petiolatus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 127. 6. Preoce.
Ageniella accepta var. conflicta Banks, 1944. Mus. Compar. Zool., Bul. 94: 176. 9.

Biology: Hurd and Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey). —Townes, 1957. U.
S. Natl. Mus., Bul. 209: 211 (prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc.
Pub. 3: 106-107 (prey capture, transport). —Kurezewski and Kurczewski, 1968. Kans. Ent.
Soc., Jour. 41: 7 (prey transport).

coronata Banks. South. B. C. to south. Calif., Utah in Transit. and U. Austr. Zones. Prey:
Clubionidae sp., prob. Liocranoides sp.
Ageniella coronata Banks, 1919. Mus. Compar. Zool., Bul. 63: 242. 9.

Biology: Wasbauer and Leech, 1973. Pan-Pacific Ent. 49: 182 (prey).

Superfamily POMPILOIDEA 1541

cupida (Cresson). N. Y., Pa., Va., W. Va., Ga., Iowa, Kans., Colo.; Guatemala.
Pompilus (Agenia) cupidus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 122. 9.
euphorbiae (Viereck). Common in Calif., less common in B. C., Oreg., Ariz., N. Mex., Colo., and
Tex. Prey: Agelenopsis sp.; Pardosa sp.
Agenia euphorbiae Viereck, 1903 (1902). Acad. Nat. Sci.Phila., Proc. p. 734. d.
Ageniella subaequalis Banks, 1919. Mus. Compar. Zool., Bul. 63: 243. ¢.

Biology: Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 7 (prey).

evansi Townes. Ariz., N. Mex.; Mexico (Teotihuacan). Prey: Lycosa sp., Tarentula kochi Keys.
Ageniella (Ageniella) evansi Townes, 1957. U. S. Natl. Mus., Bul. 209: 205. 2.

Biology: Townes, 1957. U.S. Natl. Mus., Bul. 209: 206 (prey). —Evans, 1959. Kans. Ent. Soc.,
Jour. 32: 76 (prey).
grisea Townes. Idaho (Hollister, Wendell).
Ageniella (Ageniella) grisea Townes, 1957. U.S. Natl. Mus., Bul. 209: 195. @.
mintaka Brimley. Fla., N. C., Va., W. Va.
Ageniella mintaka Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 202. d.
neglecta Banks. Colo., N. Mex., Ariz.; Mexico (Puebla, Zacatecas, Teotihuacan).
Ageniella neglecta Banks, 1944. Mus. Compar. Zool., Bul. 94: 176. ¢.
nivalis (Cameron). Ariz.; Mexico (Durango, Guerrero). Host: Lycosa sp.
Salius nivalis Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 182, pl. 11, fig. 2. °.

Biology: Evans, 1964. Kans. Ent. Soc., Jour. 37: 306-307 (prey transport).
norata Banks. Ont. and Mass. south to N. C., Ind., Kans. Ecology: Occurs in woods.
Ageniella norata Banks, 1914. N. Y. Ent. Soc., Jour. 22: 305. d.
Ageniella cupidella Banks, 1915. Canad. Ent. 47: 400. 9.
pallida Banks. Iowa, Nebr., Kans., Tex.
Ageniella pallida Banks, 1945. Psyche 52: 106. °.
partita Banks. Transcont. in U. Austr. and L. Austr. Zones; Mexico (Durango, Puebla, Nayarit,
Zacatecas, Hidalgo). Parasite: Ceropales hatoda Brim. Prey: Gnaphosa sericata (Koch),
Zelotus sp., Arctosa littoralis (Hentz), A. sp., Pardosa falcifera Camb.
Ageniella partita Banks, 1919. Mus. Compar. Zool., Bul. 63: 244. ¢.
Alasagenia rubineus Dreisbach, 1950. Ent. News. 61: 68. °.

Biology: Krombein, 1955. Brooklyn Ent. Soc., Bul. 50: 15 (parasite). —Townes, 1957. U.S.

Natl. Mus., Bul. 209: 194 (prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub.
3: 107 (prey transport). —Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 395 (prey).
—Kurczewski, 1963. Fla. Ent. 46: 209 (prey transport). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 7-8 (prey). —Kurezewski and Kurczewski, 1968. Kans. Ent.
Soc., Jour. 41: 369 (prey).

seminole Townes. Fla.
Ageniella (Ageniella) seminole Townes, 1957. U. S. Natl. Mus., Bul. 209: 196. 2, d.

submetallica (Banks). Tex. (Austin).
Pseudagenia submetallica Banks, 1917. Mus. Compar. Zool., Bul. 61: 108. °.

utilis delicata Banks. D. C., Va., S. C.
Ageniella delicata Banks, 1944. Mus. Compar. Zool., Bul. 94: 174. d.
Ageniella restricta Banks, 1944. Mus. Compar. Zool., Bul. 94: 175. d.

utilis utilis (Cameron). Fla., La., Tex.; Mexico (Tabasco), Panama.
Pseudagenia utilis Cameron, 1891. Biol. Cent.-Amer., Hym. v. 2, p. 170. °.
Ageniella obscura Banks, 1925. Mus. Compar. Zool., Bul. 67: 331. 2.

vogeli Townes. Carol. Fauna, Pa. to Ga., Kans. Prey: Lycosa avara (Keys.), juv.
Ageniella (Ageniella) vogeli Townes, 1957. U.S. Natl. Mus., Bul. 209: 202. 2, o.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 203 (prey).
Genus AGENIELLA Subgenus AMERAGENIA Banks

Ameragenia Banks, 1945. Bol. Ent. Venezolana 4: 125.
Type-species: Ameragenia irene Banks. Monotypic.

1542 Hymenoptera in America North of Mexico

Pseudageniella Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 22-23.
Type-species: Pompilus rusticus Fabricius. Orig. desig.
Allageniella Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 23.
Type-species: Allageniella obsoleta Haupt. Orig. desig.
Brachyagenia Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 25.
Type-species: Brachyagenia nigra Haupt. Orig. desig.
Parageniella Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 26.
Type-species: Priocnemis rufofemoratus Taschenberg. Orig. desig.
fasciata Townes. Tex. (Brownsville); Mexico (Cordoba).
Ageniella (Ameragenia) fasciata Townes, 1957. U. S. Natl. Mus., Bul. 209: 217. 9.
salti (Banks). Fla.; Cuba. Prey: Clubiona spp.
Priocnemella salti Banks, 1928. Studies on Cuban Insects, v. 1, p. 6. @.
Priocnemis osceola Banks, 1939. Canad. Ent. 71: 230. °.

Biology: Townes, 1957. U. S. Natl. Mus., Bul. 209: 219 (prey). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 8 (prey).

striga Townes. Tex. (Brownsville).
Ageniella (Ameragenia) striga Townes, 1957. U. S. Natl. Mus., Bul. 209: 216. °.

Genus PRIOCNEMELLA Banks

Priocnemis subg. Priocnemella Banks, 1925. Mus. Compar. Zool., Bul. 67: 337.
Type-species: Priocnemis (Priocnemella) fairchildi Banks. Desig. by Pate, 1946.
Eragenia Banks, 1946. Mus. Compar. Zool., Bul. 96: 421.
Type-species: Eragenia infelix Banks. Orig. desig.
Cosmagenia Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 27-28.
Type-species: Agenia amabilis Taschenberg. Orig. desig.
tabascoensis (Cameron). South Tex. to Ecuador.
Pseudagenia tabascoensis Cameron, 1891. Biol. Cent.-Amer., Hym., v. 3, p. 172. 3.
Ageniella rufula Banks, 1945. Bol. Ent. Venezolana 4: 117. °.

SUBFAMILY POMPILINAE

Revision: Evans, 1966. Amer. Ent. Soc., Mem. 20: 1-442, 2 text figs., 80 maps, 11 pls.
(Mexican and Cent. Amer. spp.; includes records and synonymy of some spp. north of

Mexico).
TRIBE APORINI

Wasps of this tribe, so far as known, prey upon trapdoor spiders (Ctenizidae); the burrow of
the spider is utilized as a nest by the wasp.

Revision: Bradley, 1944. Amer. Ent. Soc., Trans. 70: 32-139 (New World spp.).
Genus EPIPOMPILUS Kohl

Epipompilus Kohl, 1885. K. K. Zool.-Bot. Ges. Wien, Verh. 34: 57.

Type-species: Epipompilus maximiliani Kohl. Desig. by Ashmead, 1900.
Aulocostethus Ashmead, 1902. Canad. Ent. 54: 133.

Type-species: Aulocostethus bifasciatus Ashmead. Orig. desig.
Epicostethus Banks, 1947. Mus. Compar. Zool., Bul. 99: 445.

Type-species: E'picostethus williamsi Banks. Orig. desig.

Revision: Evans, 1961. Psyche 68: 25-37.
pulcherrimus (Evans). Fla. (Royal Palm Hammock in Everglades Natl. Park); Bahamas

(Andros Is.).
Aulocostethus pulcherrimus Evans, 1955. Ent. News 66: 150. 2.

Superfamily POMPILOIDEA 1548

Genus APORUS Spinola

Genus APORUS Subgenus APORUS Spinola

Aporus Spinola, 1808. Insectorum Liguriae, v. 2, p. 5.
Type-species: Aporus bicolor Spinola. Desig. by Latreille, 1810.
Actenopoda Ashmead, 1902. Canad. Ent. 34: 88.
Type-species: Actenopoda rileyi Ashmead, Orig. desig. (=Planiceps niger Cresson).
Melanaporus Ashmead, 1902. Canad. Ent. 34: 132.
Type-species: Planiceps euferalis Fox. Orig. desig.
Odontaporus Bradley, 1944. Amer. Ent. Soc., Trans. 70: 110.
Type-species: Planiceps notabilis Smith. Orig. desig.
calcaratus (Fox). S. Fla.
Planiceps calcaratus Fox, 1893. Canad. Ent. 25: 115. 6.
concolor (Smith). Tex. to Calif., s. to Costa Rica.
Planiceps concolor Smith, 1860. Jour. Ent. 1: 80. 9.
Pompilus monticola Cameron, 1893. Biol. Cent.-Amer., Hym. 2: 190. d.
luxus assimilis (Banks). U. Sonor. and Transit. Faunas, Calif. and Utah to B. C. and Idaho.
Planiceps assimilis Banks, 1917. Mus. Compar. Zool., Bul. 61: 100. °.
luxus luxus (Banks). L. Sonor. Fauna of Calif., Ariz. Ecology: Occurs in sandy areas.
Planiceps luxus Banks, 1914. N. Y. Ent. Soc., Jour. 22: 304. 9, d.
niger (Cresson). Carol. and Austrorip. Faunas, Tex. and Fla. to Nebr., IIll., south. Ont. and
Mass.
Planiceps niger Cresson, 1867. Amer. Ent. Soc., Trans. 1: 136. 2. Preoce. in Pompilus.
Planiceps minor Fox, 1893. N. Y. Ent. Soe., Jour. 1: 55. d.
Planiceps dubius Fox, 1893. N. Y. Ent. Soc., Jour. 1: 55. 3.
Planiceps clericus Dalla Torre. 1897. Cat. Hym., v. 8, p. 281. N. name.
Planiceps compressus Banks, 1917. Mus. Compar. Zool., Bul. 61: 99. °.
Aporus (Aporus) hermes Bradley, 1944. Amer. Ent. Soc., Trans. 70: 94. 2, d.

Taxonomy: Evans, 1956. Ent. News. 67: 6 (Syn. of hermes).
notabilis prolongatus Evans. South. Tex.; Mexico (Tamaulipas, Nuevo Leon, Durango,
Veracruz). Typical notabilis (Sm.) ranges from south. Mexico to Costa Rica.
Aporus (Aporus) notabilis prolongatus Evans, 1966. Amer. Ent. Soc., Mem. 20: 57. 2, d.
notabilis pulchritarsis (Cameron). South. Ariz. to Oaxaca and Yucatan.
Pompilus (Planiceps) pulchritarsis Cameron, 1893. Biol. Cent.-Amer., Hym. 2: 185. ?.
Planiceps bequaerti Banks, 1931. Brooklyn Ent. Soc., Bul. 26: 131. 2. Intergrade with
typical notabilis (Sm.).
Planiceps yavapai Banks, 1933. Psyche 40: 1. 2.
UNPLACED TAxA OF APORUS SuBGENUS APORUS SPINOLA

concolor (Patton). Kans.
Planiceps concolor Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 367. °.

feralis (Cresson). Tex.
Planiceps feralis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 207. “2” = 6.
Nomen NupuM IN APORUS SuBGENuS APORUS SPINOLA

Actenopoda rileyi Ashmead, 1902. Canad. Ent. 34: 88.
Genus APORUS Subgenus PLECTRAPORUS Bradley

Aporus subg. Plectraporus Bradley, 1944. Amer. Ent. Soc., Trans. 70: 97.
Type-species: Planiceps hirsutus Banks. Orig. desig.
hirsutus (Banks). Oreg. and Idaho to Calif. and Ariz.; Mexico (Baja California). Ecology: Nests
in sand dunes in burrows of trapdoor spider prey. Prey: Aptostichus sp., prob.
stanfordianus Sm.
Planiceps hirsutus Banks, 1917. Mus. Compar. Zool., BuleGl99" oe

1544 Hymenoptera in America North of Mexico

Biology: Williams, 1928. Hawaii. Sugar Planter’s Assoc. Expt. Sta., Ent. Ser., Bul. 19:
135-140, figs. 201, 204 (hunting behavior).

Genus CHELAPORUS Bradley

Chelaporus Bradley, 1944. Amer. Ent. Soc., Trans. 70: 116.
Type-species: Pedinaspis anomala Banks. Orig. desig.

anomalus (Banks). Austrorip. Fauna of Tex.; Mexico south to Veracruz.
Pedinaspis anomala Banks, 1917. Mus. Compar. Zool., Bul. 61: 100. @.

Genus ALLAPORUS Banks

Allaporus Banks, 1933. Psyche 40: 2.
Type-species: Planiceps pulchella Banks. Orig. desig. (=Aporus rufiventris
Cresson).
aurulentus Evans. Ariz. (Cochise Co.).
Allaporus aurulentus Evans, 1966. Amer. Ent. Soc., Mem. 20: 78. 2, 6.
fascipennis Evans. Tex. (Cameron Co.).
Allaporus fascipennis Evans, 1959. Kans. Ent. Soce., Jour. 32: 29. 2, ¢.
pulchellus (Banks). Transcont. in U. S., Pa. to Oreg. in U. and L. Austr. Zones, south in Mexico
to Campeche.
Planiceps pulchella Banks, 1910. N. Y. Ent. Soc., Jour. 18: 123. 9.
Planiceps hesperus Banks, 1929. Psyche 36: 327. 2, d.
Euplaniceps aquilonaris Dreisbach, 1952. Ent. News 63: 95, 3 figs. d.
rufiventris (Cresson). Tex. and southeast. Ariz. south to Morelos and Michoacan.
Aporus rufiventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 207. 2. Preoce. in Pompilus.
Aporus minimus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 207. d.
Pompilus rufiventricosus Dalla Torre, 1897. Cat. Hym., v. 8, p. 318. N. name.
Allaporus mexicanus Evans, 1950. Ent. News 61: 3. 2.
smithianus (Cameron). South. Tex., Ariz., Calif. south to Morelos and Veracruz.
Pompilus (Aporus) smithianus Cameron, 1893. Biol. Cent.-Amer., Hym. 2: 191, pl. 11, fig.
ome

Allaporus amabilis Evans, 1950. Ent. News 61: 2. 9.
Genus PSORTHASPIS Banks

Pedinaspis subg. Psorthaspis Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 231.
Type-species: Ferreola laevifrons Cresson. Orig. desig.
Idopedinaspis Haupt, 1930. Berlin Zool. Mus., Mitt. 16: 724, 783.
Type-species: Ferreola laevifrons Cresson. Desig. by Haupt, 1937.
Idiopedinaspis (!) Neave, 1939. Nomencl. Zool. 2: 762.
Dicyrtomalis Bradley, 1944. Notulae Nat. 145: 11.
Type-species: Pompilus connexus Cresson. Orig. desig.
Taxonomy: Evans, 1954. Amer. Mus. Novitates 1662: 14-17 (keys to males, and all black
females).
australis (Banks). Tex.
Pedinaspis australis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 122. 6.
brimleyi (Malloch). Austrorip. Fauna, Tex., Ark. to N. C.
Pedinaspis brimleyi Malloch, 1928. Ent. Soe. Wash., Proe. 30: 101. 2.
legata (Cresson). Austrorip. Fauna, Tex., Kans., and Fla. to Tenn. and Md. Ecology: Occurs in
woods.
Pompilus (2?) legatus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 109. 2.
luctuosa (Banks). Tex.
Pedinaspis luctuosa Banks, 1910. N. Y. Ent. Soe., Jour. 18: 128. 2.
macronotum arizonensis Dreisbach. South. Ariz. Typical macronotum (Kohl) is Mexican.
Psorthaspis arizonensis Dreisbach, 1950. Brooklyn Ent. Soc., Bul. 45: 119. 9.
macronotum cressoni Bradley. Tex. south in Mexico to Veracruz.
Psorthaspis levis Bradley, 1944. Amer. Ent. Soc., Trans. 70: 43. 6.

Superfamily POMPILOIDEA 1545
Psorthaspis nahuatlensis Bradley, 1944. Amer. Ent. Soc., Trans. 70: 46, pl. 3, figs. 2, 13, 26.

é.
Psorthaspis cressoni Bradley, 1944. Amer. Ent. Soc., Trans. 70: 59. 2, d.

Taxonomy: Evans, 1954. Amer. Mus. Novitates 1662: 11 (as first reviser placed levis as syn.
of cressoni).
magna (Banks). Tex. (Lee Co.).
Aporus magnus Banks, 1910. N. Y. Ent. Soe., Jour. 18: 126. ¢.
mariae (Cresson). Austrorip. and Carol. Faunas, Fla. and Tenn. to III. and N. J. Ecology:
Occurs in woods.
Pompilus (?) mariae Cresson, 1867. Amer. Ent. Soc., Trans. 1: 108. 9.
Pedinaspis mariae var. antennalis Banks, 1921. Ent. Soe. Amer., Ann. 14: 21. @.

Taxonomy: Krombein, 1956. Ent. Soc. Wash., Proc. 58: 155 (sex association).

nigriceps (Banks). N. Mex., Ariz., Utah.
Pedinaspis nigriceps Banks, 1933. Psyche 40: 2. 2.

planata (Fox). Calif., Utah, Ariz.; Mexico (Baja California). Prey: Bothriocyrtum californicum

(Camb.).

Planiceps planatus Fox, 1892. Ent. News 3: 171. °@.
Pompilus aequus Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 99. 2.
Sophropompilus tumifrons Banks, 1917. Mus. Compar. Zool., Bul. 61: 103. ¢.
Pedinaspis bucephala Malloch, 1928. Ent. Soc. Wash., Proce. 30: 101. ¢.
Pedinaspis albocaudata Malloch, 1928. Ent. Soc. Wash., Proc. 30: 101. ¢.
Psorthaspis morosa Bradley, 1944. Amer. Ent. Soc., Trans. 70: 42. °.

Biology: Davidson, 1915. Ent. News 16: 233 (prey). —Jenks, 1938. Natl. Geog. Mag. 74:
807-828.

portiae conocephala Bradley. Tex.; Mexico (Nuevo Leon).
Psorthaspis conocephala Bradley, 1944. Amer. Ent. Soc., Trans. 70: 67. 3.

portiae portiae (Rohwer). South. Ariz. to Sinaloa and Jalisco, east to Nuevo Leon.
Pedinaspis (Psorthaspis) portiae Rohwer, 1920. U. S. Natl. Mus., Proe. 57: 228. 2.

sanguinea (Smith). Austrorip. Fauna, Tex. and Fla. to Kans. and N. C.
Ferreola sanguinea Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 170. 9°.
Parapompilus contiguus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 206. °.

texana (Cresson). Austrorip. Fauna of Tex.
Parapompilus texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 206. 2.

vicina (Cresson). Tex.; Mexico (Tamaulipas).
Parapompilus vicinus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 206. 2. Preoccupied in
Pompilus.
Pompilus bombinator Dalla Torre, 1897. Cat. Hym., v. 8, p. 277. N. name.

TRIBE POMPILINI

Revision: Evans, 1950-51. Amer. Ent. Soc., Trans. 75: 133-270; 76: 207-361; 77: 203-340.
Taxonomy: Dreisbach, 1949. Ent. Amer. (n. s.) 29: 5-12 (key to N. Amer. and Antillean
genera).
Genus TASTIOTENIA Evans

Tastiotenia Evans, 1950. Amer. Ent. Soce., Trans. 75: 150.
Type-species: Tastiotenia festiva Evans. Orig. desig.

festiva Evans. South. Calif. to west. Tex.; Mexico (Sonora). Ecology: Occurs on sandy deserts;
probably nests on side of pre-existing burrow in sand. Prey: Latrodectus mactans F.
Tastiotenia festiva Evans, 1950. Amer. Ent. Soe., Trans. 75: 152. °.

Taxonomy: Evans, 1954. Pan-Pacifie Ent. 30: 103. d.
Biology: Evans, 1961. Southwest. Naturalist 6: 51-52 (prey, probable nest site).

1546 Hymenoptera in America North of Mexico

Genus CHALCOCHARES Banks

Psammochares subg. Chalcochares Banks, 1917. Mus. Compar. Zool., Bul. 61: 107.
Type-species: Psammochares hirsutifemur Banks. Monotypic.
Anotochares Banks, 1939. Canad. Ent. 71: 225, 228.
Type-species: Anotochares engleharti Banks. Monotypic.
engleharti (Banks). Tex.; Mexico east of Sierra Madre Occidentale, south to Hidalgo.
Anotochares engleharti Banks, 1939. Canad. Ent. 71: 228. @.
hirsutifemur (Banks). L. Sonor. Fauna, west. Tex. to Calif.; south in Mexico to Zacatecas.
Psammochares hirsutifemur Banks, 1914. N. Y. Ent. Soc., Jour. 22: 304. 2.

Genus EVAGETES Lepeletier

Evagetes Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 390.
Type-species: Evagetes bicolor Lepeletier. Monotypic (=~Aporus dubius Vander
Linden).
Sophropompilus Howard, 1901. Insect Book, pl. VII, fig. 13.
Type-species: Pompilus ingenuus Cresson. Monotypic.
Nannopompilus Ashmead, 1902. Canad. Ent. 34: 82.
Type-species: Nannopompilus argenteus Ashmead. Monotypic (~Pompilus parvus
Cresson).
Leuchimon Haupt, 1930. Zool. Mus. Berlin, Mitt. 16: 792.
Type-species: Leuchimon palmatus Haupt. Orig. desig.
Psammocharoides Moczar, 1946. Hist. Nat. Mus. Natl. Hungarici, Ann. 39: 114.
Type-species: Pompilus crassicornis Shuckard. Orig. desig.
Streptosella Dreisbach, 1950. Amer. Midland Nat. 43: 570.
Type-species: Streptosella steyskali Dreisbach. Orig. desig.
Evagetes subg. Contemptevagetes Wolf, 1970. Beitr. z. Ent. 20: 52.
Type-species: Pompilus contemptus Tournier. Orig. desig.
Evagetes subg. Carinevagetes Wolf, 1970. Beitr. z. Ent. 20: 52.
Type-species: Pompilus crassicornis Shuckard. Orig. desig.

Members of this genus are social parasites of other Pompilini. The females seek the freshly
filled nests of their hosts, usually in sandy places, dig into them, destroy the host egg, deposit
one of their own, and refill the burrow.

asignus Dreisbach. Mass. south to Fla., west to Mich., Alta., Colo., Tex.; Mexico (Chihuahua,
Durango, Zacatecas).
Evagetes asignus Dreisbach, 1956. Ent. News 67: 147, 2 figs. d, 2.
calefactus Evans. Calif., Ariz., Tex., Mont. ?
Evagetes calefactus Evans, 1966. Amer. Ent. Soc., Mem. 20: 136. 2, d.

crassicornis consimilis (Banks). Canad. and Transit. Faunas, Yukon and Man. to higher parts
of N. Mex. and Calif.
Pompiloides consimilis Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 228. 2, d.
crassicornis crassicornis (Shuckard). Canad. Zone, N. S. and Yukon to Wash., N. Dak., Mich.,
and Pa.; Europe.
Pompilus crassicornis Shuckard, 1835. Essay on Indig. Fossor. Hym., p. 63. °.
Pompiloides rufibasis Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 226. 2.
Streptosella michiganensis Dreisbach, 1950. Amer. Midland Nat. 43: 571. Nom. nud., a
lapsus for steyskali.
Streptosella steyskali Dreisbach, 1950. Amer. Midland Nat. 43: 571, figs. 1, 2. d.
Streptosella albertensis Dreisbach, 1950. Amer. Midland Nat. 43: 572, figs. 3, 4. d.
Streptosella stricklandi Dreisbach, 1950. Amer. Midland Nat. 43: 574, figs. 5, 6. d.
hyacinthinus (Cresson). Transcont., L. Austr. to Transit. Zone. Ecology: Sandy places,
especially along streams.
Pompilus hyacinthinus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 90. 2.
Pompilus brevicornis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 90. 6.
Pompilus castaneus Provancher, 1882. Nat. Canad. 138: 35, 39. ¢.
Psammochares (Arachnophila) scudderi Banks, 1917. Mus. Compar. Zool., Bul. 61: 104,
107. 2.

Superfamily POMPILOIDEA 1547

Sophropompilus bradleyi Banks, 1919. Mus. Compar. Zool., Bul. 63: 237. 9.
Nannopompilus texanus Banks, 1944. Mus. Compar., Zool., Bul. 94: 170. 9.
ingenuus (Cresson). Transcont. in U. Austr. and lower part of Transit. Zone; Mexico
(Chihuahua, Mexico). Ecology: Occurs in sandy places, and on flowers of Solidago and
Daucus carota.
Pompilus ingenuus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 89. 2, ¢.
Pompilus brevicornis Taschenberg, 1869. Ztschr. Gesell. Naturw. 34: 50. ¢, 2. Preoce.
Pompilus Fiorentini Dalla Torre, 1897. Cat. Hym., v. 8, p. 289. N. name.
macswaini Evans. Calif., Wyo.
Evagetes macswaini Evans, 1957. Pan-Pacifie Ent. 33: 181, fig. 1. d, @.
mohave (Banks). N. Y. to Fla., through southern U. S. to Calif., in western Mexico south to
Guerrero and Puebla. Host: Anoplius apiculatus autumnalis (Bks.), A. americanus
juxtus (Cr.).
Sophropompilus quadrispinosus Banks, 1919. Canad. Ent. 51: 82. Preoce. in Evagetes.
Sophropompilus mohave Banks, 1933. Psyche 40: 6. 2.

Taxonomy: Krombein, 1953. Wasmann Jour. Biol. 10: 320-323. 2, ¢.

Biology: Evans, Lin and Yoshimoto, 1953. N. Y. Ent. Soe., Jour. 61: 72-77 (hosts).
padrinus minusculus (Banks). Austrorip. to Transit. Fauna, Tex. and Fla. to Man., Mich., Ont.,
and N. H.; Mexico (Tamaulipas, Coahuila). Host: Episyron q. quinquenotatus (Say).
Psammochares minusculus Banks, 1910. N. Y. Ent. Soc., Jour. 18: 118. 9.

Biology: Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 8-9 (host).
padrinus padrinus (Viereck). B. C., N. W. T. and Man. south to Baja California and, at
moderate to high elevations, to E] Salvador. Ecology: Occurs in sandy areas.
Anoplius (Pompilinus) padrinus Viereck, 1903 (1902). Acad. Nat. Sci. Phila., Proc. 54: 734.
é.

parvus (Cresson). Transcont. in Canadian to U. Austral Zones, Yukon, N. W. T. and N. S. to
Calif. and Ga., south to Costa Rica. Ecology: Occurs in sandy places.
Pompilus parvus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 453. 9.
Pompilus subviolaceus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 91. °.
Pompilus argenteus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 98. ¢.
Nannopompilus argenteus Ashmead, 1902. Canad. Ent. 34: 82.
Pompiloides minora Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 227. °.
subangulatus (Banks). Transcont., Huds. to Transit. Zone. Ecology: Occurs in sandy places,
and on flowers of Solidago and Spiraea.
Sophropompilus subangulatus Banks, 1919. Mus. Compar. Zool., Bul. 63: 237. 2, 6.
Psammochares (Sophropompilus) tebemi Brimley, 1936. Elisha Mitchell Sci. Soc., Jour. 52:

Avene
Genus AGENIOIDEUS Ashmead

Genus AGENIOIDEUS Subgenus RIDESTUS Banks

Ridestus Banks, 1912 (1911). N. Y. Ent. Soc. Jour. 19: 223.
Type-species: Psammochares transversalis Banks. Orig. desig. (=Psammochares
biedermani Banks).
biedermani (Banks). L. Sonor. Fauna, entering U. Sonor., Kans., Utah, Oreg., Mont. ?, south
into Mexico (Baja California, Chihuahua, Nuevo Leon, Hidalgo). Prey: Loxosceles devius
Gertsch and Mulaik.
Pompilus mexicanus Taschenberg, 1869. Ztschr. Ges. Naturw. 34: 52. 2. Preoce.
Psammochares biedermani Banks, 1910 (June). N. Y. Ent. Soc., Jour. 18: 116. d.
Psammochares striatulus Banks, 1910 (June). N. Y. Ent. Soc., Jour. 18: 119. 9.
Psammochares transversalis Banks, 1910 (Dec.). Psyche 17: 248. °.
Arachnophroctonus anahuacensis Bradley, 1944. Notulae Nat. 145: 10. N. name.

Biology: Evans, 1959. Kans. Ent. Soc., Jour. 32: 76 (prey transport).

1548 Hymenoptera in America North of Mexico
Genus AGENIOIDEUS Subgenus AGENIOIDEUS Ashmead

Agenioideus Ashmead, 1902. Canad. Ent. 34: 85.
Type-species: Pompilus hwmilis Cresson. Orig. desig.
Aporoideus Ashmead, 1902. Canad. Ent. 34: 86.
Type-species: Pompilus sericeus Vander Linden. Orig. desig.
humilis (Cresson). Transit. to L. Austr. Zones, N. B. and B. C. to Fla. and Calif., south to
Panama. Ecology: Nests in sandy places, and around cliffs, walls, and buildings in
pre-existing crevices. Parasite: Bombyliidae sp.; Mutillidae sp. Prey: Acacesia hamata
(Hentz), Araneus cornutus McC., A. sp., Neosconella pegnia (Walck.), Conepeira excelsa
(Bks.).
Pompilus humilis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 91. 9.

Taxonomy: Hurd, 1947. Pan-Pacifie Ent. 23: 133. 6.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 15 (prey). —Hurd, 1947. Pan-Pacific Ent.
23: 132 (nest, parasites). —Krombein, 1953. Brooklyn Ent. Soc., Bul. 48: 113-115 (prey,
nest). —Eberhard, 1971 (1970). Psyche 77: 243-247 (prey hunting).

Genus AGENIOIDEUS Subgenus GYMNOCHARES Banks

Gymnochares Banks, 1917. Mus. Compar. Zool., Bul. 61: 107, 108.
Type-species: Psammochares birkmanni Banks. Desig. by Pate, 1946.
birkmanni (Banks). Transcont. in L. Austr. Zone, entering U. Austr. in West. N. J., Kans.,
Wyo., and Oreg. southward in Mexico to Oaxaca. Prey: Herpyllus vasifer (Walck.).
Psammochares birkmanni Banks, 1910. N. Y. Ent. Soc., Jour. 18: 116. 6.
Gymnochares texana Banks, 1944. Mus. Compar. Zool., Bul. 94: 170. 2.

Biology: Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 369 (prey).
Genus SERICOPOMPILUS Howard

Sericopompilus Howard, 1901. Insect Book, pl. XI, fig. 17.
Type-species: Pompilus cinctipes Cresson. Monotypic @Ceropales apicalis Say).
angustatus (Cresson). L. and U. Austr. Zones of the Great Plains area; Tex. and N. Mex. to
Wyo., 8. Dak., and Mich.; in Mexico south to Jalisco and Guanajuato in Central Plateau.
Pompilus angustatus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 452. 2.
Psammochares fulvoapicalis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 117. @.
Sericopompilus fumosus Banks, 1933. Psyche 40: 5. ¢.
apicalis (Say). Austrorip. and Carol. Faunas, Tex. and Fla. to Nebr., Mich., and Conn. Ecology:
Nests in sand and soft earth. Parasite: Miltogrammini sp. Prey: Phidippus audax Hentz,
P. audax var. bryantae Kast., P. whitmanii Peckh., P. putnami Peckh., P. sp.,
Paraphidippus chrysis Walck., Thiodina sylvana (Hentz); Thomisus sp., Xysticus ferox
(Hentz), X. funestus Keys., Philodromus washita Bks.; Aysha gracilis (Hentz);
Lycosidae sp.; Oxyopes scalaris Hentz; Eriophora ravilla (Koch), Acanthepeira stellata
(Walck.), Hustala anastera (Walck.), Neoscona arabesca (Walck.), N. sp.
Ceropales apicalis Say, 1835. Boston Jour. Nat. Hist. 1: 366. ¢.
Pompilus fuscipennis Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 434. 2. Preoce.
Pompilus sordidus Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 160. 2. N. name.
Pompilus cinctipes Cresson, 1867. Amer. Ent. Soc., Trans. 1: 102. ¢. N. name.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 140-144 (prey,
nest). —Krombein, 1953. Ent. Soc. Wash. Proc. 55: 116-117 (prey, parasite). —Krombein,
1955. Ent. Soc. Wash., Proc. 57: 149 (prey). —Evans and Yoshimoto, 1955. Kans. Ent. Soc.,
Jour. 28:17-18 (prey). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 101 (prey). —Krombein,
1959. Ent. Soe. Wash., Proc. 61: 193 (prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer.,
Misc. Pub. 3: 89-91 (prey, nest, parasite, life history). —Kurezewski and Kurczewski, 1973.
Kans. Ent. Soc., Jour. 46: 66 (prey).

neotropicalis (Cameron). L. Austr. Zone, Calif., Ariz., Kans., Ark., Tex., Ala., Ga., Fla., and N.
C., south to Costa Rica.
Pompilus neotropicalis Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 203. 9.

Superfamily POMPILOIDEA 1549

Pompilus guatemalensis Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 206. d.
Psammochares posticatus Banks, 1910. N. Y. Ent. Soe., Jour. 18: 119. 6.
Psammochares fuscipennis var. georgiana Banks, 1911. Ent. Soc., Wash., Proc. 13: 238. °.

Genus EPISYRON Schiodte

Episyron Schiodte, 1837. Kroyer’s Naturhist. Tidsskr. 1: 341.
Type-species: Sphex rufipes Linnaeus. Monotypic.
Spilopompilus Ashmead, 1902. Canad. Ent. 34: 81.
Type-species: Pompilus biguttulatus (!) Fabricius. Orig. desig.
Epizuron (!) Ashmead, 1902. Canad. Ent. 34: 82.

These are extremely active wasps, two of which have been dubbed the “Tornado Wasp” and
the “Hurricane Wasp” by the Peckhams. They nest in short burrows in sand, which they provi-
sion invariably with orb-weavers (Araneidae). They visit flowers of many kinds.

biguttatus biguttatus (Fabricius). Temperate N. Amer. east of the Rockies, Tex. and Fla. to
Labr. and N. W. T. Ecology: Nests in sand. Parasite: Miltogrammini sp. Prey: Araneus
diadematus Clerck, A. spp., Hustala anastera (Walck.), Metepeira labyrintha (Hentz),
Neoscona benjamina (Walck.), N. sp., near sacra (Walck.), N. sp.
Pompilus biguttatus Fabricius, 1798. Sup. Ent. System., p. 249. 9.
Episyron atrytone Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 229. 9.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 138-140 (prey
transport). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 9-10 (prey, nest, parasite).
—Krombein, 1953. Wasmann Jour. Biol. 264-266 (nest, prey, parasite). —Krombein, 1953.
Ent. Soc. Wash., Proc. 55: 115-116 (nest, prey). —Evans and Yoshimoto, 1962. Ent. Soc.
Amer., Mise. Pub. 3: 93 (prey). —Kurezewski, 1963 (1962). Brooklyn Ent. Soc., Bul. 57: 88
(prey, nest). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 9 (prey).
—Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 369 (prey). —Kurezewski
and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 66 (prey).

biguttatus californicus (Banks). U. Sonor. to Canad. Fauna, Calif. to B. C., Alta., and S. Dak.
Prey: Neoscona naiba Chamb. and Gertsch.
Psammochares californica Banks, 1910. N. Y. Ent. Soc., Jour. 18: 117. d.

Biology: Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 396 (prey).
biguttatus montezuma (Cameron). Ariz., N. Mex., west. Tex. south in Mexico to Morelos,
Guerrero and Veracruz. Prey: Neoscona naiba Chamb. and Gertsch.
Pompilus montezuma Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 193, pl. 11, fig. 10.

OF
Episyron arizonica Banks, 1933. Psyche 40: 5. °.

Biology: Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 396 (prey).
conterminus posterus (Fox). N. Y. to Fla., west to south. Calif, Ill., south to Costa Rica.

Ecology: Nests in sand. Parasite: Senotainia sp., Miltogrammini sp.; Hvagetes sp. Prey:
Eustala anastera (Walck.), E. cepina (Walck.), E. sp., Araneus ozarkensis (Arch.), A. sp.,
Larinia sp., Neoscona arabesca (Walck.), N. vertebrata McCook, N. sp., Neosconella
pegnia (Walck.), Argiope aurantia Lue., Drexelia directa (Hentz), Gea heptagon (Hentz).
Typical conterminus (Sm.) occurs in South America.

Pompilus posterus Fox, 1893 (May). Canad. Ent. 25: 115. ?, d.

Pompilus exactus Cameron, 1893 (Sept.). Biol. Cent.-Amer., Hym., v. 2, p. 202. 2, d.

Pompilus porus Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 98. 2, d.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 439 (larva).

Biology: Evans, 1950. Amer. Ent. Soc., Trans. 75: 225 (prey). —Krombein, 1953. Wasmann
Jour. Biol. 10: 266-268 (prey, nest, parasite). —Krombein, 1955. Ent. Soe. Wash., Proce. 57:
149-150 (prey, nest). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 102 (prey, nest,
parasite). —Krombein, 1959. Ent. Soe. Wash., Proc. 61: 194 (prey, nest). —Wasbauer and
Powell, 1962. Kans. Ent. Soc., Jour. 35: 396 (prey). —Kurezewski, 1963. Fla. Ent. 46:
210-212 (prey, nest). —Krombein, 1964. Amer. Mus. Novitates 2201: 10-11 (prey, nest).

1550 Hymenoptera in America North of Mexico

—Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 9 (prey). —Kurezewski and
Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 369 (prey). —Kurezewski and Kurczewski,
1973. Kans. Ent. Soe., Jour. 46: 66-67 (prey, nest, parasite).
oregon Evans. Transit. and Canad. Fauna, Yukon and Alta. to Wyo., Colo. and Calif. Prey:
Araneus trifolium (Hentz).
Episyron oregon Evans, 1950. Amer. Ent. Soc., Trans. 75: 231. 9, d.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 481 (prey, transport).
quinquenotatus hurdi Evans. Transit. and U. Sonor. Fauna, B. C. and Alta. to Utah and Calif.;
Mexico (Baja California). Prey: E'‘peira sp.
Episyron quinquenotatus hurdi Evans, 1950. Amer. Ent. Soc., Trans. 75: 221. 2, d.
quinquenotatus quinquenotatus (Say). Canad. in U. Austr. Zone, N.S., Sask., and B. C. to
Mont., Colo., Tex., and Ala. Ecology: Nests in sand. Parasite: Senotainia trilineata
(Wulp). Prey: Metazygia wittfeldae (McC.), Araneus cornutus Clerck, A. patagiatus
Clerck, A. diadematus Clerck, A. sp. near diadematus Clerck, A. thaddeus (Hentz), A.
marmoreus Clerck, A. ozarkensis (Arch.), A. sp., Epeira displicata Hentz, E. foliata
Fourer., Singa eugeni Levi, Neoscona arabesca (Walck.), N. minima (Camb.), N. sp.,
Eustala anastera (Walck.), E. triflex (Walck.), E. cepina (Walck.), Allepeira stellata
(Walek.).
Pompilus 5-notatus Say, 1835. Boston Jour. Nat. Hist. 1: 304. @.
Pompilus griseus Provancher, 1882. Nat. Canad. 13: 35, 36. 6.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 439, fig. 46 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 126-138 (prey,
nest). —Evans, 1950. Amer. Ent. Soc., Trans. 75: 220 (prey). —Evans and Yoshimoto, 1955.
Kans. Ent. Soc., Jour. 28: 18 (prey). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 102
(prey). —Kurezewski, 1961. Brooklyn Ent. Soc., Bul. 56: 24 (prey transport). —Krombein,
1961. Brooklyn Ent. Soc., Bul. 56: 63 (prey, nest). —Evans and Yoshimoto, 1962. Ent. Soc.
Amer., Mise. Pub. 3: 91-92 (prey, nest, parasite). —Evans, 1963. Ent. News 74: 238-239, fig.
2 (prey, nest). —Krombein, 1964. Brooklyn Ent. Soc., Bul. 58: 118 (prey, nest).
—Kurczewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 9-12 (prey). —Kurezewski
and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 370 (prey). —Evans, 1970. Mus. Compar.
Zool., Bul. 140: 482 (prey). —Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46:
67 (prey).

snowi (Viereck). Transcont. in L. and U. Austr. Zones, n. to N. J., Ill., S. Dak., Mont., and
Oreg.; Mexico (Baja California, Chihuahua, Durango). Parasite: Miltogrammini sp. Prey:
Neoscona benjamina (Walck.), Cyclosa conica (Pall.).
Anoplius (Pompilinus) snowi Viereck, 1906. Amer. Ent. Soc., Trans. 32: 202. d.
Psammochares maneei Banks, 1910. N. Y. Ent. Soc., Jour. 18: 117. @.
Episyron laevis Banks, 1933. Psyche 40: 4. ?.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 10 (prey transport). —Krombein, 1953.
Wasmann Jour. Biol. 10: 268-269 (prey, nest, parasite). —Krombein, 1953. Ent. Soc. Wash.,
Proc. 55: 116 (prey, nest).

Genus POECILOPOMPILUS Howard

Poecilopompilus Howard, 1901. Insect Book, pl. V. fig. 1, and pl. XI, fig. 18.

Type-species: Pompilus navus Cresson. Desig. by Ashmead, 1902 (= Ceropales
interrupta Say).

Batozonus Howard, 1901. Insect Book, pl. XI, fig. 24.
Type-species: Pompilus algidus Smith. Monotypic.

Batazonus (!) Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 221, 224.

Eubatozonus Haupt, 1950. Explor. Pare Nat. Albert, Mission de Witte, fasc. 69, p. 52.
Type-species: Eubatozonus pulcher Haupt. Orig. desig.

Wasps of this genus, as in the closely related Episyron, prey exclusively upon Araneidae, and
nest in sandy places; they are also frequent visitors on flowers.

Superfamily POMPILOIDEA 1551

algidus algidus (Smith). Austrorip. and Carol. Fauna, Tex. and Fla. to Mass., Mich., and N.
Dak.
Pompilus algidus Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 158. 9.
algidus coquilletti (Provancher). U. Sonor. Fauna, south. Calif. and N. Mex. to Utah and
north. Calif.
Pompilus coquilletti Provancher, 1887. Addit. Corr. Fauna Ent. Canad., Hym., p. 261. d.
Batazonus (!) flavipennis Banks, 1921. Ent. Soc. Amer., Ann. 14: 20. 2.
algidus marcidus (Smith). South. Tex. s. to Nicaragua.
Pompilus marcidus Smith, 1862. Jour. Ent. 1: 395. °.
Pompilus pygidialis Kohl, 1886. Zool.-Bot. Ges. Wien, Verhandl. 36: 314, 334. 9.
algidus willistoni (Patton). S. Dak. and Colo. to N. Mex. and Ariz., s. in Mexico to Durango
and Baja California.
Pompilus willistoni Patton, 1879. U.S. Geol. and Geog. Survey, Bul. 5: 352. 9.
flavopictus flavopictus (Smith). South. Tex to Surinam and Colombia. Other subspp. occur in
the Antilles and Brazil.
Pompilus flavopictus Smith, 1862. Jour. Ent. 1: 396. °.
interruptus cressoni (Banks). North. Va. and east. Ohio to Mass. Ecology: Nests in sand. Prey:
Epeira trifolium Hentz, Argiope trifasciata (Forsk.).
Batazonus (!) interruptus var. cressoni Banks, 1944. Mus. Compar. Zool., Bul. 94: 167. 2,
é.

Biology: Evans, 1950. Amer. Ent. Soc., Trans. 75: 253 (prey). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 12-13 (prey, nest).
interruptus interruptus (Say). Transcont. in L. and U. Austr. Zones, north to N. J., Ohio,
Mich., S. Dak., Utah, and south. Calif. Ecology: Nests in sand. Parasite: Miltogrammini
sp. Prey: Neoscona benjamina (Walck), N. vertebrata McCook, N. sp., Araneus
cornutus Cl., A. trifolium (Hentz), Argiope aurantia Luc., A. trifasciata (Forsk.), Epeira
foliata (Fourer.), Acanthepeira stellata Walck.; Lycosidae sp.
Ceropales interrupta Say, 1835. Boston Jour. Nat. Hist. 1: 365. d.
Pompilus navus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 105. 2, d.
Pompilus ichneumoniformis Patton, 1879. U. S. Geol. Survey, Bul. 5: 351. 2. Preoce.
Pompilus ichneumonoides Dalla Torre, 1897. Cat. Hym., v. 8, p. 295. N. name.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 438, figs. 26-30 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 152-153 (prey,
nest). —Krombein, 1953. Wasmann Jour. Biol. 10: 269-272 (nest, prey, parasite).
—Strandtmann, 1953. Kans. Ent. Soc., Jour. 26: 45-46 (prey). —Evans and Yoshimoto,
1955. Kans. Ent. Soc., Jour. 28: 18 (prey, nest). —Evans and Yoshimoto, 1962. Ent. Soe.
Amer., Misc. Pub. 3: 93-94 (nest, prey). —Kurezewski and Kurezewski, 1968. Kans. Ent.
Soc., Jour. 41: 13 (prey).

interruptus semiflavus Evans. Calif. (Central Valley).
Poecilopompilus interruptus semiflavus Evans, 1966. Amer. Ent. Soc., Mem. 20: 224. 2, d.

Genus TACHYPOMPILUS Ashmead

Tachypompilus Ashmead, 1902. Canad. Ent. 34: 83
Type-species: Tachypompilus abbotti Ashmead. Orig. desig. @Sphex analis
Fabricius).
Arachnophroctonus Ashmead, 1902. Canad. Ent. 34: 83. Preoce.
Type-species: Ceropales ferruginea Say. Orig. desig.
Balanoderes Haupt, 1929. Berlin Zool. Mus., Mitt. 15: 119, 155.
Type-species: Sphex analis Fabricius. Desig. by Haupt, 1929.
Afropompilus Arnold, 1936. Transvaal Mus., Ann. 18: 107.
Type-species: Pompilus ignitus Smith. Orig. desig.
Zarachnophroctonus Pate, 1946. Amer. Ent. Soc., Trans. 72: 130. N. name.
Wasps of this genus occur around walls, stone piles, and buildings, where they nest in crevices,
supplying the nest with Lycosid and Pisaurid spiders; they also visit flowers.

1552 Hymenoptera in America North of Mexico

ferrugineus annexus (Banks). Tex.; northeast. Mexico ? Ecology: Nests in loose dry soil
beneath house. Prey: Lycosa rabida (Walck.), L. antelucana Mont.
Arachnophroctonus ferrugineus var. annexus Banks, 1944. Mus. Compar. Zool., Bul. 94:
168. @.

Biology: Strandtmann, 1953. Kans. Ent. Soc., Jour. 26: 46-48, fig. 1 (nest, prey). —Evans and
Yoshimoto, 1962. Ent. Soe. Amer., Mise. Pub. 3: 87 (prey).
ferrugineus ferrugineus (Say). L. and U. Austr. Zones, east. Tex. and Fla. to N. J., Ohio,
Minn., and Colo. Ecology: Nests in crevices in walls. Prey: Lycosa helluo Walck., L.
osceola Wall., L. rabida Walck., L. spp.; Dolomedes tenebrosus Hentz, D. sp.
Ceropales ferruginea Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 334. d.

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 78-83 (prey, nest). —Rau, 1922. Acad.
Sci. St. Louis, Trans. 24: 12 (prey). —Evans, 1950. Amer. Ent. Soc., Trans. 75: 260 (prey).
—Evans and Yoshimoto, 1962. Ent. Soc Amer., Misc. Pub. 3: 87 (prey). —Kurezewski and
Kurcezewski, 1968. Kans. Ent. Soe., Jour. 41: 13 (prey). —Kurezewski and Kurczewski, 1973.
Kans. Ent. Soc., Jour. 46: 67 (prey).

ferrugineus nigrescens (Banks). North. Va. and west. N. Y. to N. H.
Arachnophroctonus ferrugineus var. nigrescens Banks, 1944. Mus. Compar. Zool., Bul. 94:
GE), OD. Bie
ferrugineus torridus (Smith). South. parts of Tex., N. Mex., Ariz., and Calif.; Mexico s. to
Chiapas.
Pompilus torridus Smith, 1862. Jour. Ent. 1: 396. 2.
Arachnophroctonus ferrugineus var. unicolor Banks, 1944. Mus. Compar. Zool., Bul. 94:
168. 2. Preoce.
Tachypompilus ferrugineus Yavapai Evans, 1950. Amer. Ent. Soc., Trans. 75: 263. N.
name.
unicolor cerinus Evans. South. Calif., Utah, N. Mex. and Tex. s. to Costa Rica.
Tachypompilus unicolor cerinus Evans, 1966. Amer. Ent. Soc., Mem. 20: 235, map 41. 9°,
é.

unicolor unicolor (Banks). L. and U. Sonor. Faunas, Calif. and Utah, Wyo., Idaho, and B. C.
Arachnophroctonus unicolor Banks, 1919. Mus. Compar. Zool., Bul. 63: 239, 240. 2.
Arachnophroctonus latifrons Banks, 1939. Canad. Ent. 71: 229. 9, 6.

Genus ANOPLIUS Dufour

Genus ANOPLIUS Subgenus LOPHOPOMPILUS Radoszkowski

Lophopompilus Radoszkowski, 1887. Soc. Ent. Rossica, Horae 21: 42.
Type-species: Pompilus grandis Eversmann. Desig. by Ashmead, 1902 (=Sphex
samariensis Pallas).
_Pompilogaster Howard, 1901. Insect Book, pl. V, fig. 19.
Type-species: Pompilus aethiops Cresson. Monotypic.
Pompilogastra Ashmead, 1902. Canad. Ent. 34: 81.
Type-species: Pompilus aethiops Cresson. Orig. desig.

Wide-ranging, strong-flying species which prey chiefly on Lycosidae and nest in tunnels in
sand or soft earth. They are frequent visitors to flowers and occasionally come to light at night.

Revision: Regan, 1923. Ent. Soc. Amer., Ann. 16: 177-194.
aethiops (Cresson). N.S. to B. C. s. to Ga., Tex., Calif. in Transition to L. Austral Zones;

Mexico (higher altitudes), Guatemala. Ecology: Occurs in fields, meadows, and prairies;
commonly visits flowers; nests in pre-existing crevices in soft soil. Prey: Lycosa
frondicola Em., L. helluo Walck., L. gulosa Walck., L. carolinensis Walck., L. santrita
Chamb. and Ivie, Schizocosa ocreata (Hentz).

Pompilus aethiops Cresson, 1865. N. Y. Ent. Soe., Jour. 4: 451. d (2 misdet.).

Psammochares ilione Banks, 1910. Psyche 17: 249. @.

Lophopompilus azotus Banks, 1929. Psyche 36: 326. 9.

Superfamily POMPILOIDEA 1553

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 76: 217 (ecology, prey). —Evans and
Yoshimoto, 1962. Ent. Soe. Amer., Mise. Pub. 3: 70 (prey transport, nest). —Kurezewski
and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 13 (prey transport, nest). —Kurezewski
and Kurczewski, 1968. Kans. Ent. Soc., Jour. 46: 370 (prey transport). —Kurezewski and
Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 68 (prey).

atrox (Dahlbom). L. and U. Austr. Zones, N. Mex. and Fla. to S. Dak., Ont., and Mass. Ecology:
Occurs in fields, meadows and open woods; nests in sand or soft soil. Prey: Lycosa
rabida Walck., Dolomedes tenebrosus Hentz, D. urinator Hentz.
Pompilus atrox Dahlbom, 1843. Hym. Europaea, v. 1, p. 63. 2.

Biology: Peckham and Peckham, 1900. Wis. Nat. Hist. Soc., Bul. 1: 87 (prey). —Evans, 1951.
Amer. Ent. Soc., Trans. 76: 224 (ecology, prey). —Krombein, 1955. Ent. Soc. Wash., Proc.
57: 150 (prey). —Krombein, 1959. Ent. Soc. Wash., Proc. 61: 194 (prey).
bengtssoni (Regan). Austrorip. and Carol. Faunas, Tex. and Fla. to S. Dak., Mich., and Mass.
Lophopompilus bengtssoni Regan, 1923. Ent. Soc. Amer., Ann. 16: 185, 186. 2, 3.
carolina (Banks). Alleghanian Fauna, south. Que. to north. Ga. Ecology: Occurs in open
woodlands, and on flowers of Daucus carota; nests in mammal burrows or in bare soil.
Prey: Wadotes hybridus (Em.), W. sp.; Amaurobius bennetti (Blackw.).
Lophopompilus carolina Banks, 1921. Ent. Soe. Amer., Ann. 14: 20. 2.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 442, fig. 50 (larva).

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 76: 222 (ecology, prey). —Krombein, 1958. Ent.
Soe. Wash., Proc. 60: 53 (prey transport). —Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 63
(prey transport). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 70-71 (prey
transport, nest). —Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 68 (prey).

cleora (Banks). P. E. I. to B. C., south to N. C., Ind., Tex., Ariz., mostly in U. and L. Austral
Zones; Mexico (Baja California, Sonora, Jalisco, Puebla). Ecology: Nests in sand; rarely
visits flowers. Parasite: Senotainia litoralis Allen. Prey: Arctosa littoralis (Hentz),
Lycosa helluo Walck., Trochosa avara Keys., T. sp.
Psammochares cleora Banks, 1917. Mus. Compar. Zool., Bul. 61: 105. °.

Biology: Krombein, 1952. Amer. Ent. Soc., Trans. 78: 93 (prey). —Krombein, 1953. Ent. Soe.
Wash., Proc. 55: 116 (prey). —Evans and Yoshimoto, 1955. Kans. Ent. Soc., Jour. 28: 18
(prey, nest). —Krombein, 1958. Ent. Soc. Wash., Proc. 55: 116 (prey). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 69-70 (prey transport, nest, parasite).
—Kurcezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 13-14 (prey). —Kurezewski
and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 371 (prey). —Kurezewski and Kurezewski,
1973. Kans. Ent. Soc., Jour. 46: 68 (prey).

Genus ANOPLIUS Subgenus NOTIOCHARES Banks

Notiochares Banks, 1917. Mus. Compar. Zool., Bul. 61: 107, 108.
Type-species: Pompilus philadelphicus Lepeletier. Monotypic (=Sphex amethystina
Fabricius).
amethystinus amthystinus (Fabricius). Southern Fla., Ariz., Calif. south to Panama. Prey:

Lycosa sp. Another subsp. occurs in South America.

Sphex amethystina Fabricius, 1793. Ent. System., Emend. et Aucta, v. 2, p. 210. 9.

Pompilus anceps Cresson, 1865. Ent. Soc. Phila., Proc. 4: 130. ¢. Preoce.

Pompilus cubensis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 93. N. name.

Pompilus propinquus Fox, 1891. Amer. Ent. Soc., Trans. 18: 339. 2. Preoce.

Pompilus dux Dalla Torre, 1897. Cat. Hym., v. 8, p. 286. N. name.

Pompilus amethystinoides Strand, 1911. Arch. f. Naturges., v. 77, bd. 1, sup. 2, p. 147. N.
name for amethystinus Tasch., believed to differ from amethystinus F.

Psammochares philadelphicus var. floridensis Banks, 1917. Mus. Compar. Zool., Bul. 61:
106. @.

lepidus atramentarius (Dahlbom). Southern New England, Ohio and Nebr. to Fla. and Tex. in
Austroriparian and Carolinian Faunas; Mexico (Coahuila, Durango). Ecology: Nests in
sand and firm soil; visits many flowers. Prey: Lycosa rabida (Walck.), L. avida Walck.,

1554 Hymenoptera in America North of Mexico

L. antelucana Mont., L. sp., Arctosa littoralis (Hentz). Typical lepidus (Say) occurs from
Mexico to British Guiana and Eeuador.
Pompilus atramentarius Dahlbom, 1843. Hym. Europaea, v. 1, p. 48.
Pompilus philadelphicus Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 423. 9.
Pompilus philadelphicus var. sericatus Cresson, 1872. Amer. Ent. Soe.; Trans. 4: 202. °.
Sericopompilus plutonis Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 228. d.
Notiochares angusticeps Banks, 1939. Canad. Ent. 71: 227. °.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 13-15 (prey, nest). —Evans, 1951. Amer.
Ent. Soc., Trans. 76: 233-234 (prey). —Evans and Yoshimoto, 1955. Kans. Ent. Soc., Jour.
28: 18 (prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 71-72 (prey
transport, nest).

Genus ANOPLIUS Subgenus ANOPLIODES Banks

Anopliodes Banks, 1939. Canad. Ent. 71: 225, 226.
Type-species: Anopliodes modestus Banks. Monotypic © Anoplius bolli Banks).
bolli Banks. Tex. to Kans., Mo., and N. J.; Mexico (Tamaulipas).
Anoplius bolli Banks, 1917. Mus. Compar. Zool., Bul. 61: 104. d.
Anopliodes modestus Banks, 1939. Canad. Ent. 71: 226. 9.
parsonsi (Banks). South. Fla.; Cuba, Mexico south to Costa Rica.
Psammochares parsonsi Banks, 1944. Mus. Compar. Zool., Bul. 94: 183. ?.

Genus ANOPLIUS Subgenus ARACHNOPHROCTONUS Howard

Arachnophroctonus Howard, 1901. Insect Book, pl. VII, figs. 11, 14.
Type-species: Sphex tropica Fabricius. Desig. by Pate, 1946.
Arachnophila Ashmead, 1902. Canad. Ent. 34: 86, Preoce.
Type-species: Pompilus divisus Cresson. Orig. desig.
Anoplinellus Banks, 1934. Amer. Acad. Arts and Sci., Proc. 69: 84.
Type-species: Pompilus clotho Smith. Monotypice.
Arachnodaicter Pate, 1946. Amer. Ent. Soc., Trans. 72: 74. N. name.

Day (1974) synonymized the subgenus Pompilinus Ashm. under the subgenus Arachnophroc-
tonus How. This action was taken too late to be used in this edition of the catalog.

The wasps of this group inhabit sandy places, where they nest in short burrows in the earth,
stocking their burrows with errant spiders of several families. They visit flowers of many kinds.

Taxonomy: Day, 1974. Brit. Mus. (Nat. Hist.), Ent., Bul. 30: 379-380 (subgeneric syn.). —Day,
1974. Brit. Mus. (Nat. Hist.), Ent. Bul. 31: 49-51 (identity of Anoplinellus Bks.).
acapulcoensis (Cameron). L. Austr. Zone, Ariz. to Fla. and south. N. J., south to Guatemala.
Ecology: Visits flowers of many kinds.
Pompilus acapulcoensis Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 198. 6.
Psammochares bellicosus Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 224. 6.
Lophopompilus autilone Banks, 1919. Canad. Ent. 51: 82. d.

americanus ambiguus (Dahlbom). Tropical and L. Austr. Zones, Greater Antilles and Panama

to Calif., Utah, Kans., and Ala. Ecology: Nests in sand-clay soil. Parasite: Evagetes
mohave (Bks.). Prey: Arctosa littoralis (Hentz), Lycosa spp., Schizocosa crassipes
Walck.; Peucetia viridans Hentz.

Pompilus ambiguus Dahlbom, 1845. Hym. Europaea, sup. 1, p. 452. °.

Pompilus coruscus Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 156. 2.

Pompilus juxtus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 128. °.

Pompilus subargenteus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 129. 3.

Anoplius puella Banks, 1941. Canad. Ent. 73: 121. ¢.

Pompilinus orthodes Banks, 1944. Zool. 29: 112. 2.

Anoplius varunus Banks, 1947. Mus. Compar. Zool., Bul. 99: 419. 6.

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 76: 270 (prey transport, nest, parasite).
—Evans and Yoshimoto, 1955. Kans. Ent. Soe., Jour. 28: 18 (prey, nest). —Hurd and
Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey). —Evans and Yoshimoto, 1962. Ent.

Superfamily POMPILOIDEA 1555

Soe. Amer., Misc. Pub. 3: 76-77 (prey transport, nest). —Wasbauer and Powell, 1962. Kans.
Ent. Soc., Jour. 35: 397 (prey).
americanus americanus (Palisot de Beauvois). Cent. Tex. and north. Ga. to S. Dak., Minn.,

south Ont., and southeast. N. Y. Ecology: Nests in clay. Prey: Pardosa milvina (Hentz),
P. sp.; Philodromus sp.

Pompilus americanus Palisot de Beauvois, 1811. Ins. Afr. Amer., p. 117. 2.

Pompilus plebejus Dahlbom, 1845. Hym. Europaea, v. 1, p. 60. 2.

Psammochares albomarginatus Banks, 1910. N. Y. Ent. Soe., Jour. 18: 114. 6.

Pompiloides agnema Brimley, 1928. Elisha Mitchell Sci. Soe., Jour. 43: 204. 3.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 10-11 (prey transport, nest).
—Kurczewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 15 (prey).
americanus trifasciatus (Palisot de Beauvois). Austrorip. Fauna, N. C., Ga. and Fla. Prey:
Geolycosa hubbelli Wall.
Pompilus trifasciatus Palisot de Beauvois, 1811. Ins. Afr. Amer., p. 118. 2.
Psammochares eurydice Banks, 1921. Ent. Soc. Amer., Ann. 14: 19. 9.

Biology: Kurezewski, 1963. Fla. Ent. 46: 212 (prey).
apiculatus apiculatus (Smith). Tropical and L. Sonor. Zones, south Calif., Ariz., and cent. Tex.
to Panama. Ecology: Occurs along sandy streambanks and seashores.
Pompilus apiculatus Smith, 1855. Cat. Hym. Brit., Mus., v. 3, p. 157. 2.
Pompilus coruscus Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 155. 2. Preoce.
Arachnophroctonus apiculatus hondurensis Dreisbach, 1954. Amer. Midland Nat. 52: 439.
3.
apiculatus autumnalis (Banks). Austrorip. and Carol. Faunas, Tex. and Fla. to Ga., west. N.
Y., south. Ont., Minn., and Kans. Ecology: Nests in sand. Parasite: Evagetes mohave
(Bks.). Prey: Arctosa littoralis (Hentz).
Pompiloides autumnalis Banks, 1914. N. Y. Ent. Soc., Jour. 22: 301. 2.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 442, fig. 49 (larva).

Biology: Evans, Lin and Yoshimoto, 1953. N. Y. Ent. Soc., Jour. 61: 61-78 (prey transport,
nest, parasite, life history). —Evans and Yoshimoto, 1955. Kans. Ent. Soce., Jour. 28: 18
(prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 74-75 (prey transport,
nest, parasite). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 15 (prey).
—Kurczewski and Kurczewski, 1968. Kans. Ent. Soe., Jour. 41: 371 (prey). —Kurezewski
and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 68 (prey, nest).

apiculatus pretiosus (Banks). Coastal plain, Fla. to Maine. Ecology: Nests in sand, especially
along bodies of water. Parasite: Senotainia litoralis Allen. Prey: Arctosa littoralis
(Hentz), Lycosa helluo (Walck.).
Psammochares pretiosa Banks, 1910. N. Y. Ent. Soc., Jour. 18: 119. 6.
Pompiloides autumnalis var. atlanticus Banks, 1914. N. Y. Ent. Soc., Jour. 22: 301. °.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 442 (larva).

Biology: Krombein, 1953. Wasmann Jour. Biol. 10: 272-275 (prey, nest, life history, parasite).
—Krombein and Evans, 1954. Ent. Soc. Wash., Proc. 56: 230 (prey). —Krombein, 1959. Ent.
Soc. Wash., Proc. 61: 194 (prey). —Krombein, 1964. Amer. Mus. Novitates 2201: 11 (nest,
prey). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 15-16, fig. 3 (prey,
nest).

marginalis (Banks). Ariz., Tex., and Fla. to Utah, Man., south. Ont., and Mass. Ecology: Nests

in sand. Parasite: Evagetes mohave (Bks.). Prey: Geolycosa pikei (Marx), G. wrighti Em.,
G. missouriensis Bks., G. spp., Lycosa baltimoriana Keys., L. carolinensis Walck., L.
frondicola Em., L. avida Walck., L. sp. near lenta Hentz, L. sp. near ceratiola Gertsch
and Wall., L. spp., Arctosa littoralis Hentz; Pellenes coronatus (Hentz).

Sphex tropica Fabricius, 1775. Systema Ent., p. 350. 2. Preoce.

Psammochares marginalis Banks, 1910 (June). N. Y. Ent. Soc., Jour. 18: 118. 2, 6.

Psammochares castella Banks, 1910 (Dec.). Psyche 17: 248. 6.

Psammochares fabricii Banks, 1933. Psyche 40: 6. N. name.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 441, figs. 31-34, 48 (larva).

1556 Hymenoptera in America North of Mexico

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 45-58 (prey, nest). —Krombein, 1953.
Wasmann Jour. Biol. 10: 275-276 (prey, nest, life history). —Krombein, 1953. Ent. Soe.
Wash., Proc. 55: 116-117 (prey, nest). —Evans and Yoshimoto, 1955. Kans. Ent. Soc., Jour.
28: 18 (prey, nest). —Krombein, 1958. Ent. Soc. Wash., Proe. 60: 102-103 (prey, nest).
_Evans and Yoshimoto, 1962. Ent. Soe. Amer., Misc. Pub. 3: 73-74 (nest, prey, parasite).
—Wasbauer and Powell, 1962. Kans. Ent. Soe., Jour. 35: 397 (prey). —Kurczewski, 1963
(1962). Brooklyn Ent. Soe., Bul. 57: 88-89 (prey). —Krombein, 1964. Amer. Mus. Novitates
2201: 11-13 (prey, nest). —Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 68
(prey, nest, parasite).

moestus (Banks). L. Sonor. Fauna, east. Tex. to southeast. Ariz.; Mexico.
Pompiloides moestus Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 226. °.
Anoplius comanche Banks, 1941. Canad. Ent. 73: 121. 6.
occidentalis (Dreisbach). Calif. (Redwood City).
Arachnophroctonus occidentalis Dreisbach, 1954. Amer. Midland Nat. 52: 437, figs. 13, 17.
Bo
relativus (Fox). Mass., Ont. and B. C. to Fla., Tex. and Calif., mostly in U. and L. Austral
Zones, south in Mexico at higher altitudes to Oaxaca and Guerrero. Ecology: Occurs in
fields and prairies; nests in sand and heavier soil. Prey: Lycosa rabida Walck., L.
antelucana Mont., Arctosa littoralis (Hentz), Geolycosa missouriensis (Bks.), G. sp.;
Agelenopsis naevia (Walck.), A. pennsylvanica (Koch).
Pompilus relativus Fox, 1893. Canad. Ent. 25: 114. °.
Psammochares (Allocyphonyx) hesione Banks, 1910. Psyche 17: 250. d.
Psammochares difficilis Banks, 1917. Mus. Compar. Zool., Bul. 61: 105. °.
Anoplius confraternus Banks, 1926. Canad. Ent. 58: 201. °.
Psammochares henshawi Banks, 1939. Canad. Ent. 71: 226. °. -
Arachnoproctonus (!) variegatus Dreisbach, 1957. Ent. News 68: 72, 2 figs. d.

Biology: Evans, 1951. Amer. Ent. Soe., Trans. 76: 251-252 (nest, prey). —Evans and
Yoshimoto, 1955. Kans. Ent. Soc., Jour. 28: 18 (nest, prey). —Evans and Yoshimoto, 1962.
Ent. Soc. Amer., Misc. Pub. 3: 72-73 (prey, nest). —Kurezewski and Kurczewski, 1968.
Kans. Ent. Soc., Jour. 41: 14-15 (prey, nest). —Kurezewski and Kurczewski, 1968. Kans.
Ent. Soe., Jour. 41: 371-372 (prey hunting). —Kurezewski and Kurezewski, 1973. Kans. Ent.
Soe., Jour. 46: 68-69 (prey, nest).

semirufus (Cresson). Tex. and Fla. to Mont., Man., and N. B. Ecology: Nests in sand. Prey:

Lycosa gulosa Walck., L. rabida Walck., L. punctulata Hentz, L. avida Walck., L. spp.,
Schizocosa crassipalpis (Em.), S. crassipes (Walck.), S. saltatrix (Hentz), S. bilineata
(Em.), S. ocreata (Hentz), S. spp., Pardosa milvina Hentz, P. falcifera Camb., P. sp. near
modica (Blackw.), P. distincta (Blackw.), P. ap. near floridana Bks., P. spp., Trochosa
pratensis (Em.), T. avara Keys.; Agelenopsis pennsylvanica (Koch), A. sp.; Amaurobius
sp. ;

Pompilus semirufus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 100. °.

Pompilus divisus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 100. 2.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 442 (larva).

Biology: Krombein, 1953. Wasmann Jour. Biol. 10: 277-280 (prey, nest, life history). —Evans,
1953. Syst. Zool. 2: 168-169 (nest, prey). —Evans and Yoshimoto, 1955. Kans. Ent. Soc.,
Jour. 28: 18 (prey, nest). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 103 (prey, nest).
—Krombein, 1959. Ent. Soc. Wash., Proc. 61: 194 (prey). —Evans and Yoshimoto, 1962.
Ent. Soc. Amer., Misc. Pub. 3: 75-76 (prey, nest, life history). —Kurezewski, 1963. Brooklyn
Ent. Soc., Bul. 57: 89 (prey transport). —Evans, 1964. Kans. Ent. Soc., Jour. 37: 306 (prey
transport). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 16-18 (prey).
—Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 372 (prey). —Kurezewski
and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 69 (nest, prey).

xerophilus Evans. L. Sonor. Fauna, Tex., N. Mex., to cent. Calif. and Utah; Mexico (Baja
California).
Anoplius xerophilus Evans, 1947. Ent. News 58: 10. 2, do.
Arachnophroctonus gaigei Dreisbach, 1954. Amer. Midland Nat. 52: 488, figs. 4, 11. d.

Superfamily POMPILOIDEA 1557

Genus ANOPLIUS Subgenus POMPILINUS Ashmead

Psammochares Latreille, 1796. Precis Caract. Gen. Ins., p. 115. Suppressed by Internatl.
Comn. Zool. Nomencl., Op. 166, 1945.
Pompilinus Ashmead, 1902. Canad. Ent. 34: 85.
Type-species: Pompilus cylindricus Cresson. Orig. desig.
Anopliella Banks, 1939. Canad. Ent. 71: 225, 227.
Type-species: Pompilus tenebrosus Cresson. Orig. desig.

Day (1974) synonymized the subgenus Pompilinus Ashm. under the subgenus Arachnophroc-
tonus How. This action was published too late to be used in this edition of the catalog.

Wasps of this group inhabit gardens, waste places, clay banks, and sandy areas; they prey
upon a wide variety of spiders and nest in short galleries in the earth.

Revision: Dreisbach, 1949. Ent. Amer. (n. s.) 29: 12-58.

Taxonomy: Day, 1974. Brit. Mus. (Nat. Hist.), Ent., Bul. 30: 379-380 (subgeneric syn.).
californiae Evans. L. Sonor. Fauna, N. Mex. to Calif.; Mexico (Sonora, Baja California).
Ecology: Has been taken on Helianthus and Tamarix.
Anoplius (Pompilinus) californiae Evans, 1948. Pan-Pacifie Ent. 24: 128. 2, ¢.
clystera (Banks). L. and U. Sonor. Fauna, Tex. and south. Calif. to Colo. and cent. Calif.;
Mexico (Chihuahua, Baja California). Prey: Lycosa sp.
Pompiloides clystera Banks, 1914. N. Y. Ent. Soe., Jour. 22: 302. 6.
Pompilinus submarginatus Dreisbach, 1952. Amer. Midland Nat. 48: 146, figs. 14, 17. ¢.

Taxonomy: Evans, 1968. Ent. News 79: 254-255, fig. 4 (redescription female).

Biology: Wasbauer and Powell, 1962. Pan-Pacific Ent. 35: 398 (prey).

cylindricus (Cresson). Conn., south. Ont., N. W. T., cent. Oreg. s. to Fla.,Tex. and Calif.; Mexico
(Chihuahua). Ecology: Nests in sandy places. Prey: Geolycosa wrightii (Em.), G. sp.
Pompilus cylindricus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 92. ¢.
Arachnophila brevihirta Banks, 1945. Psyche 52: 105. 2.
Pompilinus truncatus Dreisbach, 1949. Ent. Amer. (n. s.) 29: 15. 3.
Pompilinus subtruncatus Dreisbach, 1949. Ent. Amer. (n. s.) 29: 17. 6.
Pompilinus hispidus Dreisbach, 1949. Ent. Amer. (n. s.) 29: 23. ¢.
Pompilinus clavipes Dreisbach, 1958. Ent. News 69: 61, figs. d.

Biology: Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 19 (prey).
—Kurczewski and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 69 (prey).
estellina (Banks). U. Sonor. and Transit. Faunas, Calif. and N. Mex. to Alta., s. to Guatemala.
Ecology: Nests in pre-existing crevices or burrows in soil. Prey: Lycosidae sp. ?
Pompiloides estellina Banks, 1914. N. Y. Ent. Soc., Jour. 22: 303. 3d, (2 misdet.).
Pompilinus utahensis Dreisbach, 1952. Amer. Midland Nat. 48: 145, figs. 15, 16. ¢.
Pompilinus minutus Dreisbach, 1952. Amer. Midland Nat. 48: 147, figs. 7, 8. d.

Biology: Evans, 1964. Kans. Ent. Soc., Jour. 37: 305-306 (prey, nest).
fraternus (Banks). L. and U. Austr. Zones, N. Mex. and Fla. to Colo., Nebr., and southeast. N.

Y.; Mexico (Tamaulipas, Sonora). Ecology: Nests on protected beaches and on elevated
salt flats inland in pre-existing crevices or burrows. Parasite: Ephuta s. sabaliana
Schus. Prey: Lycosa watsoni Gertsch, L. carrana Bryant, Sosippus floridanus Sim.,
Pardosa longispinnata Tull.; Habronattus sp., Pellenes sp.

Lophopompilus fraternus Banks, 1941. Canad. Ent. 73: 120. 2.

Pompilinus dowi Dreisbach, 1949. Ent. Amer. (n. s.) 29: 12. d.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 440, figs. 35-37 (larva).

Biology: Krombein and Evans, 1954. Ent. Soc. Wash., Proc. 56: 231 (prey transport, nest).
—Evans, Krombein, and Yoshimoto, 1955. Brooklyn Ent. Soc., Bul. 50: 77-84, 3 figs.
(hunting behavior, prey, nest, life history).

insolens (Banks). Transcont. in Transit. and U. Austr. Zones, Maine, Ont. and B. C. s. to Ga.,
Tex. and Calif., s. in Mexican central plateau to Veracruz and Morelos. Prey: Maevia
vittata (Hentz).
Pompiloides insolens Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 226. 3.

1558 Hymenoptera in America North of Mexico

krombeini Evans. Austrorip. Fauna, Fla. to N. J. Ecology: Nests in sand. Prey: Pardosa

floridana Bks., Schizocosa sp.
Anoplius (Pompilinus) krombeini Evans, 1950. Kans. Ent. Soce., Jour. 23: 88. , d.

Taxonomy: Evans, 1959. Ent. Soe. Amer., Ann. 52: 440-441 (larva).

Biology: Krombein, 1953. Ent. Soc. Wash., Proce. 55: 117-118 (prey, nest). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 80 (prey transport, nest, life history).
—Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 69 (prey).

leona (Cameron). West. Tex. to Ariz., south to El Salvador.
Pompilus leona Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 198, pl. 11, fig. 20. “3” =

OF

Anoplius (Pompilinus) grandiflexionis Evans, 1950. Kans. Ent. Soc., Jour. 23: 84. 9, d.

marginatus (Say). L. Austr. to Canad. Zone, Ariz., Colo., and Alta. east to the Atlantic.

Ecology: Nests in sand, gravel and other light soils. Parasite: Evagetes parvus (Cr.) ?
Prey: Odiellus pictus (Wood); Argiope aurantia Luc.; Agelenopsis naevia (Walck.), A.
pennsylvanica (Koch), Pellenes borealis (Bks.), P. viridipes (Hentz), P. sp., Arctosa
emertoni Gertsch, Geolycosa fatifera Comst., Lycosa avida Walck., L. domifex Hancock;
Pardosa saxatilis (Hentz), P. sp., Pirata insularis Em., Schizocosa crassipes Walck., S.
sp., Trochosa avara (Keys.), T. pratensis (Em.), T. sp.; Callilepis imbecillus (Keys.),
Drassodes neglectus (Keys.), Drassylus niger (Bks.), Gnaphosa muscorum (Koch), G. sp.,
Haplodrassus signifer (Koch), Herpyllus vasifer (Walck.); Castianeira longipalpus
(Hentz), Clubiona kastoni Gertsch, C. mixta Em., C. sp.; Anyphaena sp.; Philodromus
infuscatus Keys., P. washita Bks., Thanatus formicinus (Oliv.), Tibellus duttoni (Hentz),
Xysticus bicuspis Keys., X. elegans Keys., X. ferox (Hentz), X. funestus Keys.; Evarcha
hoyi (Peckh.), Habrocestum decorus (Blackw.), Maevia vittata (Hentz), Phidippus audax
(Hentz), P. clarus Keys., P. purpuratus Keys.; Amaurobius bennetti (Blackw.). Females
of bequaerti (Dreis.), rectangularis (Dreis.) and townesi Evans are unknown and are
probably confused with females of marginatus; accordingly, some of the prey records
and biological references may refer to one of the former three taxa.

Pompilus marginatus Say, 1824. In Keating’s, Narr. Long’s 2nd Exped., v. 2, p. 333. °.

Pompilus (Miscus) petiolatus Say, 1835. Boston Jour. Nat. Hist. 1: 305. 2.

Pompiloides reducta Banks, 1914. N. Y. Ent. Soe., Jour. 22: 302. 2.

Pompiloides hageni Banks, 1919. Mus. Compar. Zool., Bul. 63: 235. °.

Pompilinus basirufus Dreisbach, 1952. Amer. Midland Nat. 48: 149, figs. 9, 10. d (°
misdet.).

Pompilinus basirufous (!) Dreisbach, 1952. Amer. Midland Nat. 48: 158.

Taxonomy: Evans, 1959. Ent. Soe. Amer., Ann. 52: 440, fig. 39 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 144-152 (prey
transport, nest). —Hartman, 1905. Tex. Univ., Bul. 65: 52-54 (prey, nest). —Rau and Rau,
1918. Wasp Studies Afield, pp. 58-63 (prey hunting, transport). —Evans, 1948. Ent. News
59: 183-184 (prey, nest). —Evans, 1950. Amer. Ent. Soc., Trans. 75: 164 (parasite).
—Krombein, 1953. Wasmann Jour. Biol. 10: 280-281 (prey, nest). —Evans and Yoshimoto,
1955. Kans. Ent. Soce., Jour. 28: 19 (prey). —Krombein, 1959. Ent. Soc. Wash., Proc. 61: 194
(prey). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 78-80 (prey transport,
nest, life history). —Kurezewski, 1963 (1962). Brooklyn Ent. Soc., Bul. 57: 89 (prey
transport). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 19-20 (prey).
—Kurczewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 373-374 (prey).
—Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 69-70, fig. 1 (prey, nest).

percitus Evans. Austrorip. and Carol. Faunas, Tex. to Mass. Prey: Trochosa sp. probably
gosiuta Chamb., immature.
Anoplius (Pompilinus) percitus Evans, 1950. Kans. Ent. Soc., Jour. 23: 85. 2, d.
rectangularis gillaspyi Evans. Tex., N. C., N. J. in Austrorip. Fauna.
Anoplius (Pompilinus) rectangularis gillaspyi Evans, 1951. Amer. Ent. Soc., Trans. 76:
310, fig. 101. 3.
rectangularis rectangularis (Dreisbach). Transit. and U. Austr. Zones, Mass. to Minn., s. to
Colo., Kans., Ohio and N. C. Prey: Lycosa avida Walck., Schizocosa sp., Pardosa

Superfamily POMPILOIDEA 1559

distincta (Blackw.); Xysticus elegans Keys., X. ferox (Hentz); Phidippus whitmanii
Peckh.
Pompilinus rectangularis Dreisbach, 1949. Ent. Amer. (n. s.) 29: 18. d.

Biology: Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 20-21 (prey).
—Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 374 (prey). —Kurezewski
and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 70 (prey).

splendens (Dreisbach). L. and U. Austr. Zones, Fla., Tex., and Ariz. to Alta., Man., Ont., and

Maine; Mexico (Chihuahua, Durango). Ecology: Nests in sandy areas. Prey: Agelenopsis
pennsylvanica (Koch), A. spp.; Araneus patagiatus Clerck; Pisaurina mira (Walck.);
Amaurobius bennetti (Blackw.); Pardosa distincta (Blackw.), P. milvina (Hentz), P. sp.
near floridana Bks., Schizocosa crassipes (Walck.), S. saltatrix (Hentz), S. sp., Trochosa
avara Keys., Arctosa littoralis (Hentz), Geolycosa wrightii Em., Lycosa avida Walck., L.
frondicola Em., L. sp.; Xysticus pellax Camb., X. ferox (Hentz), X. funestus Keys., X.
elegans Keys., X. sp.; Thanatus formicinus (Clerck); Marpissa pikei (Peckh. and
Peckh.), Pellenes coronatus (Hentz), P. agilis Bks., P. sp., Phidippus audax (Hentz), P.
clarus Keys., Habronattus viridipes (Hentz); Castianeira longipalpus (Hentz), C. sp.

Pompilinus splendens Dreisbach, 1949. Ent. Amer. (n. s.) 29: 20. 6.

Pompilinus pseudoreductus Dreisbach, 1949. Ent. Amer. (n. s.) 29: 21. d.

Pompilinus ohioensis Dreisbach, 1949. Ent. Amer. (n. s.) 29: 24. 3.

Pompilinus shappirioi Dreisbach, 1952. Amer. Midland Nat. 48: 148, figs. 5, 6. ¢. N. syn.
(H. E. Evans).

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 76: 320 (prey). —Krombein, 1959. Ent. Soc.
Wash., Proc. 61: 194 (prey). —Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 398
(prey). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 21-22 (prey).
—Kurczewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 374-375 (prey).
—Kurezewski and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 70 (prey).

stenotus bequaerti (Dreisbach). Tex. and Fla. to Alta., Minn., Mich., and N. C. Ecology: Nests
in sand. Prey: Lycosa sp.
Pompilinus bequaerti Dreisbach, 1949. Ent. Amer. (n. s.). 29: 16. ¢.

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 13 (prey transport).
stenotus stenotus (Banks). South. Fla. and Ala. Ecology: Nests in sand. Prey: Lycosa sp.,
Trochosa sp.
Pompiloides stenotus Banks, 1914. N. Y. Ent. Soc., Jour. 22: 302. ¢.

Biology: Krombein and Evans, 1955. Ent. Soc. Wash., Proc. 57: 230 (prey, nest).
—Kurezewski and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 70 (prey).
subcylindricus (Banks). L. Austr. to Transit. Zones, Fla., Tex., and Ariz. to Utah, Minn., Que.,
and Vt.; Mexico to Morelos and Guerrero. Ecology: Nests in sand and gravelly soil.
Parasite: Evagetes parvus (Cr.). Prey: Xysticus gulosus Keys., X. banksi Bry., X. ferox
(Hentz), X. triguttatus Keys., X. funestus Keys.; Micaria sp.
Pompiloides subcylindricus Banks, 1917. Mus. Compar. Zool., Bul. 61: 103. 9, d.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 440 (larva).

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 76: 288 (prey). —Evans, 1951. Amer. Ent. Soc.,
Trans. 77: 315 (prey). —Krombein, 1953. Wasmann Jour. Biol. 10: 281 (prey). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 77-78 (prey transport, nest, life history,
parasite). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 18-19 (prey).

tenebrosus (Cresson). Transcont. in Canad. and Transit. Zones, entering U. Austr. Ecology:

Nests in sand. Prey: Schizocosa saltatrix Hentz, Trochosa avara Keys., Tarentula kochi
Keys., Lycosa frondicola Em., L. avida Walek., L. baltimoriana Keys.; Xysticus gulosus
Keys., X. ferox Hentz, Thanatus formicinus (Oliv.).

Pompilus tenebrosus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 453. °.

Pompilus compactus Provancher, 1895. Nat. Canad. 22: 111. °.

Pompiloides canadensis Banks, 1919. Canad. Ent. 51: 82. d.

Pompilinus drakei Dreisbach, 1958. Ent. News 69: 62, 2 figs. d. N. syn (H. E. Evans). The
unique holotype is a teratological specimen.

1560 Hymenoptera in America North of Mexico

Biology: Evans, 1951. Amer. Ent. Soe., Trans. 76: 303 (prey). —Evans and Yoshimoto, 1962.
Ent. Soc. Amer., Mise. Pub. 3: 81 (prey). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 482
(prey). —Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 399 (prey). —Kurezewski
and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 70-72 (prey transport, nest).

texanus (Dreisbach). La. and Tex. s. in Mexico to Morelos and Yucatan.
Pompilinus texanus Dreisbach, 1949. Ent. Amer. (n. s.) 29: 14. 6.
townesi Evans. Alleghan. Fauna, N. H. w. to Man., Conn. to Fla.
Anoplius (Pompilinus) townesi Evans, 1951. Amer. Ent. Soc., Trans. 76: 318, fig. 102. d.

Genus ANOPLIUS Subgenus ANOPLIUS Dufour

Anoplius Dufour, 1834. Soe. Ent. France, Ann. 2: 483.
Type-species: Sphex nigerrimus Scopoli. Desig. by Internatl. Comn. Zool. Nomencl.,
Op. 997, 1973.
Pompilioides Radoszkowski, 1887. Soc. Ent. Rossica, Horae 21: 94.
Type-species: Pompilioides unicolor Radoszkowski. Desig. by Ashmead, 1902.
Aphiloctenus Ashmead, 1902. Canad. Ent. 34: 87.
Type-species: Pompilus virginiensis Cresson. Orig. desig.

Wasps of this subgenus nest in ready-made cavities of various kinds, and occupy a variety of
habitats.

depressipes Banks. Carol. and Alleghanian Faunas, Tex., Ala. and Fla. to Wis. and Maine.
Ecology: Nests in pre-existing cavities in wood and soil; occasionally stores more than
one cell per cavity, each cell separated by, a partition of the substrate. Prey: Dolomedes
triton sexpunctatus Hentz, D. scriptus Hentz, D. striatus Giebel, D. tenebrosus Hentz.
Anoplius depressipes Banks, 1919. Canad. Ent. 51: 81. 2.

Biology: Evans, 1949. Ent. Soe. Wash., Proc. 51: 206 (prey transport). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 83-84 (prey transport, nest). —Kurezewski
and Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 22 (prey).

dreisbachi Evans. Ariz. and south. Calif. to Colo., Wyo. and B. C., chiefly in Transit. Zone.
Anoplius (Anoplius) dreisbachi Evans, 1966. Amer. Ent. Soc., Mem. 20: 353, fig. 49. gd, 2.
elongatus Dreisbach. Mich., Minn.
Anoplius elongatus Dreisbach, 1950. Amer. Midland Nat. 43: 576, figs. 11, 12. d.
fulgidus (Cresson). Extreme southern U. S., Fla., Tex., Ariz., Utah, Calif.; Antilles, Mexico,
Central and South America to Peru and Brazil. Ecology: Nests in heavy loam along
stream. Prey: Pirata sedentarius Montg., Arctosa sp. near littoralis (Hentz).
Pompilus fulgidus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 181. 9.
Pompilus aeneopurpureus Fox, 1891. Amer. Ent. Soc., Trans. 18: 339. ?, d.
Pompilus championi Cameron, 1893. Biol. Cent.-Amer. Hym., v. 2, p. 196. °.
Pompilus mundulus Fox, 1897. Acad. Nat. Sci. Phila., Proc. 49: 243. °.
Anoplius amarus Banks, 1947. Mus. Compar. Zool., Bul. 99: 416. °.

Biology: Wasbauer, 1955. Pan-Pacific Ent. 31: 63-66 (prey transport). —Kurcezewski and

Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 22 (prey).
hispidulus Dreisbach. IIl., Mich., N. Y., Conn., Md.

Anoplius hispidulus Dreisbach, 1950. Amer. Midland Nat. 43: 577, figs. 25, 26. d.
illinoensis (Robertson). U. Austr. and Transit. Zones, Ga., Tex., Utah, and Colo. to Mont.,

re and Que. Ecology: Nests in firm soil. Prey: Lycosa avida Walck., L. helluo Walck.,
. Sp.
Beep illinoensis Robertson, 1901. Amer. Ent. Soc., Trans. 27: 202. 2, ¢.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 489, figs. 20-25, 47 (larva).

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Misc. Pub. 3: 82-83 (prey transport,
nest). —Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 72 (prey).
imbellis Banks. Transcont. in U. Austr. and Transit. Zones south at higher altitudes to Costa
Rica. Ecology: Nests in soil near still water. Prey: Pardosa ramulosa McCook, P.
milvina (Hentz), P. distincta group, P. sp., Trochosa avara Keys., Arctosa sp.
Anoplius imbellis Banks, 1944. Mus. Compar. Zool., Bul. 94: 169. 2, 6.

Superfamily POMPILOIDEA 1561

Anoplius imbellis var. major Dreisbach, 1950. Amer. Midland Nat. 43: 581, figs. 20, 24. d.
Anoplius subimbellis Dreisbach, 1952. Amer. Midland Nat. 48: 155, figs. 23, 25. ¢.

Biology: Wasbauer, 1957. Wasmann Jour. Biol. 15: 81-97, 7 figs. (prey, nests). —Wasbauer
and Powell, 1962. Kans. Ent. Soc., Jour. 35: 399 (prey). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 22 (prey). —Kurezewski and Kurezewski, 1973. Kans. Ent.
Soc., Jour. 46: 72 (prey).

ithaca (Banks). Transcont. in Transit. and U. Austr. Zones; Mexico (Nuevo Leon). Ecology:
Occurs along rocky stream-beds and nests under stones. Prey: Pardosa lapidicina Em.,
P. milvina (Hentz), P. groenlandica Th., P. steva Lowrie and Gertsch, P. sp., Arctosa
littoralis (Hentz), Lycosa sp.
Psammochares ithaca Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 224. 9.
Anoplius selkirkensis Banks, 1919. Mus. Compar. Zool., Bul. 63: 234. 2, d.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 440 (larva).

Biology: Evans, 1948. Ent. News 59: 180-183 (prey transport, nest). —Evans and Yoshimoto,
1955. Kans. Ent. Soc., Jour. 28: 19 (prey). —Krombein, 1956. Brooklyn Ent. Soc., Bul. 51: 42
(prey transport). —Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 83 (prey
transport, nest). —Kurezewski, 1963 (1962). Brooklyn Ent. Soc., Bul. 57: 89 (prey hunting
and transport). —Kurezewski and Kurcezewski, 1968. Kans. Ent. Soc., Jour. 41: 22 (prey).
—Ricards, 1969. Ent. News 80: 149-157, 2 figs. (prey transport, nesting behavior).
—Kurczewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 72 (prey).

nigerrimus (Scopoli). Transcont. in Hudsonian and Canadian Zones, entering Transition Zone

rarely; Holarctic. Ecology: Not known for North America; in Europe it nests in
pre-existing cavities in twigs or soil, or under stones. Prey: Not known for North
America; in Europe it preys chiefly on Lycosidae, occasionally on Pisauridae and
Gnaphosidae.

Sphex nigerrima Scopoli, 1763. Ent. Carniolica, p. 295. 2 ?

Pompilus incisus Tischbein, 1850. Stettin. Ent. Ztg. 11: 8. d.

Pompilus excerptus Tournier, 1889. Ent. Genevois 1: 159. 9.

Pompilus nigerrimus var. Kohli Verhoeff, 1892. Ent. Nachr. 18: 71. ¢.

Anoplius wheeleri Banks, 1939. Canad. Ent. 71: 228. 2, 6.

Anoplius banksi Dreisbach, 1950. Amer. Midland Nat. 43: 579, figs. 13, 14. 3.

Taxonomy: van der Vecht and Menke, 1968. Bul. Zool. Nomencl. 25: 120-124 (request for
designation of nigerrimus as type-species of Anoplius, and of a neotype for nigerrimus;
request approved in Op. 997, Internatl. Comn. Zool. Nomencl., 1973).

Biology: Richards and Hamm, 1939. Soc. Brit. Ent., Trans. 6: 98-100 (summarizes European *
literature on prey, nests).
papago Banks. D. C. to Kans., s. to Fla. and Ariz. and Costa Rica. Ecology: Nests in loamy soil.
Prey: Lycosa sp.
Anoplius papago Banks, 1941. Canad. Ent. 73: 120. d.
Anoplius subtarsatus Dreisbach, 1950. Amer. Midland Nat. 43: 578, figs. 18, 22. 3.
Anoplius guatemalensis Dreisbach, 1952. Amer. Midland Nat. 48: 154. d.

Biology: Evans, 1964. Kans. Ent. Soc., Jour. 37: 304 (prey, nest).

tenuicornis (Tournier). Transcont. in Canad. Zone, s. to N. B., Vt., Man. and in mts. to N. Mex.
and Calif.; Holarctic.
Pompilus tenuicornis Tournier, 1889. Ent. Genevois 1: 159. 9.
Pompilus piliventris Morawitz, 1889 Soc. Ent. Rossica, Horae. 23: 122. 6.
Anoplius basalis Dreisbach, 1950. Amer. Midland Nat. 43: 578, figs. 9, 10. 6.
toluca (Cameron). Calif. to west. Tex., s. to Costa Rica; adventive in Hawaii. Ecology: Nests in
soil. Prey: Alopecosa gertschii Schenkel, Lycosa sp., Schizocosa sp. in avida group,
Tarentula kochii Keys., Trochosa gosiuta Chamb., T. pratensis (Em.); adults.
Pompilus toluca Cameron, 1893. Biol. Cent.-Amer., Hym. 2: 195. 2.
Anoplius tarsatus Banks, 1919. Mus. Compar. Zool., Bul. 63: 233. °.

Biology: Evans, 1964. Kans. Ent. Soce., Jour. 37: 305 (prey).

1562 Hymenoptera in America North of Mexico

ventralis (Banks). Austrorip. to Alleghanian Fauna, Tex. and Fla. to Man., Ont., and N.S.
Ecology: Nests in sand and in hard-packed soil Prey: Agelenopsis naevia (Walck.);
Lycosa helluo Walck., Schizocosa saltatrix (Hentz), Trochosa frondicola (Em.).

Psammochares ventralis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 120. 6.

Biology: Wasbauer and Powell, 1962. Kans. Ent. Soe., Jour. 35: 399 (prey). —Kurezewski and
Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 23 (prey). —Kurezewski and Kurezewski, 1968.
Kans. Ent. Soc., Jour. 41: 375-376 (prey transport, nest).

virginiensis (Cresson). N. S., Sask., and B. C., s. to Wash., Colo., Mo., and Ga. Ecology: Occurs
in open woods; nests in pre-existing cavities in dead wood. Prey: Agelenopsis
pennsylvanica (Koch), A. utahana (Chamb. and Ivie), Wadotes calcaratus (Keys.), W.
hybridus (Em.), Coras juvenilis (Keys.); Amaurobius bennetti (Blackw.); Pardosa
lapidicina Em.
Pompilus virginiensis Cresson, 1867. Amer. Ent. Soc., Trans. 1: 92. d.

Taxonomy: Evans, 1959. Ent. Soc. Amer., Ann. 52: 440 (larva).

Biology: Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 81-82 (prey hunting and
transport, nest, life history). —Kurezewski and Kurezewski, 1968. Kans. Ent. Soce., Jour. 41:
22 (prey transport). —Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46: 72

(prey).
Genus POMPILUS Fabricius

The species of this genus prey upon errant spiders, principally of the families Lycosidae, Sal-
ticidae, and Pisauridae; they nest in sand or soft earth so far as is known.

Genus POMPILUS Subgenus POMPILUS Fabricius

Pompilus Fabricius, 1798. Sup. Ent. System., p. 212.
Type-species: Pompilus pulcher Fabricius. Desig. by Internatl. Comn. Zool.
Nomencl., Op. 166, 1945.

Members of the typical subgenus occur only in the Old World.
Genus POMPILUS Subgenus HESPEROPOMPILUS Evans

Pompilus subg. Hesperopompilus Evans, 1948. Ent. Soc. Wash., Proc. 50: 141.
Type-species: Pompilus orophilus Evans. Orig. desig.

The seven known species of this subgenus occur in western North America south into Baja
California and Sonora.

Revision: Evans, 1966. Amer. Ent. Soc., Mem. 20: 358-370.
hilli Evans. Calif. (Pomona Mts.).
Pompilus (Hesperopompilus) hilli Evans, 1957. Pan-Pacifie Ent. 33: 183. d.
idahoensis Evans. Idaho (Gooding).
Pompilus (Hesperopompilus) idahoensis Evans, 1966. Amer. Ent. Soc., Mem. 20: 367. °.

jacintoensis Evans. South. Calif. and Ariz.; Mexico (Baja California, Sonora).
Pompilus (Hesperopompilus) jacintoensis Evans, 1948. Ent. Soc. Wash., Proc. 50: 146. 2
(3d misdet.).
Pompilus (Hesperopompilus) evagetoides Evans, 1951. Amer. Ent. Soc., Trans. 77: 217. ¢.
orophilus Evans. Transit. Fauna, Alta. and Nebr. to N. Mex. and Calif.; Mexico (Sonora).
Pompilus orophilus Evans, 1947. Ent. News 58: 14. °. :
rufopictus Evans. South. Calif. and Ariz. in deserts.
Pompilus (Hesperopompilus) rufopictus Evans, 1948. Ent. Soc. Wash., Proc. 50: 144. 2.
Pompilus (Hesperopompilus) boharti Evans. 1951. Amer. Ent. Soc., Trans. 77: 216, figs.
176, 204. ¢.
serrano Evans. South. Calif.; Mexico (Baja California).
Pompilus (Hesperopompilus) serrano Evans, 1966. Amer Ent. Soc., Mem. 20: 369, figs. 58,
Sarie sl.

Superfamily POMPILOIDEA 1563

Genus POMPILUS Subgenus XENOPOMPILUS Evans

Pompilus subg. Xenopompilus Evans, 1953. Ent. Soc. Amer., Ann. 46: 531.
Type-species: Pompilus (Xenopompilus) tlahuicanus Evans. Orig. desig.
tlahuicanus Evans. Ariz. (Portal): Mexico (Puebla, Morelos, Jalisco, Durango, Chihuahua).
Pompilus (Xenopompilus) tlahuicanus Evans, 1953. Ent. Soc. Amer., Ann. 46: 534, fig. 1.
ORROr
Pompilus (Xenopompilus) tarahumarae Evans, 1953. Ent. Soc. Amer., Ann. 46: 535. °.

Genus POMPILUS Subgenus PERISSOPOMPILUS Evans

Pompilus subg. Perissopompilus Evans, 1951. Amer. Ent. Soc., Trans. 77: 222.
Type-species: Pompilus (Ammosphex) phoenix Evans. Orig. desig.
perfasciatus Evans. South. Calif. and Ariz. in deserts; Mexico (Zacatecas).
Pompilus (Perissopompilus) perfasciatus Evans, 1951. Amer. Ent. Soc., Trans. 77: 225. ©

Taxonomy: Evans, 1958. Ent. News 69: 147, figs. 1-2. 3.

phoenix Evans. Calif., Nev., Utah, west. Tex.; Mexico (Baja California and Sonora s. to Puebla
and Guerrero). Ecology: Occurs in open, semiarid areas. Prey: Filistata sp.
Pompilus (Ammosphex) phoenix Evans, 1948. Pan-Pacifie Ent. 24: 123. d, 2

Biology: Evans, 1966. Ent. Soc. Wash., Proc. 68: 339 (prey).

Genus POMPILUS Subgenus XEROCHARES Evans

Pompilus subg. Xerochares Evans, 1951. Amer. Ent. Soc., Trans. 77: 218.
Type-species: Pompilus connexus Fox. Orig. desig.
expulsus Schulz. South. Ariz.; Mexico to Nicaragua.
Pompilus connexus Fox, 1893. Calif. Acad. Sci., Proc. (2) 4: 23. 2. Preoce.
Pompilus expulsus Schulz, 1906. Spolia Hym., p. 170. N. name.
Psammochares arizonica Banks, 1910. N. Y. Ent. Soc., Jour. 18: 115. °.
Pompilus rubriventris Bradley, 1944. Notulae Nat. 145, p. 9. N. name.

Genus POMPILUS Subgenus AMMOSPHEX Wilcke

Ammosphex Wilcke, 1942. Ent. Ber. 11: 25.
Type-species: Pompilus unguicularis Thomson. Orig. desig.
Anopompilinus Dreisbach, 1949. Ent. Amer. (n. s.) 29: 7, 10, 11.
Type-species: Anopompilinus michiganensis Dreisbach. Monotypic.
Pompilus subg. Holarctopompilus Wolf, 1965. Schweiz. Ent. Gesell., Mitt. 38: 101. N. syn.
(H. E. Evans).
Type-species: Psammochares gibbomimus Haupt. Orig. desig.
Pompilus subg. Boreopompilus Wolf, 1965. Schweiz. Ent. Gesell., Mitt. 38: 101. N. syn. (H.
E. Evans).
Type-species: Pompilus trivialis Dahlbom. Orig. desig.
angularis angularis (Banks). Transcont. in Transit. and U. Austr. Zones; Mexico (Baja
California). Ecology: Occurs in sandy areas, and also nests in hard-packed soil. Prey:
Callilepis altitudonis Chamb., C. imbecilla (Keys.); Xysticus spp.; Pellenes spp.,
Habronattus ? sp., Thomisidae sp. Another subsp. occurs in Mexico.
Psammochares angularis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 115. 6.

Biology: Evans, 1959. Kans. Ent. Soc., Jour. 32: 76 (prey transport). —Wasbauer and Powell,
1962. Kans. Ent. Soc., Jour. 35: 399-400 (prey). —Evans, 1963. Ent. News 74: 239 (prey
transport). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 482 (prey transport).
—Kurczewski and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 72-73 (prey transport).
—Aleock, 1973. Wasmann Jour. Biol. 31: 323-324, fig. 1 (nest, prey).

anomalus anomalus (Dreisbach). U. Sonor. and Transit. Faunas, Alta. to Ariz., Calif.; Mexico
(Baja California). Prey: Xysticus conctator Thor. Another subsp. occurs in Mexico.
Anopompilinus anomalus Dreisbach, 1950. Amer. Midland Nat. 42: 725, 737. d.
Anopompilinus arnaudi Dreisbach, 1952. Amer. Midland Nat. 48: 153, figs. 1, 2. 3.

Biology: Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 400 (prey).

1564 Hymenoptera in America North of Mexico

dakota dakota (Dreisbach). Transit. Zone, Alaska, Mont. and N. Dak. s. to Ariz. and N. Mex.
Prey: Thanatus formicinus (Oliv.). Another subsp. occurs in Europe.
Anopompilinus dakota Dreisbach, 1950. Amer. Midland Nat. 42: 728, figs. 9, 10. d.

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 77: 246 (prey).
imbecillus imbecillus (Banks). Transcont. in Transit. Zone except Pacific States. Ecology:
Occurs in open woods.
Anopliella imbecilla Banks, 1939. Canad. Ent. 71: 227. °.
Anopompilinus coloradensis Dreisbach, 1950. Amer. Midland Nat. 42: 724, 737. 6.
Anopompilinus banksi Dreisbach, 1950. Amer. Midland Nat. 42: 728, 737. 6.
imbecillus ojibwae Evans. Canad. and Hudson. Zones, N. B. and B. C., n. to N. W. T.
Pompilus (Ammosphex) imbecillus ojibwae Evans, 1951. Amer. Ent. Soc., Trans. 77: 249.
OMe
luctuosus luctuosus Cresson. Transcont. in Canad. Zone, entering Transit. Zone in West.
Ecology: Occurs in open woods. Other subspp. occur in Palaearctic Region.
Pompilus luctuosus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 452. °.
Anopompilinus hirsutus Dreisbach, 1950. Amer. Midland Nat. 42: 733, 737. 6.
Anopompilinus mainensis Dreisbach, 1950. Amer. Midland Nat. 42: 733, 737. d.
michiganensis michiganensis (Dreisbach). Hudson., Canad., and Transit. Zones from N. B., N.
W. T., Yukon, s. to Alta., Minn., N. Y., and in mts. to Colo. and Ga. Ecology: Occurs in
open woods and fields; nests in sand and sandy loam. Prey: Xysticus conctator Thor., X.
transversatus (Walck.), X. ferox (Hentz), X. funestus (Keys.), X. sp. Another subsp.
occurs in the Palaearctic Region.
Anopompilinus michiganensis Dreisbach, 1949. Ent. Amer. (n. s.) 29: pl. 1, fig. 3. 3.
Anopompilinus aspinosus Dreisbach, 1950. Amer. Midland Nat. 42: 730, 738. 3, @.

Biology: Kurezewski and Snyder, 1964. Biol. Soc. Wash., Proc. 77:215-222, 6 figs. (prey
transport, nest). —Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 23 (prey).
—Kurezewski and Kurczewski, 1968. Kans. Ent. Soc., Jour. 41: 376 (prey, nest).
—Kurcezewski and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 73 (prey).

occidentalis (Dreisbach). Transit. Fauna, B. C. and Alta. to Colo. and Calif., s. in Mexico to
Morelos and Puebla. Ecology: Nests in gravel bank. Prey: Pardosa uintana Gertsch, P.
sp.
Anopompilinus occidentalis Dreisbach, 1950. Amer. Midland Nat. 42: 726, 736, 738. 6, 9.

Biology: Powell, 1957. Pan-Pacific Ent. 33: 39-40 (prey). —Evans, 1963. Ent. News 74: 239

(prey transport).
parvulus (Banks). U. Sonor. and Transit. Faunas, N. Mex. and Alta., w. to the Pacific, s. in
Mexico to Baja California and Zacatecas.
Pompiloides parvulus Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 227. 2, 3.
Pompilus (Ammosphex) parvulus tepahuanus Evans, 1953. Ent. Soc. Amer., Ann. 46: 542.
OE
silvivagus Evans. Transit. Zone forests, Tex., N. Mex., Colo., Ariz., Calif.
Pompilus (Ammosphex) solonus silvivagus Evans, 1951. Amer. Ent. Soc., Trans. 77: 241,
figs. 184, 218. 6, @.
solonus solonus (Banks). L. and U. Sonor. Faunas, Calif., Colo., and cent. Tex.; Mexico (Baja
California). Prey: Lycosa sp. Another subsp. occurs in Mexico.
Pompiloides solonus Banks, 1914. N. Y. Ent. Soc., Jour. 22: 303. 9.

Biology: Evans, 1951. Amer. Ent. Soe., Trans. 77: 241 (prey).

wasbaueri Evans. Calif. (Contra Costa and Tuolumne Co’s.).
Pompilus (Ammosphex) wasbaueri Evans, 1966. Amer. Ent. Soc., Mem. 20: 396. 3.

Genus POMPILUS Subgenus ARACHNOSPILA Kincaid

Arachnospila Kineaid, 1900. Wash. Acad. Sci., Proe. 2: 509.
Type-species: Arachnospila septentrionalis Kineaid. Monotypic.
Pycnopompilus Ashmead, 1902. Canad. Ent. 34: 83.
Type-species: Pompilus scelestus Cresson. Orig. desig.

Superfamily POMPILOIDEA 1565

arctus Cresson. Transcont. in Canad. and Transit. Zones; Mexico (Hidalgo, Mexico). Ecology:

Occurs in open woods; nests in gravelly soil. Prey: Arctosa sp., Schizocosa saltatrix
(Hentz), Trochosa avara Keys.; Amaurobius bennetti (Blackw.), A. ferow Walck.;
Clubiona sp.; Orodrassus coloradensis Em.

Pompilus arctus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 453. “2” = ¢.

Psammochares anoplinus Banks, 1919. Mus. Compar. Zool., Bul. 63: 231, 232. 9.

Pycnopompilus stouxensis Dreisbach, 1950. Amer. Midland Nat. 43: 592, figs. 46, 47. ¢.

Pycnopompilus sculleni Dreisbach, 1950. Amer. Midland Nat. 43: 594, figs. 33, 37. d.

Pycnopompilus parvus Dreisbach, 1952. Amer. Midland Nat. 48: 152, figs. 3, 4. d.

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 77: 260-261 (nest, prey). —Evans and
Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 86 (prey). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 376 (prey).

fumipennis eureka (Banks). Canad. and Transit. Zones, B. C. and Mont. to Calif. and N. Mex.;
Mexico (Baja California). Ecology: Nests off side of mammal burrow. Prey: Tarentula
kochi Keys., Trochosa pratensis Em., Lycosa sp.
Psammochares eureka Banks, 1919. Mus. Compar. Zool., Bul. 63: 231. 2 (d¢ misdet.).
Psammochares catalinae Banks, 1933. Psyche 40: 7. 2, 3.

Biology: Evans, 1951. Amer. Ent. oc., Trans. 77: 268 (prey, nest). —Evans, 1959. Kans. Ent.
Soc., Jour. 32: 75 (prey). —Wasbauer and Powell, 1962. Kans. Ent. Soc., Jour. 35: 400
(prey).

fumipennis fumipennis Zetterstedt. Transcont. in Canad. Zone, Alaska, N. W. T., and
Labrador to Wash., Alta., N. Dak., and N. Y.; north. Europe.
Pompilus fumipennis Zetterstedt, 1838. Ins. Lapponica, v. 1, p. 438. 2.
Arachnospila septentrionalis Kincaid, 1900. Wash. Acad. Sci., Proe. 2: 509. 2.
Psammochares lasiope Banks, 1919. Canad. Ent. 51: 81. °.
scelestus Cresson. Transcont. in Transit. and U. Austral Zones, southern Canada to Calif. and
Ga., s. in Mexico at higher altitudes to Guerrero and Oaxaca. Ecology: Occurs in open
woods and on sand dunes; nests in light soil or sand. Parasite: Evagetes parvus (Cr.),
Ceropales sp. Prey: Lycosa gulosa Walck., L. frondicola Em., L. sp., Trochosa avara
Keys., Lycosidae sp.; Dolomedes sp.; Phidippus sp.
Pompilus scelestus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 451. 2, 6.
Pompilus pulchrinellus Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 194, pl. 11, figs. 12,
12a. 9.
Pompilus Omiltemensis Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 197, pl. 11, figs.
18, 18a. 6.
Psammochares astur Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 225. 9°.
Psammochares sublaevis Banks, 1921. Ent. Soc. Amer., Ann. 14: 20. 9.
Pycnopompilus subscelestus Dreisbach, 1950. Amer. Midland Nat. 43: 591, figs. 39, 40. 3.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 153-163 (prey,
nest, parasite). —Peckham and Peckham, 1905. Wasps, social and solitary, pp. 230-242
(prey, nest, parasite). —Rau and Rau, 1918. Wasp Studies Afield, pp. 64-67 (prey, nest).
—Evans, 1951. Amer. Ent. Soc., Trans. 77: 264 (prey, nest). —Kurezewski and Kurezewski,
1968. Kans. Ent. Soc., Jour. 41: 23 (prey). —Kurezewski and Kurezewski, 1968. Kans. Ent.
Soc., Jour. 41: 377 (prey). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 483 (prey transport,
nest). —Kurcezewski and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 73-74, fig. 2 (prey,
nest).

Genus POMPILUS Subgenus ANOPLOCHARES Banks

Anoplochares Banks, 1939. Canad. Ent. 71: 225, 229.
Type-species: Pompiloides rectus Banks. Orig. desig.

The North American species are found in open woods. The lack of a tarsal pecten suggests
that the females nest in pre-existing cavities, such as those of the prey as in the European spis-
sus Schiodte.

apicatus Provancher. Newfoundland, Sask., and B. C. s. to Fla., Tex., and Calif.
Pompilus apicatus Provancher, 1882. Nat. Canad. 13: 35, 38.

1566 Hymenoptera in America North of Mexico

Pompiloides rectus Banks, 1914. N. Y. Ent. Soc., Jour. 22: 308. 2, 6.
Pompiloides elsinore Banks, 1919. Mus. Compar. Zool., Bul. 63: 235, 236. °.
similaris (Banks). Alleghanian and Carol. Faunas, Ont. and New England s. to Fla. and w. to

Anoplius similaris Banks, 1919. Canad. Ent. 51: 82. °.
Genus APORINELLUS Banks

Aporinellus Banks, 1912 (1911). Ent. Soc. Wash., Proe. 13: 238.
Type-species: Aporus fasciatus Smith. Desig. by Banks, 1912.
Ferreoloides Haupt, 1929. Zool. Mus. Berlin, Mitt. 15: 150.
Type-species: Pompilus moestus Klug. Desig. by Bradley, 1944.
Eupompiloides Gussakovsky, 1935. Konowia 14: 141.
Type-species: Pompilus moestus Klug. Monotypice.
Ceratopompilus Bradley, 1944. Eos 20: 95, 97.
Type-species: Pompilus sexmaculatus Spinola. Orig. desig.

Occur chiefly in sandy places, and visit honeydew and occasionally flowers; prey usually upon
spiders of the family Salticidae.

Revision: Bequaert, 1919. Psyche 26: 115-123. —Evans, 1966. Amer. Ent. Soc., Mem. 20:
402-423.

Taxonomy: Evans, 1957. Pan-Pacific Ent. 33: 185-186 (revised key).
basalis Banks. Kans., Tex. and Oreg. s. to Costa Rica. Ecology: Nests in sand. Prey: Oxyopes
helius Chamb.
Aporinellus basalis Banks, 1933. Psyche 40: 3. 9.
Aporinellus bridwelli Evans, 1951. Amer. Ent. Soc., Trans. 77: 291, figs. 197, 265. ¢.

Biology: Evans, 1959. Kans. Ent. Soc., Jour. 32: 76 (prey transport).

borregoensis Evans. South. Calif., Ariz. and Nev.
Aporinellus borregoensis Evans, 1957. Pan-Pacific Ent. 33: 184, fig. 2. 5, 2.
completus Banks. Transcont. in south. Canada and U.S., s. in Mexico to Chiapas at higher
altitudes. Ecology: Nests in soil. Prey: Phidippus clarus Keys., Habronattus sp.,
Pellenes oregonensis Peckh., P. viridipes (Hentz), P. borealis (Bks.), P. sp., Evarcha
hoyi (Peckh.), Habrocestum pulex (Hentz), Maevia vittata (Hentz), Sitticus palustris
(Peckh.).
Aporinellus completus Banks, 1917. Mus. Compar. Zool., Bul. 61: 97. 2.

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 77: 294 (prey). —Evans, 1962. Kans. Ent. Soc.,
Trans. 32: 76 (prey). —Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 63 (prey, nest).
—Kurezewski and Kurezewski, 1968. Kans. Ent. Soe., Jour. 41: 377 (prey). —Kurezewski
and Kurczewski, 1973. Kans. Ent. Soc., Jour. 46: 74 (prey).

fasciatus (Smith). N. C., Ill, Alta. and B. C., s. to Fla., Tex. and Calif., s. in Mexico at higher
altitudes to Puebla and Morelos.
Aporus fasciatus Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 175. ¢. Preoce. in Pompilus.
Pompilus unionis Dalla Torre, 1897. Cat. Hym., v. 8, p. 330. N. name.
Aporus apicatus Banks, 1910. N. Y. Ent. Soc., Jour. 18: 126. d.
medianus Banks. Calif., Wyo., Mich. and N. Y. s. to Costa Rica. Ecology: Nests in sand. Prey:
Oxyopes salticus Hentz, Maevia vittata Hentz, Phidippus whitmanii Peckh. ?, P. sp.,
Salticus sp.; Tibellus duttoni (Hentz), Xysticus sp. near gulosus Keys. This sp. was
consistently misdet. as fasciatus Sm. until 1966.
Aporinellus medianus Banks, 1917. Mus. Compar. Zool., Bul. 61: 97. 9.
Aporinellus intermedius Banks, 1919. Mus. Compar. Zool., Bul. 63: 240. °.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 55-57 (prey, nest).
—Krombein, 1959. Ent. Soc. Wash., Proc. 61: 194-195 (prey, nest). —Kurezewski and
Kurcezewski, 1968. Kans. Ent. Soc., Jour. 41: 23-24 (prey hunting and transport, nest).

taeniatus baboquivari Evans. Ariz. (Baboquivari Mts.).
Aporinellus taeniatus baboquivari Evans, 1951. Amer. Ent. Soc., Trans. 77: 286. @.

Superfamily POMPILOIDEA 1567

taeniatus rufus Banks. Colo., Minn., Nebr.
Aporinellus rufus Banks, 1912 (1911). N. Y. Ent. Soe., Jour. 19: 230. 2.
taeniatus taeniatus (Kohl). B. C., Minn. and Mass. s. to Calif., La. and Fla., and to Guatemala
from sea level to at least 7500 ft. Ecology: Nests in sand. Prey: Habronattus calcaratus
(Bks.), Pellenes agilis (Bks.), P. borealis (Bks.), P. viridipes (Hentz), P. sp.
Pompilus taeniatus Kohl, 1886. Zool.-Bot. Gesell. Wien, Verh. 36: 315. °.
Pompilus taeniolatus Dalla Torre, 1897. Cat. Hym., v. 8, p. 326. N. name.
Aporus ferrugineipes Viereck, 1906. Amer. Ent. Soc., Trans. 32: 203. 2.
Aporinellus californicus Rohwer, 1917. U.S. Natl. Mus., Proce. 53: 240. 9.
Aporinellus banksi Bequaert, 1919. Psyche 26: 118. °.
Aporinellus semirufus Banks, 1929. Psyche 36: 326. °.
Aporinellus bequaerti Banks, 1933. Psyche 40: 3. °.

Biology: Evans, 1951. Amer. Ent. Soc., Trans. 77: 284 (prey). —Krombein, 1964. Amer. Mus.
Novitates 2201: 13 (prey). —Kurezewski and Kurezewski, 1973. Kans. Ent. Soc., Jour. 46:
74 (prey, nest).
taeniatus wheeleri Bequaert. Alleghanian Fauna, Mass. to N. C.
Aporinellus wheeleri Bequaert, 1919. Psyche 26: 118. °.
yucatanensis (Cameron). N. J., Iowa, Mont. and Oreg. s. to Costa Rica except at high altitudes.
Ecology: Nests in sand. Prey: Xysticus sp.
Pompilus (Aporus) yucatanensis Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 189. °.
Aporinellus laticeps Banks, 1912 (1911). N. Y. Ent. Soc., Jour. 19: 230. °.
Aporinellus sinuatus Evans, 1951. Amer. Ent. Soc., Trans. 77: 298, figs. 199, 246, 250, 253,
256. 3d, @.

Biology: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 194 (prey, nest; misdet. as apicatus).
Genus ALLOCHARES Banks

Allochares Banks, 1917. Mus. Compar. Zool., Bul. 61: 98.
Type-species: Allochares bruesi Banks. Monotypic.

Revision: Bradley, 1944. Amer. Ent. Soc., Trans. 70: 148.
azureus (Cresson). Fla. to south. Calif., s. in Mexico to Puebla and Veracruz.
Pompilus (Agenia) azureus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 131. 2.
Pompilus sinaloae Cameron, 1893. Biol. Cent.-Amer., Hym., v. 2, p. 192. @.
Allochares bruesi Banks, 1917. Mus. Compar. Zool., Bul. 61: 98. °.

Genus PARACYPHONONYX Gribodo

Paracyphononyx Gribodo, 1884. Mus. Civ. Stor. Nat. Genova, Ann. (2) 1: 306.
Type-species: Paracyphononyx melanicrus Gribodo. Monotypic.

Schistosalius Saussure, 1892. In Grandidier, Hist. Nat. Madagascar, v. 20, Hym., v. 1, p.

313.

Type-species: Salius ellioti Saussure. Desig. by Ashmead, 1902.

Paracyphonyx Ashmead, 1902. Canad. Ent. 34: 81.
Type-species: Paracyphonyx metemmensis Magretti. Orig. desig.

Allocyphonyx Ashmead, 1902. Canad. Ent. 34: 136.
Type-species: Pompilus maurus Cresson. Orig. desig.

Pompiloides Sustera, 1913. Zool.-bot. Gesell. Wien, Verhandl. 62: 180.
Type-species: Pompilus ruficrus Klug. Monotypic.

Dicyrtomus Haupt, 1927. Deut. Ent. Ztschr., Beih., pp. 150, 256.
Type-species: Pompilus ruficrus Klug. Orig. desig.

Atopopompilus Arnold, 1937. Transvaal Mus., Ann. 19: 22.
Type-species: Pompilus venans Kohl. Orig. desig.

Anacyphononyx Haupt, 1950. Explor. Pare. Nat. Albert, Miss. de Witte, fasc. 69, p. 59.
Type-species: Pompilus semiplumbeus Taschenberg. Orig. desig.

funereus (Lepeletier). Mass., Mich., S. Dak. and Utah s. to Fla., Tex. and Ariz., s. in Mexico to
Yucatan. Prey: Lycosa lenta (Hentz).
Anoplius funereus Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 449. d.

1568 Hymenoptera in America North of Mexico

Pompilus maurus Cresson, 1867 Amer. Ent. Soc., Trans. 1: 88. 2, d.
Psammochares (Allocyphonyx) harpalyce Banks, 1910. Psyche 17: 250. 3.

Biology: Hurd and Wasbauer, 1956. Kans. Ent. Soc., Jour. 29: 169 (prey).
SUBFAMILY CEROPALINAE

This group is considered a family by some workers.
Revision: Townes, 1957. U. S. Natl. Mus., Bul. 209: 221-272, 32 figs.

TRIBE MINAGENIINI

Genus MINAGENIA Banks

Pseudagenia subg. Minagenia Banks, 1934. Amer. Acad. Arts and Sci., Proc. 69: 40, 64.
Type-species: Pseudagenia (Minagenia) brevicornis Banks. Orig. desig.
Nannochilus Banks, 1934. Mus. Compar. Zool., Bul. 94: 171.
Type-species: Pseudagenia externa Banks. Orig. desig.
Compsagenia Haupt, 1959. Nova Acta Leopoldina 21, no. 141: 29.
Type-species: Compsagenia laevipes Haupt. Orig. desig.

Biological notes have been reported on osoria (Bks.) only. The larva lives as an external
parasite of an active lycosid spider. The wasp cocoon is spun under bark.

clypeata (Banks). Md. to N. C., Mich.
Ageniella clypeata Banks, 1914. N. Y. Ent. Soc., Jour. 22: 306. ¢.
Minagenia shappirioi Dreisbach, 1953. Amer. Midland Nat. 49: 839, figs. 1, 5. 3.
congrua (Cresson). Alleghan. Fauna, Que. to Va., W. Va., Mich. M. michiganensis Dreisb. is
possibly a syn.
Pompilus (Agenia) congruus Cresson, 1867. Amer. Ent. Soc., Trans. 1: 129. 9.
Agenia rufigastra Provancher, 1889. Addit. Corr. Faune Ent. Canada, Hym., p. 264. 2.
Minagenia semirufa Dreisbach, 1953. Amer. Midland Nat. 49: 841, figs. 10, 14. ¢.
externa (Banks). Tex. (Lee Co.).
Pseudagenia externa Banks, 1910. N. Y. Ent. Soc., Jour. 18: 124. 9.
julia (Brimley). Atlantic and Gulf Coast States, Md. to Tex.
Ageniella julia Brimley, 1934. Ent. News 45: 42. 6.
lata Townes. S. C. (McClellanville, Columbia).
Minagenia lata Townes, 1957. U.S. Natl. Mus., Bul. 209: 230, pl. 3, fig. 36, pl. 4, fig. 44. ¢.
lutea Dreisbach. Okla. (Vinita).
Minagenia lutea Dreisbach, 1955. Ent. News 66: 106. °.
major Townes. D. C., Ga.
Minagenia major Townes, 1957. U. S. Natl. Mus., Bul. 209: 229. dg, @.
michiganensis Dreisbach. Mich. (Roscommon Co.). Possibly a syn. of congrua (Cr.).
Minagenia michiganensis Dreisbach, 1953. Amer. Midland Nat. 49: 842, figs. 9, 12. 3.
montisdorsa Dreisbach. Ohio, Ga., La., Tex.; Mexico (Baja California, Zacatecas).
Minagenia montisdorsa Dreisbach, 1953. Amer. Midland Nat. 49: 840, figs. 3, 7. 6.
osoria (Banks). Md., Va., W. Va., Ga., Tex. Host: Lycosidae spp.
Nannochilus osoria Banks, 1944. Mus. Compar. Zool., Bul. 94: 172. 6, 2.

Biology: Kaston, 1959. Brooklyn Ent. Soc., Bul. 54: 103-106, fig. 9 (host, life history).
TRIBE NOTOCYPHINI

Genus NOTOCYPHUS Smith

Notocyphus Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 172.
Type-species: Notocyphus saevissimus Smith. Desig. by Smith, 1873.
dorsalis arizonicus Townes. L. Sonoran Fauna of Ariz. and Calif.
Notocyphus dorsalis arizonicus Townes, 1957. U. . Natl. Mus., Bul. 209: 223. 6, 9.

Superfamily POMPILOIDEA 1569

dorsalis dorsalis Cresson. Tex.; Mexico (Chihuahua, Durango).
Notocyphus dorsalis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 207. 2.
Notocyphus tevanus Cresson, 1872. Amer. Ent. Soe., Trans. 4: 207. d.

TRIBE CEROPALINI

Genus CEROPALES Latreille

Ceropales Latreille, 1796. Precis Caract. Gen. Ins., p. 123.
Type-species: Evania maculata Fabricius. Desig. by Curtis, 1839.

Ceratopales (!) Howard, 1901. Insect Book. pl. 5, fig. 10.

Ceratopales Schulz, 1906. Spolia Hym., p. 174. Emend.

Agenioxenus Ashmead, 1902. Canad. Ent. 34: 137.
Type-species: Ceropales rufiventris Walsh. Orig. desig. (=Ceropales robinsonii

Cresson).

Hypsiceraeus Morice and Durrant, 1915. Ent. Soc. London, Trans. pp. 403, 405.

Type-species: Evania maculata Fabricius. Orig. desig.

Members of this genus are social parasites of other spider wasps. The female Ceropales lays
her egg in the book lung of the spider prey of the host either during transport of the prey or
while the prey is lying unguarded. The Ceropales larva hatches first, devours the host egg, and
then feeds on the host prey.

Revision: Fox, 1892. Amer. Ent. Soc., Trans. 19: 49-63.

Taxonomy: Dreisbach, 1948 (1946). Mich. Acad. Sci., Arts, and Letters, Papers 32: 249 (key to
Mich. spp.). —Dreisbach, 1948. N. Y. Ent. Soc., Jour. 56: 233-268 (key to U.S. spp.).
bipunctata bipunctata Say. Atlantic to 100° W. in Transit. and U. Austr. Zones.
Ceropales bipunctata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app., p. 334. d,
Q.

bipunctata tibialis Banks. N. C. to Fla.
Ceropales bipunctata var. tibialis Banks, 1910. N. Y. Ent. Soc., Jour. 18: 126. d, °.
Ceropales floridensis Dreisbach, 1948. N. Y. Ent. Soc., Jour. 56: 233. 2.
brevicornis Patton. La., Tex., N. Mex. n. to Pa. and Alta., Wash.
Ceropales brevicornis Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 368. d.
cubensis albopicta Cresson. South. Tex. to Panama, Baja California. Typical cwbensis Cresson
occurs in the West Indies.
Ceropales albopicta Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 378. d.
elegans aquilonia Townes. Alta., Minn.
Ceropales elegans aquilonia Townes, 1957. U. S. Natl. Mus., Bul. 209: 256. ¢.
elegans elegans Cresson. Pacific to 100° W. in U. Austr. and L. Austr. Zones.
Ceropales elegans Cresson, 1872. Amer. Ent. Soc., Trans. 4: 208. 9.
Ceropales Cressoni Fox, 1892. Amer. Ent. Soc., Trans. 19: 58. d, 2.
elegans quaintancei Viereck, Md. to Fla., IIll., Kans.
Ceropales quaintancei Viereck, 1902. Ent. News 13: 275. “3” = @.
femoralis Cresson. Va. to Kans., s. to Panama.
Ceropales femoralis Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 378. °.
Ceropales foxii Rohwer, 1916. Canad. Ent. 48: 369. 6.
fulvipes Cresson. Tex.
Ceropales fulvipes Cresson, 1872. Amer. Ent. Soc., Trans. 4: 208. 9.
hatoda Brimley N. Y. to N. C., Minn. Host: Ageniella partita Bks.
Ceropales hatoda Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 48: 201. d.

Biology: Krombein, 1955. Brooklyn Ent. Soc., Bul. 50: 15 (host).

longipes Smith. N. J. to Ga., Mo.
Ceropales fasciata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app., p. 333;
Leconte Ed. v. 1, p. 224 Preoce.
Ceropales longipes Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 179. 9.
Ceropales frigida Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 180. N. name.

1570 Hymenoptera in America North of Mexico

maculata caenosa Townes. Calif. (Sierra Nevada Mts.).
Ceropales maculata caenosa Townes, 1957. U. S. Natl. Mus., Bul. 209: 242. 2, do.
maculata fraterna Smith. Transcont., chiefly in Canad. and Transit. Zones, but replaced in far
West by maculata stretchii Fox and maculata caenosa Townes, and in the
Alberta-Dakotas area by maculata rhodomerus Townes. Host: Pompilus scelestus Cr.,
Priocnemis sp., prob. germana Cr. Typical maculata (Fabricius) is European.
Ceropales fraterna Smith, 1855. Cat. Hym. Brit. Mus., v. 3, p. 180. °.
Ceropales minima Provancher, 1887. Addit. Corr. Faune Ent. Canada, Hym., p. 265. d.
Ceropales fraternus occidentalis Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 455. 3.

Taxonomy: Rohwer, 1916. Canad. Ent. 48: 369 (notes on type of minima).

Biology: Peckham and Peckham, 1898. Wis. Nat. Hist. Survey, Bul. 2: 154-155, 239 (host).
—Evans and Yoshimoto, 1962. Ent. Soc. Amer., Mise. Pub. 3: 112 (host).
maculata rhodomerus Townes. South. Alta. and Mont. to N. and S. Dak., and occasional
individuals from Minn., Mich., N. Y., Mass. in range of maculata fraterna Smith.
Ceropales maculata rhodomerus Townes, 1957. U. S. Natl. Mus., Bul. 209: 243. 2, d.
maculata stretchii Fox. Calif. east of Sierra Nevada Mts., n. to B. C. and e. to Idaho, north.
Nev., and Utah.
Ceropales Stretchii Fox, 1892. Amer. Ent. Soc., Trans. 19: 52. 2.
neomexicana Rohwer. N. Mex., Calif.; Mexico (Zacatecas, Durango).
Ceropales neomexicana Rohwer, 1915. U. S. Natl. Mus., Proc. 49: 236. d.
nigripes Cresson. Pacific to 100° W. in Transit. and U. Austr. Zones; Mexico (Durango,
Zacatecas).
Ceropales nigripes Cresson, 1867. Amer. Ent. Soc., Trans. 1: 139. @.
Ceropales texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 208. d.
pacifica Townes. Oreg., Calif.
Ceropales pacifica Townes, 1957. U. S. Natl. Mus., Bul. 209: 264. d, 2.
robinsonii robinsonii Cresson. Que. to Fla., Ont., Ohio, Ill. Host: Phanagenia bombycina (Cr.).
Ceropales Robinsonii Cresson, 1867. Amer. Ent. Soc., Trans. 1: 140. d.
Ceropales rufiventris Walsh and Riley, 1869. Amer. Ent. 1: 136, 163. 3, °.
Ceropales superba Provancher, 1883. Nat. Canad. 14: 36. °.

Biology: Walsh and Riley, 1869. Amer. Ent. 1: 136, 163 (host).
robinsonii stigmatica Banks. Kans., Tex.
Ceropales robinsonii var. stigmatica Banks, 1910. N. Y. Ent. Soc., Jour. 18: 126. 2.
rugata Townes. Oreg., Calif., Wyo., Utah, Gulf and South Atlantic States; Mexico (Zacatecas,
Teotihuacan).
Ceropales rugata Townes, 1957. U. S. Natl. Mus., Bul. 209: 264. 5, °.

UNPLACED TAXA OF POMPILIDAE

Pompilus bipartitus Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 439. ?. Philadelphia.
Preocc.; replaced by Pompilus semipartitus Dalla Torre.

Pompilus semipartitus Dalla Torre, 1897. Cat. Hym., v. 8, p. 321. N. name for Pompilus
bipartitus Lepeletier.

Family RHOPALOSOMATIDAE

During their larval stages the two North American rhopalosomatids are external parasites of
nymphal crickets. Gurney (1953) undoubtedly had material of both species before him in his
study of the biology and larval stages of what he supposed to be Rhopalosoma only. He
recorded as hosts Hapithis a. agitator Uhler, H. agitator quadratus Scudd., H. brevipennis
Sauss., H. sp., and Orocharis sp.

Taxonomy: Gurney, 1953. U. S. Natl. Mus., Proc. 103: 25-31, figs. 8, 9 (larva).
Biology: Gurney, 1953. U. S. Natl. Mus., Proc. 103: 19-25, pl. 1.

Genus RHOPALOSOMA Cresson
The adults are fully winged, have enlarged ocelli and are nocturnal.

Superfamily POMPILOIDEA 1571

Rhopalosoma Cresson, 1865. Ent. Soc. Phila., Proc. 4: 58.
Type-species: Rhopalosoma Poeyi Cresson. Monotypic.
Sibyllina Westwood, 1868. Ent. Soc. London, Trans. p. 329.
Type-species: Sibyllina aenigmatica Westwood. Monotypic (=Rhopalosoma poeyi
Cr.).
Revision: Brues, 1943. Ent. Soc. Amer., Ann. 36: 310-318, 1 pl.
nearcticum Brues. Md. south to Fla., Ky., Mo. Host: Orocharis saltator Uhler.
Rhopalosoma nearcticum Brues, 1943. Ent. Soc. Amer., Ann. 36: 316. 2, d.
Biology: Hood, 1914 (1913). Ent. Soc. Wash., Proc. 15: 145-147, 1 fig. (Misdet. as R. poeyi
Cresson.)

Genus OLIXON Cameron

Olixon Cameron, 1887. Biol. Cent.-Amer. Hym., v. 1, p. 412.
Type-species: Olixon testacewm Cameron. Monotypic.

Saphobethylus Kieffer, 1911. Soc. Sci. Bruxelles, Ann. (2) 35: 216.
Type-species: Saphobethylus pallidus Kieffer. Monotypic.

Nealgoa Brues, 1922. Psyche 29: 105. cs
Type-species: Nealgoa banksii Brues. Monotypic.

This genus is unusual in that extreme brachyptery is equally developed in both sexes. The
species are much smaller than Rhopalosoma, have tiny ocelli and are diurnal.

Taxonomy: Reid, 1939. Roy. Ent. Soc. London, Proc. 8: 101.
banksii (Brues). N. Y., D. C., N. C., Ga., Fla., Ohio, Minn., Kans. Host: Nemobius sp.; Hapithis
agitator Uhler.
Nealgoa banksii Brues, 1922. Psyche 29: 106. °.

Biology: Krombein, 1950 (1949). Elisha Mitchell Sci. Soc., Jour. 65: 264-265 (male).

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1573

Superfamily SPHECOIDEA

By KARL V. KROMBEIN

For nearly a century most specialists in this group, influenced by Kohl’s ultra-conservative
views, considered that the superfamily contained a single family, the Sphecidae. The monumen-
tal generic reclassification by Bohart and Menke (1976) embraces this opinion. However,
Brothers (1975) demonstrates convincingly that each of the aculeate superfamilies should com-
prise several families if the family-level groups are to represent categories of equal phylogenetic
value. The classification used elsewhere in this catalog supports the latter conviction. Ac-
cordingly the major subfamilies recognized by Bohart and Menke are restored to family rank, a
position accorded them by most specialists of the previous century.

Some authors believe that the sphecoid wasps and the bees belong to a single superfamily, the
Sphecoidea. For example, Brothers divides the Sphecoidea into two informal groups, the
Spheciformes and Apiformes, with eight and nine families respectively. However, on the basis of
the presence or absence of a hind tibial strigil, Boerner (1919) divides the Aculeata into two sub-
sections, the sphecoids, pompiloids and vespoids, and the formicoids, scolioids and apoids. The
phylogeny of the sphecoid wasps and bees requires more intensive investigation than they have
had hitherto, for the possibility exists that the bees may not be so closely related to the sphecoid
wasps as supposed by some workers. At this time the Sphecoidea and Apoidea are maintained as
separate superfamilies.

The behavior and life history of this diverse assemblage of wasps has attracted a host of ob-
servers both in the United States and abroad. Many species are ground nesters and are known
popularly as digger wasps; most of them dig their own nests but some species appropriate
pre-existing burrows of other arthropods and modify them as needed. Numerous species nest
above ground in pre-existing cavities such as abandoned borings of beetle larvae in wood, old in-
sect galls and old mud-dauber nests; many of these species can be induced to nest in borings in
wood called trap-nests. Some of our species excavate their own nests in soft pith of shrubs such
as sumac and elderberry, or in rotten wood. Relatively few North American species are mason
wasps, building various kinds of mud cells. Several genera are cleptoparasites of other
ground-nesting sphecoids. So far as known the North American species are all solitary wasps,
but apparent eusociality has been discovered in the Neotropical genus Microstigmus Ducke.

Members of the Sphecoidea prey upon a great variety of terrestrial insect orders as well as
upon spiders. Varying degrees of host specificity are found among the several families and
lesser categories. In general the more primitive sphecoids prey upon the more primitive and an-
cient groups of Hemimetabola while the more advanced groups prey upon the higher groups of
Holometabola.

The pre-1920 references listed below under the side-head Taxonomy are not reliable for
generic or specific discrimination.

Revision: Bohart and Menke, 1976. Sphecid wasps of world, 695 pp., 190 figs. (reclassification
of world genera, lists of species-level taxa).

1574 Hymenoptera in America North of Mexico

Taxonomy: Ashmead, 1899. Canad. Ent. 31: 145-155, 161-174, 212-225, 238-251, 291-300,
322-330, 345-357 (keys to families and genera, and lists of No. Amer. spp.). —Smith, 1908.
Nebr. Univ., Studies 8: 323-410, 1 pl. (keys to Nebr. spp.). —Mickel, 1918 (1917). Nebr.
Univ., Studies 17: 342-456, 2 figs. (keys to Nebr. spp.). —Pate, 1937. Amer. Ent. Soc., Mem.
9: 1-103 (generic names and type-species). —Evans and Lin, 1956. Amer. Ent. Soe., Trans.
81: 131-166, 13 pls. (larvae of Sphecinae). —Evans and Lin, 1956. Amer. Ent. Soc., Trans.
82: 35-66, 13 pls. (larvae of Nyssoninae). —Evans, 1957. Amer. Ent. Soc., Trans. 83: 79-117,
12 pls. (larvae of Philanthinae, Trypoxyloninae and Crabroninae). —Evans, 1959. Amer.
Ent. Soce., Trans. 85: 137-191, 7 pls. (addendum to larvae, keys to subfamilies and genera
based on larval characters, and remarks on evolution and classification based on larval
characters). —Evans, 1964. Amer. Ent. Soc., Trans. 90: 235-299, 12 pls. (larvae,
supplement). —Evans, 1964. Ent. News 75: 225-237, 3 figs. (classification and evolution of
digger wasps based on larvae). —Brothers, 1975. Kans. Univ. Sci. Bul. 50: 586-587, 640-641
(phylogeny).

Biology: Evans, 1963. Sci. Amer. 208 (4): 145-154 (evolution as evidenced by predatory
behavior). —Evans, 1966. Science 152: 465-471, 6 figs. (accessory burrows of digger wasps).
—Kurezewski and Snyder, 1968. Conservationist 23 (2): 28-31, 11 figs. (evolution of
cliff-nesting in digger wasps). —Miller and Kurezewski, 1973. In Dindal, Proc. First Soil
Microcommunities Conf., USAEC, CONF-711076; Natl. Tech. Inform. Serv., USDC, pp.
204-217 (ecology of digger wasps). —Evans, 1975 (1974). N. Y. Ent. Soc., Jour. 82: 259-267,
4 figs. (digger wasps as colonizers of new habitats). — Alcock, 1975. Anim. Behaviour 23:
893-894 (male behavior and territoriality).

Family AMPULICIDAE

These primitive sphecoid wasps occur principally in the tropics of the Old and New Worlds.
Appropriately, they prey upon cockroaches, one of the most primitive of insect orders. The biolo-
gy of only a few species has been observed. The nests are constructed in pre-existing cavities or
crevices, such as in twigs, beneath bark of trees or in the soil. Unlike other sphecoids, the prey is
carried backwards on foot as in most pompilid wasps.

Taxonomy: Kohl, 1893. K. K. Naturhist. Hofmus., Ann. 8: 455-516, 3 pls. (generic diagnoses,

key to and descriptions of Ampulex spp.). —Evans, 1959. Ent. News 70: 57-61, 6 figs.
(larvae).

SUBFAMILY DOLICHURINAE

Genus DOLICHURUS Latreille

Dolichurus Latreille, 1809. Gen. Crust. Ins., v. 4, p. 387.
Type-species: Pompilus corniculus Spinola. Monotypie.
Thyreosphex Ashmead, 1904. Canad. Ent. 36: 282.
Type-species: Thyreosphex Stantoni Ashmead. Monotypice.
Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 21, figs. F, G (male
genitalia).
greenei Rohwer. Ont. to Fla., Mo. Ecology: Nests under leaf litter in woods. Prey: Parcoblatta
sp., nymph.
Dolichurus greenei Rohwer, 1916. Ent. Soe. Wash., Proc. 18: 212. 9.
Taxonomy: Bradley, 1934. Ent. News 45: 33-34, 6.

Biology: Krombein, 1955. Brooklyn Ent. Soe., Bul. 50: 15 (prey, nest site).

Genus PARADOLICHURUS Williams

Dolichurus subg. Paradolichurus Williams, 1960. Wasmann Jour. Biol. 17: 299.
Type-species: Dolichurus (Paradolichurus) californicus Williams. Orig. desig.
californicus (Williams). Calif. (San Diego Co.).
eae rus (Paradolichurus) californicus Williams, 1960. Wasmann Jour. Biol. 17: 300,
HS Maal Oe

Superfamily SPHECOIDEA 1575

SUBFAMILY AMPULICINAE

Genus AMPULEX Jurine

Ampulex Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 132.
Type-species: Sphex compressus Fabricius. Desig. by Audouin, 1822.
Pronoeus Latreille, 1809. Gen. Crust. Ins., v. 4, p. 56.
Type-species: Dryinus aeneus Fabricius. Monotypic.
Lorrheum Leach, 1837. In Shuckard, Essay on Indig. Fossor. Hym., p. 18. A manuscript
name of Leach, validated by Shuckard.
Type-species: Sphex compressus Fabricius. Desig. by Shuckard, 1837.
Rhinopsis Westwood, 1844. Arcana Ent., v. 2, p. 68.
Type-species: Rhinopsis Abbottii Westwood. Monotypic.
Waagenia Kriechbaumer, 1874. Stettin Ent. Ztg. 35: 55.
Type-species: Waagenia sikkimensis Kriechbaumer. Monotypic.
Chlorampulex Saussure, 1892. In Grandidier, Hist. Nat. Madagascar, v. 20, p. 441.
Type-species: Sphex compressus Fabricius. Desig. by Pate, 1937.
canaliculata Say. Mass. to Ga., Ohio, Wis., Mo., Kans. Ecology: Nests in cavities in twigs and
in borings in wood. Prey: Parcoblatta virginica (Brunner), P. sp., Ischnoptera sp.
Ampulex canaliculatus Say, 1823. West. Quart. Rptr. 2: 76.
Rhinopsis Abbottii Westwood, 1844. Arcana Ent., v. 2, p. 68, pl. 65, fig. 5. 2.
Ampulex pensylvanica Haldeman, 1849. Acad. Nat. Sci. Phila., Proc. 4: 203.
Rhinopsis melanognathus Rohwer, 1912. U. S. Natl. Mus., Proc. 41: 464.“?”= 6.

Taxonomy: Schulz, 1911. Zool. Ann. 4: 149-152. —Bradley, 1934. Ent. News 45: 32-33.
—Evans, 1959. Ent. News 70: 57-58, 6 figs. (larva).

Biology: Williams, 1929. Hawaii. Ent. Soc., Proc. 7: 318-329, 10 figs. (nest, prey, life history).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 173-175, figs. 49-51 (nest, prey, life
history).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): 50, pl. 22, figs. A-D (male
genitalia).
ferruginea Bradley. Fla., Tex.
Ampulex (Rhinopsis) ferruginea Bradley, 1934. Ent. News 45: 274. 6.

Taxonomy: Strandtmann, 1943. Ent. Soc. Amer., Ann. 36: 46-48. 2.
Family SPHECIDAE

Collectively, the members of this family are known as thread-waisted wasps because of the
slender, elongate abdominal petiole. Most North American species are moderately large wasps,
many of them with conspicuous coloration. The nesting habits are quite varied: The majority of
species are digger or sand wasps, excavating their nests in soil; others utilize pre-existing cavi-
ties or borings in wood, or abandoned mud-dauber cells; a few are mud-daubers. The prey is also
quite varied and includes spiders, cockroaches, crickets, grasshoppers, katydids and larvae of
Lepidoptera and Hymenoptera. Normally, the species of a genus or higher category prey upon
species of only one of the foregoing groups.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 131-153, 73 figs. (larvae).
—Evans, 1959. Amer. Ent. Soc., Trans. 85: 147-149, figs. 31-35 (larvae). —Bohart and
Menke, 1963. Univ. Calif. Pubs. Ent. 30: 91-182, 115 figs. (reclassification). —Evans, 1964.
Amer. Ent. Soc., Trans. 90: 237-245, pls. 8-9 (larvae). —Menke, 1965. Tijdschr. v. Ent. 108:
205-217, 8 figs. (syn. of some New World spp.).

SUBFAMILY SCELIPHRONINAE
Revision: Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 99-117 (Nearctic spp.).
Genus CHLORION Latreille

Chlorion Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 333.
Type-species: Sphex lobata Fabricius. Monotypic.

1576 Hymenoptera in America North of Mexico

Chlorium Schulz, 1906. Spolia Hym., p. 193. Emend.

Revision: Menke, 1961. Ent. Soc. Amer., Ann. 54: 667-669, 9 figs. (N. Amer. spp.).
aerarium Patton. Entire U. S. Ecology: Nests in sand, probably constructing several cells per
nest. Prey: Gryllus rubens Seud., G. pennsylvanicus (Burm.), G. sp., Anurogryllus
muticus (De Geer).
Chlorion aerarium Patton, 1879. Canad. Ent. 11: 133. °.
Sphex (Chlorion) nearcticus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 186. 2, od.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 242-244, figs. 9-12 (larva).

Biology: Peckham and Peckham, 1900. Wis. Nat. Hist. Soc., Bul. 1: 85-87 (nest, prey,
parasite). —Peckham and Peckham, 1905. Wasps, Social and Solitary, pp. 256-261 (nest,
prey, parasite). —Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 283 (prey). —Krombein,
1958. Ent. Soc. Wash., Proc. 60: 104 (nest). —Krombein, 1959. Ent. Soc. Wash., Proce. 61:
195 (prey).

cyaneum Dahlbom. Tex., N. Mex., Ariz.; Mexico.
Chlorion cyaneum Dahlbom, 1843. Hym. Europaea, v. 1, p. 24.
Sphex (Chlorion) occultus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 187. 2, 6.

Genus PODIUM Fabricius

Podium Fabricius, 1804. Systema Piezatorum, pp. x, 183.
Type-species: Podium rufipes Fabricius. Desig. by Latreille, 1810.
Talthybius Rafinesque, 1815. Analyse Nature ou Tabl. Univers, Palermo, p. 124. N. name.
Ammophilus Perty, 1833. Delect. Anim. Artic. Brasil, p. 141.
Type-species: Ammophilus fumigatus Perty. Desig. by Pate, 1937.
Parapodium Taschenberg, 1869. Ztschr. Gesam. Naturw. Halle 34: 423.
Type-species: Parapodium biguttatum Taschenberg. Monotypic.

Most of the species are Neotropical. Biological information has been published on only a few
species, all of which prey upon cockroaches and nest in pre-existing cavities in wood, termite
mounds or mud-dauber nests.

Revision: Kohl, 1902. Zool.-Bot. Gesell. Wien, Abh. 1 (4): 1-101.
krombeini Bohart and Menke. Tex., southern Calif.; Mexico (Morelos, Puebla, Oaxaca).
Podium krombeini Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 106, figs. 4, 49, 60.
Grae
luctuosum Smith. N. Y. to Tex., Mo., Kans. Ecology: Nests in borings in wood or in dead tree
trunks; closing plug of rotten wood particles at inner end, the outer end plastered with
mud. Parasite: Melittobia chalybii Ashm.; Neochrysis panamensis (Cam.)? Prey:
Parcoblatta uhleriana (Sauss.), adults.
Podium luctuosum Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 235. ©.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 244-245, figs. 14, 15 (larva).

Biology: Krombein, 1964. Brooklyn Ent. Soc., Bul. 58: 118-119 (nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 255-257, figs. 64, 65 (nest, prey, life history, parasites).
rufipes Fabricius. N. Y. to Fla., Iowa, Kans., Tex.; Mexico, Central and South America.

Ecology: Nests in borings in wood and in abandoned mud-dauber nests; closing plug of a
variety of compacted debris or earth with an outer coating of resin. Parasite:
Histiostoma myrmicarum Scheuch.; Lepidophora appendiculata (Macq.); Megaselia sp.;
Melittobia chalybii Ashm.; Neochrysis panamensis (Cam.). Prey: Parcoblatta
pennsylvanica (DeG.), P. sp., Chorisoneura texensis Sauss. and Zehnt., Cariblatta lutea
(Sauss. and Zehnt.), C. minima Heb., Latiblatella rehni Heb., Eurycotis floridana
(Wlkr.), adults and nymphs. Predator: Cymatodera undulata Say.

Podium rufipes Fabricius, 1805. Systema Piezatorum, p. 183.

Parapodium biguttatum Taschenberg, 1869. Ztschr. Gesam. Naturw. Halle 34: 423. 6, 9.

Podium carolina Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 556. 9.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soe., Trans. 81: 150, figs. 69-73 (larva).

Superfamily SPHECOIDEA 1577

Biology: Rau, 1937. Ent. News 48: 91-93 (nest, prey, life history). —Krombein, 1958. Amer.
Ent. Soe., Trans. 84: 147-149 (nest, prey, life history, parasite). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 251-255, figs. 62, 63 (nest, prey, life history, parasites,
predator). —Krombein, 1970. Smithsn. Contrib. Zool. 46: 12-22, figs. 40-61 (nest behavior,
prey, life history).

Genus CHALYBION Dahlbom

Chalybion Dahlbom, 1843. Hym. Europaea, v. 1, p. 21.

Type-species: Sphex cyanea Fabricius. Desig. by Patton, 1880.
Chalybiwm Agassiz, 1847. Nomencl. Zool., p. 77. Emend.
Chalybium Schulz, 1906. Spolia Hym., p. 192. Emend.

Only the typical subgenus occurs in North America. These wasps nest in pre-existing cavities,
most commonly in abandoned nests of mud-daubers, but also in holes in wood, bamboo and plant
stems, and crevices in walls. There is one atypical record of californicum opening a sealed nest
of Sceliphron caementarium (Dru.), the black and yellow mud-dauber, extracting the enclosed
spiders, and restocking the nest with its own spiders.

Revision: Kohl, 1918. K. K. Naturhist. Hofmus. Ann. 32: 1-171 (World spp.). —Hutson, 1919.
Amer. Ent. Soc., Trans. 45: 203-227 (N. Amer. spp.).
californicum (Saussure). Transcontinental in U. S., B. C., northern Mexico; adventive in
Hawaii and Bermuda. Ecology: Nests in abandoned mud nests of Sceliphron
caementarium (Dru.). Parasite: Anthrax limatulus artemisia Marst., Sphaeropthalma
(S.) a. auripilis (Bl.). Prey: Latrodectus mactans (F.) most commonly, Asagena
americana Em., Enoplognatha puritana Chamb. and Ivie, Theridion tepidariorum
(Koch), T. frondeum Hentz, T. australe Bks., Steatoda borealis (Hentz); Neoscona sp.,
Epeira foliata (Fourcr.), Araneus sp., Gea heptagon (Hentz); Misumeninae sp.,
Thomisidae spp.; Oxyopes scalaris Hentz, Oxyopidae sp.; Paraphidippus marginatus
(Walck.), Salticidae sp.
Sphex caerulea Linnaeus, 1763. Centuria Ins. Rar., p. 30. Preoce.
Sphex cyanea Fabricius, 1775. Systema Ent., No. 5, p. 346. Preoce.
Pelopoeus (Chalybion) californicus Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p.
26.

Taxonomy: Rau, 1915. Psyche 22: 62-63 (cocoon). —Evans and Lin, 1956. Amer. Ent. Soc.,
Trans. 81: 149, figs. 67, 68 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2 (Sci. Ser. 1):
176-199 (nest, prey). —Peckham and Peckham, 1905. Wasps Social and Solitary, pp. 265-274
(nest, prey). —Rau, 1915. Ent. News 26: 469-471 (number of generations per year). —Rau,
1928. Ent. Soc. Amer., Ann. 21: 25-35 (nesting habits). —Rau, 1928. Acad. Sci. St. Louis,
Trans. 25: 443-448 (nesting habits). —Rau, 1935. Ent. News 46: 259-260 (prey). —Irving
and Hinman, 1935. Science 82: 395-396 (prey). —Muma and Jeffers, 1945. Ent. Soc. Amer.,
Ann. 38: 245-255 (prey). —Ward, 1972. Ind. Acad. Sci., Proc. 81: 177-181 (sleeping
aggregations).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14: 50, pl. 23, figs. H, J, L
(male genitalia).
zimmermanni aztecum (Saussure). Western Tex. to Ariz., Utah; Mexico to Costa Rica, Clarion
Is. Ecology: Nests in old mud nests of Sceliphron assimile (Dahlb.). Parasite: Anthrax

pluricellus Will.
Pelopeus (!) aztecus Saussure, 1867. Reise der Ost. Freg. Novara, Zool. 2: 26. 6.
Sceliphron (Chalybion) monstrosum Kohl, 1918. K. K. Naturhist. Hofmus., Ann. 32: 61. d.

Biology: Rau, 1940. Ent. Soc. Amer., Ann. 33: 591 (nests, parasite). —Rau, 1942. Canad. Ent.
74: 196 (sleeping aggregation).
zimmermanni zimmermanni Dahlbom. Tenn., N. C., Fla. to eastern Tex., Ind.; Hispaniola.
Ecology: Nests in borings in wood and in old mud nests of Sceliphron caementarium
(Dru.). Parasite: Melittobia chalybii Ashm. Prey: Araneus spp., Argiope aurantia Luc.,

1578 Hymenoptera in America North of Mexico

A. trifasciata (Forsk.), Cyclosa conica (Pall.); Theridion frondeum Hentz, Asagena
americana Em.
Chalybion zimmermanni Dahlbom, 1843. Hym. Europaea, v. 1, p. 22. 2, d.
Pelopoeus tecanus Cresson, 1872. Amer. Ent. Soe., Trans. 4: 210. °, 3.

Biology: Ward, 1970. Ind. Acad. Sci., Proc. 79: 231-233 (nest, prey). —Ward, 1971. Ind. Acad.
Sci., Proc. 80: 264-266, 1 fig. (nests). —Ward, 1973. Ind. Acad. Sci., Proe. 82: 231-233
(growth in captivity). —Ward, 1973. Ind. Acad. Sci., Proc. 82: 233-234 (parasite).

Genus SCELIPHRON Klug

Genus SCELIPHRON Subgenus SCELIPHRON Klug

Sceliphron Klug, 1801. Neue Schr. Gesell. Naturf. Freunde Berlin 3: 561.
Type-species: Sphex spirifex Linnaeus. Desig. by Bingham, 1897.
Pelopoeus Latreille, 1803 (1802). Hist. Nat. Crust. Ins., v. 3, p. 334.
Type-species: Sphex spirifex Linnaeus. Desig. by Latreille, 1810.
Pelopaeus Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, p. 180. Emend. or error.
Sceliphrum Schulz, 1906. Spolia Hym., p. 192. Emend.

Only the typical subgenus occurs in North America. Most species are tropical.
These are the common black and yellow mud-daubers which build mud cells in sheltered situa-
tions. Each cell is stored with a number of small, paralyzed spiders.

Revision: Kohl, 1918. K. K. Naturhist. Hofmus., Ann. 32: 1-171 (world spp.). —Porter, 1926.
U.S. Natl. Mus., Proc. 70 (1): 1-22 (N. Amer. spp.). —van der Vecht and van Breugel, 1968.
Tijdschr. Ent. 111: 185-255 (world spp.).

assimile (Dahlbom). Tex.; Mexico south to Panama, Cuba. Parasite: Acroricnus cubensis (Cr.).
Prey: Epeira oaxacensis Keys., E. fuscovittata Keys.; Misumena sp.; Scytodes sp.;
Phidippus sp.
Pelopoeus assimilis Dahlbom, 1843. Hym. Europaea, v. 1, p. 23. 2, d.
Sceliphron caementarium var. nicaraguanum Kohl, 1918. K. K. Naturhist. Hofmus., Ann.
32: 118. 2.

Biology: Dow, 1932. Psyche 39: 10-12 (nest, prey, parasite).
caementarium (Drury). South. Canada and entire U. S. south to Central America, West

Indies; adventive in Bermuda, Peru, Japan, Mariana Is., Marshall Is., Hawaii, Australia,
New Caledonia, Fiji, Samoa, Society Is., Marquesas Is., Gambier Is., France, Germany,
Madeira Is. Parasite: Anthrax limatulus fur (O. S.), A. l. artemisia Marst.; Amobia
floridensis (Tns.); Acroricnus s. stylator (Thunb.), A. s. edwardsvi (Cr.), A. s. junceus
(Cr.); Chrysis fuscipennis Br.; Sphaeropthalma (Photopsioides) sp., S. (S.) a. auripilis
(B1.), S. (S.) p. pensylvanica (Lep.), S. (S.) p. scaeva (Bl.). Prey: Neoscona arabesca
(Walck.), N. benjamina (Walck.), N. sp., Acanthepeira stellata (Walck.), Argiope
trifasciata (Forsk.), A. aurantia Luce., Epeira foliata (Fourcr.), E. displicata Hentz, E.
sp., Aranea nivea Hentz, A. miniata (Walck.), A. cornuta Cl. A. sp., Metepeira
labyrinthea (Hentz), Eustala anastera (Walck.), Mangora gibberosa (Hentz.);
Philodromus pernix Blackw., Misumenops asperatus (Hentz), Misumena calycina (L.),
M. sp., Misumenoides aleatorius (Hentz), Misumeninae sp., Thomisidae sp.; Phidippus
mystaceus Hentz, P. clarus Keys., P. sp., Xysticus ferox (Hentz), Marpissa undata
(DeG.), Salticidae sp.; Schizocosa crassipes (Walck.), Lycosidae sp.; Dolomedes sp.;
Anyphaenidae sp.; Oxyopes scalaris Hentz, O. salticus Hentz, Oxyopidae sp.;
Clubionidae sp. Predator: Lecontella cancellata (LeC.).

Sphex caementaria Drury, 1773. Illus. Nat. Hist., v. 2, index.

Sphex flavomaculata DeGeer, 1773. Mem. Hist. Ins., v. 3, p. 588.

Sphex lunata Fabricius, 1775. Systema Ent. p. 347.

Sphex flavipes Fabricius, 1781. Species Ins., p. 444.

Sphex flavipunctata Christ, 1791. Naturgesch. Class. Nomencl., p. 301.

Sphex affinis Fabricius, 1793. Ent. System., v. 2, p. 203.

Pelopoeus architectus Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 318. @.

Pelopoeus servillei Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 313. @.

Pelopoeus solieri Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 318. 9.

Superfamily SPHECOIDEA 1579

Pelopeous canadensis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 233. 6.
Pelopoeus nigriventris Costa, 1864. Mus. Zool. Napoli, Ann. 2: 60.
Pelopoeus tahitensis Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 27, pl. 2, fig. 17.

Sphex economica Curtiss, 1938. Short Zoology of Tahiti, p. 155.

Taxonomy: Rau, 1915. Psyche 22: 62-63 (cocoon). —Evans and Lin, 1956. Amer. Ent. Soc.,
Trans. 81: 147, figs. 50-56 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. and Nat. Hist. Survey, Bul. 2 (Sci. Ser. 1):
176-199 (nest, prey). —Peckham and Peckham, 1905. Wasps, Social and Solitary, pp.
265-274 (nest, prey). —Rau and Rau, 1913. Ent. News 24: 392-401 (nest, prey). —Rau,
1915. Jour. Anim. Behavior 5: 240-249 (experiments on prey recognition). —Rau, 1915. Ent.
News 26: 469-471 (number of generations per year). —Rau and Rau, 1918. Wasp Studies
Afield, pp. 118-124 (nest). —Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 443-466 (nest, prey,
experiments with substitute prey). —Rau, 1935. Ent. News 46: 267-270 (prey). —Muma
and Jeffers, 1945. Ent. Soc. Amer., Ann. 38: 246-255 (prey). —Rau, 1946. Brooklyn Ent.
Soc., Bul. 41: 10-11 (parasites, mating). —Shafer, 1949. Ways of a mud dauber, 78 pp., 10
pls., 9 figs. (nest, life history, physiology). —Eberhard, 1971 (1970). Psyche 77: 247-251
(predatory behavior).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99: no. 14: 50, pl. 23, figs. A-F
(male genitalia).

SUBFAMILY SPHECINAE

All of our species are ground-nesting except for those belonging to Jsodontia which nest in
pre-existing cavities in wood, stems or in the ground.

Revision: Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 77-194, 317-462 (world spp.).
—Fernald, 1906. U. S. Natl. Mus., Proc. 31: 291-423 (New World spp.). —Bohart and
Menke, 1963. Univ. Calif. Pubs., Ent. 30: 117-160 (Nearctic spp.).

TRIBE SPHECINI
Genus SPHEX Linnaeus

Genus SPHEX Subgenus SPHEX Linnaeus

Sphex Linnaeus, 1758. Syst. Nat., ed. 10, v. 1, p. 569.

Type-species: Sphex flavipennis Fabricius. Desig. by Internat]. Comn. Zool.

Nomencl., Op. 180, 1946.

Sphaex Scopoli, 1772. Observ. Zool., Hist.-Nat., v. 5, p. 122. Emend. or error.
Ammobia Billberg, 1820. Enum. Ins., p. 105.

Type-species: Pepsis argentata Fabricius. Desig. by Rohwer, 1911.
Proterosphex Fernald, 1905. Ent. News 16: 165.

Type-species: Sphex maxillosus Fabricius. Orig. desig.

Members of this subgenus are all fossorial, and many of them nest gregariously in the same
site year after year. So far as known the Nearctic species construct multicelled nests, each cell
at the end of a lateral from the vertical or oblique burrow. Preferred prey are nymphs or adults
of Tettigoniidae, but occasionally Gryllacrididae are also stored. Our species practice mass provi-
sioning, but one Oriental species practices progressive provisioning and is also unusual in con-
structing 1-celled nests.

ashmeadi (Fernald). Calif., Nev., Utah, Colo., Ariz., N. Mex., Tex.; Mexico (Tamaulipas, Nuevo

Leon).
Chlorion (Proterosphex) ashmeadi Fernald, 1906. U. S. Natl. Mus., Proce. 31: 389. 9, 6.

dorsalis Lepeletier. Southern U. S., Fla. and Ga. to Calif.; Central and South America.
Ecology: Nests in firm soil, the burrow vertical or nearly so, terminating in one or more
cells, each cell stored with 1-3 prey specimens. Prey: Conocephalus fasciatus (DeG.), C.

sp.
Sphex dorsalis Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 347. ¢.

1580 Hymenoptera in America North of Mexico

Sphex singularis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 261. d.

Sphex chlorargyrica Costa, 1862. Mus. Zool. Napoli, Ann. 1: 66.

Sphex micans Taschenberg, 1869. Ztschr. Gesam. Naturw. Halle 34: 419. 9. Preoce.
Sphex dubitata Cresson, 1872. Amer. Ent. Soe., Trans. 4: 213. 9.

Sphex spiniger Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 428. d.

Biology: Dow, 1932. Psyche 39: 8-9 (nest, prey). —Krombein and Evans, 1954. Ent. Soc.
Wash., Proce. 56: 233-234 (nest, prey, life cycle).
flavovestitus flavovestitus Smith. Va. to Fla., west to Tex.; Mexico (Durango). Parasite:
Pseudoxenos smithii (Heyd.). Another subspecies occurs in Mexico.
Sphex flavovestita Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 253. ¢.
Sphex flavipes Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 268. 2. Preoce.
Chlorion (Proterosphex) flavitarsis Fernald, 1906. U.S. Natl. Mus., Proc. 31: 379. 2, d. N.
name.
habenus Say. Md. to Fla., west to Ark. and Tex.; Mexico (Sinaloa). Parasite: Psewdoxenos
smithii (Heyd.). Prey: Tettigoniidae sp., nymph.
Sphex habena Say, 1832. New Sp. N. Amer. Ins. chiefly of Louisiana, p. 14. 2.
Sphex lauta Cresson, 1872. Amer. Ent. Soc., Trans. 4: 212. °.
Sphex lauta var. illustris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 210. °.
Sphex princeps Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 398. ¢.
Sphex chrysophorus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 300. °.
Sphex lanciger Kohl, 1895. K. K. Naturhist. Hofmus., Ann. 10: 55. 3.

Biology: Strandtmann, 1953. Kans. Ent. Soc., Jour. 26: 51-52 (nest, prey).
ichneumoneus (Linnaeus). Southern Canada, U. S., south to Brazil and Peru. Ecology: Nests

in hard-packed soil or sand, the burrow vertical or nearly so, each nest with 2-7 cells,
each cell provisioned with several prey specimens. Parasite: Pseudoxenos smithii
(Heyd.); Metopia argyrocephala (Meig.), M. campestris (Fall.), Senotainia trilineata
(Wulp); Nysson plagiatus Cr. Prey: Neoconocephalus ensiger (Harr.), N. sp., Acanthodis
SP) Atlanticus dorsalis (Burm.), Conocephalus attenuatus (Seud.), C. fasciatus (DeG.),
C. brevipennis (Seud.), C. triops (L.), C. sp., Neduba sp., Scudderia texensis (Sauss.), S.
pistillata (Brunn.), Orchelimum vulgare Harr., O. calcaratum R. and H., O. delicatum
Brun., Amblycorypha oblongifolia (DeG.); Oecanthus n. nigricornis (Wlkr.);
Brachybaenus sp., Gryllacris sp. This handsome wasp is commonly called “The Great
Golden Digger”.

Apis ichneumonea Linnaeus, 1758. Syst. Nat., ed. 10, p. 578.

Nomada surinamensis Retzius, 1783. Genera et Species Insectorum, p. 62. N. name.

Sphex auriflua Perty, 1834. Delect. Anim. Articul. Brasil., p. 142. A questionable synonym.

Sphex aurocapillus Templeton, 1841. Ent. Soc. London, Trans. 3: 51. A questionable
synonym.

Sphex Croesus Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 351. 2.

Sphex dimidiatus Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 352. ¢. Preoce.

Sphex sumptuosa Costa, 1862. Mus. Zool. Napoli, Ann. 1: 66. ¢. A questionable synonym.

Sphex ichnewmoneus var. ignotus Strand, 1916. Arch. f. Naturgesch. 81: 99. 9.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 139, figs. 1-8 (larva).

Biology: Packard, 1869. Guide Study Ins., pp. 167-168 (nest, prey). —Peckham and Peckham,
1898. Wis. Geol. Nat. Hist. Survey, Bul. 2, Sci. Ser. 1: 33-41, pl. 2, fig. 4, pl. 11, fig. 1, pl. 12,
figs. 1-2 (nest, prey). —Rau and Rau, 1918. Wasp studies afield, pp. 193-198, fig. 43 (nest,
prey). —Reinhard, 1929. Witchery of wasps, pp. 141-164, 2 pls. (prey). —Abbott, 1931.
Iowa Acad. Sci., Proc. 38: 255-258 (nesting behavior). —Frisch, 1937. Amer. Midland Nat.
18: 1043-1062 (nesting aggregation, prey, life cycle, parasite). —Fernald, 1945. Ent. Soc.
Amer., Ann. 38: 458-460 (nesting aggregation, prey). —Ristich, 1953. Canad. Ent. 85:
374-386, 1 pl., 4 text figs. (nest, prey, parasites).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14: 50, pl. 22, figs. K-N, Q,
R (male genitalia).
jamaicensis (Drury). Fla.; West Indies.

Vespa Jamaicensis Drury, 1770. Illus. Nat. Hist., v. 1, p. 104.
Sphex Jamaica Christ, 1791. Naturgesch. Class. Nomencl. Ins., p. 292. Emend.

Superfamily SPHECOIDEA 1581

Sphex aurulenta Guerin, 1835. Ieonogr. Regne Anim., Planches Anim. Invert., pl. 70, fig. 2.
Lapsus for lanierii Guer., 1844.

Sphex Lanierii Guerin, 1844. Iconogr. Regne Anim., Ins., v. 3: 433. d.

Sphex ornata Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 344. 2, d.

Sphex ichneumoneus var. fulviventris Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 431.
nudus Fernald. Md. to IIl., south to Fla. and La. Prey: Camptonotus carolinensis Gerst.

Sphex nudus Fernald, 1903. Psyche 10: 201. 3.

Sphex bridwelli Fernald, 1903. Psyche 10: 202. °.

Biology: Rau and Rau, 1918. Wasp studies afield, p. 206 (prey).
pensylvanicus Linnaeus. Transcont. in U. S. except northwestern states; northern Mexico.
Ecology: Nests in soft earth in sheltered areas, the burrow oblique; several cells
provisioned with 2-6 prey specimens. Parasite: Pseudoxenos smithii (Heyd.); Senotainia
trilineata (Wulp). Prey: Microcentrum retinerve (Burm.), M. rhombifolium (Sauss.),
Scudderia furcata Brunn. This wasp is commonly called “The Great Black Wasp.”
Sphex pensylvanica Linnaeus, 1763. Centuria Ins. Rar., p. 30.
Sphex pensylvanicus var. robustisoma Strand, 1916. Arch. f. Naturgesch. 81: 101. 9.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 140, figs. 9-11 (larva).

Biology: Reinhard, 1929. Witchery of wasps, pp. 165-191, 1 pl., 1 text fig. (nest, prey).
—Frisch, 1938. Amer. Midland Nat. 19: 673-677 (nest, prey, life cycle, parasites). —Rau,
1944. Ent. Soc. Amer., Ann. 37: 439-440 (nest, prey). —Krombein, 1955. Brooklyn Ent. Soc.,
Bul. 50: 16-17 (nesting aggregation, prey).
servillei Lepeletier. Southern Tex. to Argentina.

Sphex fuliginosa Dahlbom, 1843. Hym. Europaea, v. 1, p. 425. Preoce.

Sphex Servillei Lepeletier, 1845. Hist. Nat. Ins. Hym. 3: 336. 6.

Sphex Chichimecus Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 40. d.

Sphex congener Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 418. 9.

Sphex Joergenseni Brethes, 1913. Buenos Aires Mus. Nac. de Hist. Nat., An. 24: 120. 6.
tepanecus Saussure. Tex. to Ariz.; Mexico (Chihuahua). Ecology: Nests gregariously in fine

sandy loam soil. Prey: Tettigoniidae sp., nymph.
Sphex tepanecus Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 41. 6.
Sphex mexicana Taschenberg, 1869. Ztschr. Gesam. Naturw. Halle 34: 416. 6. Preoce.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 237-238, figs. 1-4 (larva).

Biology: Gillaspy, 1962. Brooklyn Ent. Soc., Bul. 57: 15-17 (nest, prey, mating, cocoon).

texanus Cresson. Kans. to Tex., Ariz.
Sphex Texana Cresson, 1872. Amer. Ent. Soc. Trans. 4: 212. 2, 3.

Genus SPHEX Subgenus FERNALDINA Bohart and Menke

Fernaldina Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 130.
Type-species: Sphex lucae Saussure. Monotypic.

Our single species builds a 1-celled nest in the soil which it provisions with nymphal and adult
Tettigoniidae.

lucae Saussure. Southeastern and western U.S., B. C.; northern Mexico. Ecology: Nests in
compacted sandy soil or rocky ground. Parasite: ? Senotainia sp. in trilineata (Wulp)
complex. Prey: Jnsara sp., probably elegans Scud., nymphs and adults.
Sphex lucae Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 41. 2.
Sphex belfragei Cresson, 1872. Amer. Ent. Soc., Trans. 4: 212. 9.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 156-157, fig. 4 (sleeping aggregation).
—Cazier and Mortenson, 1965. Pan-Pacifie Ent. 41: 34-43, 6 figs. (nesting behavior, prey,
parasite).

Genus ISODONTIA Patton

This is the only non-fossorial genus among our native Sphecinae.

1582 Hymenoptera in America North of Mexico
Genus ISODONTIA Subgenus ISODONTIA Patton

Isodontia Patton, 1881. Boston Soe. Nat. Hist. Proe. 20: 380.
Type-species: Sphex philadelphica Lepeletier. Orig. desig.
Leontosphex Arnold, 1945. Sphecidae of Madagascar, p. 90.
Type-species: Sphex leoninus Saussure. Monotypic.
apicalis (Smith). N. J. to Fla., west to Tex., Tenn., Nebr., Ariz.
Sphex apicalis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 262. 9.
Isodontia macrocephala var. cinerea Fernald, 1903. Canad. Ent. 35: 271.
Chlorion (Isodontia) harrisi Fernald, 1906. U. S. Natl. Mus., Proc. 31: 359. N. name.

exornata Fernald. N. C. to Fla., west to Tex.
Isodontia exornata Fernald, 1903. Canad. Ent. 35: 270. °.
philadelphica (Lepeletier). Eastern states, N. Y. to Kans., south to Fla. and Tex., Ariz., Calif;
Mexico. Ecology: Nests in cavities in log, and in rotten limb. Parasite: Amobia
floridensis (Tns.). Prey: Orocharis sp.
Sphex philadelphica Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 340. 2.
Sphex (Isodontia) macrocephalus Fox, 1890. Ent. News 1: 137. 2.
Spheax aztecus var. digueti Berland, 1927. Paris Mus. d’Hist. Nat., Bul. 32: 283. °.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 148, figs. 29, 30 (larva; misdet.
as azteca Sauss.).

Biology: Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 135 (nest). —Krombein, 1967.
Trap-nesting wasps and bees, p. 239 (nest).

Genus ISODONTIA Subgenus MURRAYELLA Bohart and Menke

Isodontia Subg. Murrayella Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 137.
Type-species: Sphex elegans Smith. Orig. desig.

Species of this subgenus are sometimes called “grass carrier” wasps because they use grass
stems and blades to form partitions between cells and to make the closing plug in their nests in
cavities in wood, plants and even abandoned bee burrows in clay banks. Other plant materials,
such as the fibrous inner bark of certain trees and Spanish moss, are also used in nest construc-
tion. Our three species show an interesting evolutionary development in nest structure; elegans
always makes individual cells separated by partitions, mexicana sometimes makes similar nests
but usually constructs just one larger brood chamber in which several larvae develop without
cannibalism, and awripes always makes a nest in which there is just a brood chamber. Our spe-
cies provision their nests with both Gryllidae and Tettigoniidae, but gryllids of the genus
Oecanthus are preferred when they are available.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 239-251, figs. 57-61 (U. S. spp.).
auripes (Fernald). N. Y. to Fla., west to Mich., Kans. and Tex. Ecology: Nests in borings in

wood and abandoned bee burrows in clay banks; nest contains only a single large brood
cell in which several larvae are reared. Parasite: Pseudoxenos auripedis (Pierce);
Anthrax aterrimus (Big.); Miltogrammini sp.; Phoridae sp.; Melittobia chalybui Ashm.;
Chrysididae sp. Prey: Neoxabea bipunctata (DeG.), Oecanthus exclamationis Davis, ?
O. angustipennis Fitch, O. latipennis Riley, O. sp.; Orocharis saltator Uhl., O. luteolira
Wlkr., O. vulgare Harr., O. sp., Conocephalus memorale Seud., C. sp., Scudderia sp.
Predator: Lecontella cancellata (LeC.).

Sphex tibialis Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 3389. 2. Preoce.

Chlorion (Isodontia) auripes Fernald, 1906. U.S. Natl. Mus., Proc. 31: 356. N. name.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 144, figs. 31, 32 (larva).
—Evans, 1959. Amer. Ent. Soc., Trans. 85: 147-148 (larva).

Biology: Packard, 1869. Guide Study Ins., pp. 168-169 (nest, cocoon). —Rau and Rau, 1918.
Wasp studies afield, pp. 203-205 (nest). —Rau, 1926. St. Louis Acad. Sci., Trans. 25:
200-201 (nest). —Rau, 1928. St. Louis Acad. Sci., Trans. 25: 362-368 (prey, parasite).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 246-251, ? figs. 60, 61 (nest, prey, life
cycle, parasites). —Krombein, 1970. Smithsn. Contrib. Zool. 46: 3-12, figs. 6-39 (nesting
behavior, prey, life cycle).

Superfamily SPHECOIDEA 1583

elegans (Smith). B. C., U. S. west of 100th Meridian. Ecology: Nests in borings in wood or
stems, and in abandoned borings of mining bees, each cell being separated by a
partition. Parasite: Sphaeropthalma ferruginosa (D. T.), S. wnicolor (Cr.); Epistenia
caeruleata Westw.; Amobia floridensis (Tns.). Prey: Oecanthus quadripunctatus Beut.,
O. c. californicus Sauss., O. niveus (DeG.), O. spp.; Dichopetala sp., ? Eremopedes sp.; all
nymphs.
Sphex elegans Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 262. ¢.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 238-239, figs. 18, 16 (larva).

Biology: Ashmead, 1894. Psyche 7: 64 (prey). — Davidson, 1899. Ent. News 10: 179-180 (nest,
prey, parasites). —Fernald, 1906. U.S. Natl. Mus., Proc. 31: 364 (nest, prey, cocoon).
—Ainslie, 1924. Canad. Ent. 56: 269-270 (nest, prey). —Parker and Bohart, 1966.
Pan-Pacifie Ent. 42: 94 (nest, parasites). —Krombein, 1967. Trap-nesting wasps and bees,
pp. 240-242, fig. 57 (nest, prey, life cycle).

mexicana (Saussure). U.S. east of Rockies, Ariz.; Mexico, Central America; introduced into

Hawaii, France. Ecology: Nests in borings in wood, hollow stems, pitcher plants and
glass tubes; nest usually contains a single large brood cell, but occasionally cells of
individual larvae are separated by flimsy partitions. Parasite: Amobia distorta (Wulp),
Senotainia trilineata (Wulp), S. sp., Sarcophaga sp.; Megaselia aletiae (Comst.);
Eustalomyia vittipes (Zett.). Prey: Oecanthus angustipennis Fitch, O. quadripunctatus
Beut., O. argentinus Sauss., O. fultoni Wlkr., O. nigricornis Wlkr., O. niveus (DeG.), O.
fasciatus Fitch, O. spp., Gryllus assimilis F., Neoxabea bipunctata (DeG.); Orocharis
saltator Uhl., Odontoxiphidium apterum Morse, Conocephalus fasciatus DeG., C. spp.,
Neoconocephalus sp., Orchelimum sp., Rehnia spinosa Caud.; both adults and nymphs
are stored. Predator: Crematogaster sp.

Sphex apicalis Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p. 588.
Nom. nud.

Sphex apicalis Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 38. Preoce.

Sphex apicalis var. mexicana Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 38. 2,

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 143, figs. 23-28 (larva; misdet.
as harrisi Fern.). —Evans, 1964. Amer. Ent. Soc., Trans. 90: 239-240 (larva).

Biology: Ashmead, 1895. U. S. Dept. Agr., Insect Life 7: 241 (prey). —Jones, 1904. Ent. News
15: 14-17, 2 pls. (nest, prey). —Engelhardt, 1929. Brooklyn Ent. Soc., Bul. 23: 269-271 (nest,
prey). —Rau, 1935. Brooklyn Ent. Soc., Bul. 30: 65-68, 1 pl. (nest, prey, life cycle).
—Suehiro, 1937. Hawaii. Ent. Soc., Proc. 9: 358 (nest, life cycle). —Rau, 1943. Ent. Soe.
Amer., Ann. 36: 648 (nest). —Swezey, 1947. Hawaii. Ent. Soc., Proc. 13: 8 (nest). —Lin,
1962. Tex. Jour. Sci. 14: 429-430 (nest, prey, life cycle). —Medler, 1965. Ent. Soc. Amer.,
Ann. 58: 137-142, 4 figs. (nest, prey, life cycle, cocoon, parasites). —Lin, 1966. Wasmann
Jour. Biol. 24: 239-247, 2 figs. (nest, prey, life cycle). —Krombein, 1967. Trap-nesting wasps
and bees, pp. 242-246, figs. 58, 59 (nest, prey, life cycle, predator).

TRIBE PRIONYXINI

Genus PALMODES Kohl

Palmodes Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 112.
Type-species: Sphex occitanica Lepeletier and Serville, Desig. by Fernald, 1906.

So far as known these are solitary ground-nesting wasps except for carbo which has been re-
ported once as nesting gregariously. Most of our species prey upon Tettigoniidae except for
carbo which uses Gryllacrididae.

Revision: Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 179-191, 17 figs.

californicus Bohart and Menke. Calif., Nev., Oreg., B. C. Prey: Immature adult, Platylyra
californica Seud., Neduba morsei Caud.
Palmodes californicus Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 182, figs. 3, 13.
Gree

1584 Hymenoptera in America North of Mexico

carbo Bohart and Menke. B. C. to Calif., east to N. W. T., Mont., Colo. and N. Mex. Ecology:
Nests gregariously in sand with one cell per nest. Prey: Cyphoderris monstrosa Uhl.,
nymphs. Predator: Philanthus zebratus nitens (Bks.).
Sphex morio Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 321. d. Preoce.
Palmodes carbo Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 144. N. name.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 483-484 (nest, prey).
dimidiatus (DeGeer). U. S. except northwest; Mexico (Chihuahua, Coahuila). Ecology: Nests in

sand, the burrow short and oblique, the cell stored with one prey specimen. Prey:
Atlanticus pachymerus (Burm.), A. sp., Dissosteira carolina (L.), Pediodectes sp.,
probably stevensonii (Thom.).

Sphex dimidiatus DeGeer, 1773. Mem. pour servir a I’Hist. des Ins. 3: 577, pl. 30, fig. 5. ¢.

Sphex violaceipennis Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 349.

Sphex rufiventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 211. 2.

Sphex abdominalis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 211. d. Preoce.

Chlorion (Palmodes) rufiventris var. opuntiae Rohwer, 1911. U.S. Natl. Mus., Proc. 40:
557. 2.

Sphex (Palmodes) daggyi Murray, 1951. U.S. Dept. Agr., Monog. 2: 974. N. name.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 141, figs. 12-14 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2, Sci. Ser. 1:
174-175, pl. 2, fig. 1 (nest, prey). —Williams, 1913. Kans. Univ. Sci. Bul. 8: 227 (prey).
—Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 281-282 (nest, prey, life cycle).
—Krombein, 1955. Ent. Soc. Wash., Proc. 57: 150-151 (nest, prey, life cycle, cocoon).

hesperus Bohart and Menke. Calif. to B. C., Nev., Utah, Wyo., Colo. Ecology: Nests in sandy
loam, one cell per nest. Prey: Anabrus simplex Hald., nymph.
Palmodes hesperus Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 184, fig. 6. 5, °.

Biology: Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 185 (prey). —Evans, 1970. Mus.
Compar. Zool., Bul. 140: 484 (nest, prey).
insularis Bohart and Menke. Calif. (Channel Is.).
Palmodes insularis Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 186. d, @.
laeviventris (Cresson). A Great Basin species, Mont., Wyo. and Colo. west to Wash. and
eastern Calif. Ecology: Makes a 1-celled nest and stores 2-4 crickets in it. Parasite:
Sphenometopa tergata (Coq.); Stizoides unicinctus (Say). Prey: Anabrus simplex Hald.,
Pediodectes sp., probably stevensonii (Thos.).
Sphex laeviventris Cresson, 1865. Ent. Soc. Phila., Proc. 4: 463. 2, ¢.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 227 (prey). —LaRivers, 1945. Amer.
Midland Nat. 33: 743-763 (nest, prey, parasites).
lissus Bohart and Menke. Southern Calif., Ariz., Tex.
Palmodes lissus Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 187, figs. 5, 12. d, 9.
pacificus Bohart and Menke. Coastal Calif.
Palmodes pacificus Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 188, figs. 8, 9, 14.
Gq Pe
praestans (Kohl). Oreg., Calif., Nev., Utah, Ariz., N. Mex., west. Tex.; Mexico (Coahuila). Prey:
Capnobates fuliginosus Thom.
Sphex (Palmodes) praestans Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 323. @.

Biology: Caudell, 1919. Ent. Soc. Wash., Proc. 21: 40 (prey).
stygicus Bohart and Menke. Calif., Nev., Utah, Ariz., N. Mex., a Great Basin species.

Palmodes stygicus Bohart and Menke, 1961. Ent. Soc. Wash., Proc. 63: 191, figs. 1, 11, 16.
On er

Genus PRIONYX Vander Linden

Prionyx Vander Linden, 1827. Nouv. Mem. Acad. Roy. Sci. Bruxelles 4: 362.
Type-species: Ammophila kirbii Vander Linden. Monotypic.
Priononyx Dahlbom, 1843. Hym. Europaea, v. 1, p. 28.
Type-species: Sphex thomae Fabricius. Monotypic.

Superfamily SPHECOIDEA 1585

Enodia Dahlbom, 1843. Hym. Europaea, v. 1, p. 28. Preoce.
Type-species: Sphex albisectus Lepeletier and Serville. Desig. by Kohl, 1885.
Harpactopus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 264.
Type-species: Harpactopus crudelis Smith. Desig. by Patton, 1881.
Parasphex Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 267.
Type-species: Sphex albisectus Lepeletier and Serville. Desig. by Kohl, 1885.
Gastrosphaeria Costa, 1858. Fauna Regn. Napoli, Imen. Acul., Sphecidea, p. 10.
Type-species: Gastrosphaeria anthracina Costa. Monotypic.
Pseudosphex Taschenberg, 1869. Ztschr. Gesam. Naturw. Halle 34: 420. Preoce.
Type-species: Pseudosphex pumilio Taschenberg. Monotypic.
Calosphex Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 113.
Type-species: Sphex niveatus Dufour. Desig. by Pate, 1937.
Neosphex Reed, 1894. Ann. Univ. Chile 85: 627.
Type-species: Neosphex albospiniferus Reed. Monotypic.

Our North American species dig simple, shallow unicellular nests in a variety of soil types.
The grasshopper prey is captured before the nest is dug, although some extralimital species are
known to prepare the nest before capturing the single prey specimen.

Taxonomy: Parker, 1960. Pan-Pacific Ent. 36: 205-208, 1 pl. (key to N. A. spp.).

Biology: Evans, 1958. Ent. Soc. Amer., Ann. 51: 177-186, 3 figs. (nesting behavior).
atratus (Lepeletier). Transcont. in southern Canada and all of U. S.; Mexico (Durango).

Ecology: Nests in a variety of soils, the burrow varying from L-shaped to curved to
oblique; nest has a single cell and is provisioned with one prey specimen. Parasite:
Pseudoxenos duryi (Pierce); Metopia argyrocephala (Meig.), Senotainia sp.; Stizoides
unicinctus (Say). Prey: Ageneotettix d. deorum Sceud., Aulocara elliotti Thom., Mermiria
neomexicana Thom., Arphia xanthoptera Burm., Dissosteira carolina L., Pardalophora
phoenicoptera Burm., Spharagemon collare Scud., Trimerotropis citrina Scud.,
Melanoplus angustipennis Dodge, M. arizonae Scud., M. bispinosus Seud., M. bivittatus
Say, M. devastator Seud., M. differentialis Thom., M. femurrubrum DeG., M. foedus
Seud., M. lakinus Seud., M. spretus Walsh, M. spp., Schistocerca lineata Scud. Predator:
Apiomerus spissipes (Say).

Sphex labrosa Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p. 588.
Nom. Nud.

Sphex atrata Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 355. °.

Priononyx brunnipes Cresson, 1872. Amer. Ent. Soc., Trans. 4: 213. d.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 142, figs. 15-20 (larva). —Evans,
1959. Amer. Ent. Soc., Trans. 75: 147 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 171-174 (nest,
prey). —Bradley, 1908. Ent. Soc. Amer., Ann. 1: 128-129 (sleeping aggregation).
—Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 227-230, pl. 33, fig. 2 (nest, prey, parasite).
—Rau and Rau, 1918. Wasp studies afield, pp. 159-175, figs. 36-39 (nest, prey, parasite).
—Rau, 1922. St. Louis Acad. Sci., Trans. 24: 23 (prey). —Strandtmann, 1945. Ent. Soc.
Amer., Ann. 38: 308-310 (nest). —Evans, 1958. Ent. Soc. Amer., Ann. 51: 178-181, fig. 1
(prey transport, nest, parasite, life cycle).

canadensis (Provancher). B. C. to Man., south to Calif., Ariz., Colo. and Nebr., Ont., N. J., Va.
Priononyx Canadensis Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym,, p. 258. °.
Sphex excisus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 362. 3.
fervens (Linnaeus). Calif. to Tex., south to South America, West Indies. Ecology: Nests in
sand, the burrow oblique, the cell horizontal and provisioned with one prey specimen.
Prey: Xyleus sp., probably centralis Rehn, Schistocerca cancellata (Serv.).
Sphex fervens Linnaeus, 1758. Syst. Nat., ed. 10, p. 569. 2. Type from West Indies or
Surinam, not Indies.
Pepsis Johannis Fabricius, 1804. Systema Piezatorum, p. 208. 2.
Sphex Doumerci Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 357. °.
Priononyz striata Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 266.
Sphex (Priononyx) laerma Cameron, 1897. Ann. and Mag. Nat. Hist. (6) 19: 370.

1586 Hymenoptera in America North of Mexico

Biology: Evans, 1958. Ent. Soc. Amer., Ann. 51: 184, figs. 2-3 (nest, prey).
foxi Bohart and Menke. Tex., Nev., Utah; northern Mexico.
Sphex (Priononyx) ferrugineus Fox, 1892. Ent. News 3: 170. ?. Preoce.
Prionyx foxi Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 152. N. name.
parkeri Bohart and Menke. U. S. south to Isthmus of Tehuantepec. Ecology: Nests in a variety
of soils, the burrow varying from oblique to nearly vertical, the cell provisioned with
one prey specimen. Parasite: Senotainia rubriventris Macq. Prey: Melanoplus scudderi
Uhl., M. femurrubrum propinquus Seud., M. sp., probably femurrubrum (DeG.),
Trimerotropis citrina Seud., Scirtetica marmorata picta (Seud.), Chortophaga australior
R. and H.
Prionyx parkeri Bohart and Menke, 1963. Univ. Calif. Pubs. Ent. 30: 154, figs. 35, 54, 96,
102: ¢, ¢.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 23 (nest, prey; misdet. as bifoveolatum
Tasch.). —Evans, 1958. Ent. Soc. Amer., Ann. 51: 183-184 (nest, prey, parasite; misdet. as
pubidorsus Costa). —Linsley, 1962. Ent. Soc. Amer., Ann. 55: 156, fig. 4 (sleeping
aggregation; misdet. as pubidorsum). —Krombein, 1964. Amer. Mus. Novitates 2201:
18-19 (nest, prey).

subatratus Bohart. Oreg. and Idaho south and east to Calif., Utah, Ariz., N. Mex., western

Tex.; Mexico (Chihuahua).

Priononyx subatrata Bohart, 1958. Brooklyn Ent. Soc., Bul. 53: 90. d, 9.

thomae (Fabricius). Southeastern and western states, south to Argentina. Ecology: Nests in a

variety of soils, the burrow varying from L-shaped to curved to oblique; nest with a
single cell and provisioned with one prey specimen. Parasite: Stizoides wnicinctus (Say).
Prey: Amphitornus sp., Aulocara sp., Orphulella p. pelidna Burm., Arphia xanthoptera
Burm., Dissosteira carolina L., Encoptolophus subgracilis texensis Br., Paraidemona
sp., probably fratercula Heb.

Sphex thomae Fabricius, 1775. Systema Ent., p. 346. d.

Pepsis crucis Fabricius, 1804. Systema Piezatorum, p. 209. °.

Enodia pubidorsum Costa, 1862. Mus. Zool. Napoli, Ann. 1: 69. 3.

Priononyx thomae var. antillarwm Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p.
43. 2.

Priononyx thomae var. mexicanus Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p.
43. 2, 6.

Sphex (Harpactopus) Edwardsi Cameron, 1903. Amer. Ent. Soc., Trans. 29: 230. 2, ¢.

Sphex platensis Brethes, 1908. Buenos Aires Mus. Nac. de Hist. Nat., An. 17: 146. 6, 2.

Sphex thomae var. altibia Strand, 1911. Arch. f. Naturgesch. 77, sup. 2: 152. ¢. This is a
questionable synonym.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 142, figs. 21, 22 (larva).

Biology: Hartman, 1905. Tex. Univ., Bul. 65, Sci. Ser. 6: 62-65 (nest, prey, life cycle).
—Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 227 (prey, sleeping aggregation). —Rau
and Rau, 1918. Wasp studies afield, pp. 175-186, figs. 40-42 (nest, prey, parasite). —Evans,
1958. Ent. Soe. Amer., Ann. 51: 181-183 (nest, prey transport, life cycle).

UNPLACED TAXON OF SPHECINAE

Sphex instabilis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 263. 2. N. Amer. Possibly a senior
syn. of Isodontia exornata Fern.

SUBFAMILY AMMOPHILINAE

The North American genera are all fossorial in nesting habits, but one extralimital genus is
known to nest in cavities in wood.

Taxonomy: Menke, 1966. Canad. Ent. 98: 147-152, 12 figs. (key to genera).
Genus PODALONIA Fernald

Psammophila Dahlbom, 1842. Dispos. Method. Spee. Scand. Ins. Hym., pt. 1, pp. 2, 8.
Preoce.

Superfamily SPHECOIDEA 1587

Type-species: Ammophila affinis Kirby. Desig. by Fernald, 1927.
Podalonia Fernald, 1927. U. S. Natl. Mus., Proe. 71 (9): 11.
Type-species: Ammophila violaceipennis Lepeletier. Desig. by Internat]. Comn.
Zool. Nomencl., Op. 857, 1968. Op. 857 also suppressed Podalonia Spinola, 1853
and validated Podalonia Fernald, 1927.

These wasps are fossorial and construct a single cell at the end of a short burrow. They prey
upon caterpillars, usually those of the soil-burrowing cutworm type, and place only one prey
specimen in the cell.

Revision: Melander, 1903. Psyche 10: 156-164. —Fernald, 1927. U. S. Natl. Mus., Proe. 71 (9):
1-42. —Murray, 1940. Ent. Amer. (n. s.) 20: 1-82.

Taxonomy: Menke, Bohart and van der Vecht, 1966. Bul. Zool. Nomencl. 23: 48-51 (request
for suppression of Podalonia Spinola, 1853, validation of Podalonia Fernald, 1927, and
desig. of Ammophila violaceipennis Lepeletier as type-species).

argentifrons (Cresson). West. States and Provinces. Parasite: Pseudoxenos luctuosae (Pierce).
Ammophila argentifrons Cresson, 1865. Ent. Soe. Phila., Proc. 4: 462. ¢.

argentipilis (Provancher). Ariz., Calif.
Pelopoeus argentipilis Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 256. °.
Ammophila morrisoni Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 21. ¢.
Psammophila nicholi Carter, 1924. Ent. News 35: 366. d.

caerulea Murray. Calif., Idaho.
Podalonia caerulea Murray, 1940. Ent. Amer. (n. s.) 20: 67. 6.

clypeata Murray. West. States to Minn.
Podalonia clypeata Murray, 1940. Ent. Amer. (n. s.) 20: 49. d, 2.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 144, figs. 33-38 (larva).
communis communis (Cresson). U. S. west of 100th meridian; Mexico. Ecology: Nests in sand.
Parasite: Hilarella hilarella Zett., Metopia argyrocephala (Meig.). Prey: Noctuidae spp.,
larvae. Predator: Philanthus zebratus nitens (Bks.). Other subspp. occur in Mexico and
Central America.
Ammophila communis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 462. 6.

Biology: Newcomer, 1930. Ent. Soc. Amer., Ann. 23: 552-563 2 pls. (nest, prey, parasite, life
cycle). —Hicks, 1931. South. Calif. Acad. Sci., Bul. 30: 75-82, pls. 22-25 (nest, prey).
—Hicks, 1931. Pan-Pacifie Ent. 8: 49-51 (hibernation). —Hicks, 1932. Psyche 39: 150-154
(nest, prey, parasite). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 484-485 (nest, prey,
parasites, predator). The observations by both Newcomer and Hicks on luctwosa were
made on both luctwosa and c. communis.

compacta Fernald. Calif., Oreg.
Podalonia violaceipennis var. compacta Fernald, 1927. U.S. Natl. Mus., Proce. 71 (9): 33.
OeGe
luctuosa (Smith). Transcont. in northern tier of States and Canada, as far north as N. W. T.
and Yukon. Ecology: Nests in sand. Parasite: Pseudoxenos luctuosae (Pierce); Hilarella
hilarella (Zett.), Metopia argyrocephala (Meig.), Taxigramma heteroneura (Meig.). Prey:
Lycophotia saucia Hbn., L. margaritosa Haw. ?, Chorizagrotis agrestis Grt., Noctuidae
spp. Predator: Philanthus zebratus nitens (Bks.).
Ammophila luctuosa Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 224. 2.
Psammophila pacifica Melander and Brues, 1902. Biol. Bul. 3: 40. ¢.

Biology: Newcomer, 1930. Ent. Soc. Amer., Ann. 23: 552-563, 2 pls. (nest, prey, parasite, life
cycle). —Hicks, 1931. South. Calif. Acad. Sci., Bul. 30: 75-82, pls. 22-25 (nest, prey).
—Hicks, 1931. Pan-Pacifie Ent. 8: 49-51 (hibernation). —Hicks, 1932. Psyche 39: 150-154
(nest, prey, parasite). The observations by both Newcomer and Hicks on luctuosa were
made on both luctwosa and c. communis.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99, no. 14: 50, pl. 22, figs. O, P, S,
T (male genitalia).

melaena Murray. West. States.
Podalonia melaena Murray, 1940. Ent. Amer. (n. s.) 20: 34. d, 2.

1588 Hymenoptera in America North of Mexico

mexicana (Saussure). West. States and Provinces; Mexico.

Ammophila mexicana Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 25. d, 2.
mickeli Murray. West. Provinces and States to Minn.

Podalonia mickeli Murray, 1940. Ent. Amer. (n. s.) 20: 68. 6, 9.
occidentalis Murray. Alta., West. States. Prey: Tent-caterpillar.

Podalonia occidentalis Murray, 1940. Ent. Amer. (n. s.) 20: 54. d, °.

Biology: Murray, 1940. Ent. Amer. (n. s.) 20: 13 (prey).
parallela Murray. Calif.
Podalonia parallela Murray, 1940. Ent. Amer. (n. s.) 20: 65. 6, °.
pubescens Murray. Tex. to Ariz.; Mexico.
Podalonia pubescens Murray, 1940. Ent. Amer. (n. s.) 20: 47. d, °.
puncta Murray. Kans., Colo., Okla., Tex., N. Mex.
Podalonia puncta Murray, 1940. Ent. Amer. (n. s.) 20: 36. d, 9.
robusta (Cresson). Transcont. in Canada and U. S., as far north as N. W. T. and Yukon; south
to Costa Rica. Prey: Acronyctinae sp. ?
Ammophila robusta Cresson, 1865. Ent. Soc. Phila., Proc. 4: 461. 9.

Biology: Krombein, 1936. Ent. News 47: 93-99 (nest, prey; misdet. as violaceipennis).
sericea Murray. West. Provinces and States east to the Dakotas, very rare to Mich. Ecology:
Nests along dirt road. Prey: Zale lunata (Dru.), Homoptera salicis Behr., Noctuidae sp.
Podalonia sericea Murray, 1940. Ent. Amer. (n. s.) 20: 57. d, 2.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 240-241, figs. 5, 8 (larva; misdet. as
robusta).

Biology: Hicks, 1933. Pan-Pacific Ent. 9: 49-52 (prey; misdet. as violaceipennis). —Evans,
1963. Ent. News 74: 287, fig. 4 (nest, prey; misdet. as robusta). —Evans, 1970. Mus.
Compar. Zool., Bul. 40: 485, fig. 18 (nest, prey).

sonorensis (Cameron). Alta., U. S. west of 100th meridian; Mexico.
Ammophila sonorensis Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 21. d, @.
Podalonia sonorensis differentia Murray, 1940. Ent. Amer. (n. s.) 20: 33. 6, °.
valida (Cresson). West. Provinces and States east to Minn. Ecology: Nests in bare or densely
vegetated soil, makes cluster of unicellular nests in a restricted site, digs burrow before
hunting prey. Parasite: Bombyliidae sp. probably Ligura or Exoprosopa. Prey:
Estigmene acraea (Dru.).
Ammophila valida Cresson, 1865. Ent. Soc. Phila., Proc. 4: 461. 2.
Ammophila grossa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 209. °.

Biology: Steiner, 1974. Pan-Pacifie Ent. 50: 73-77, 1 fig. (prey hunting and transport).
—Steiner, 1975. Quaestiones Ent. 11: 113-127, 6 figs. (female territorial behavior, nest,
prey, parasite).

violaceipennis (Lepeletier). East. States west to Colo. Ecology: Nests in sand or heavier soil.
Parasite: Pseudoxenos luctuosae (Pierce); Hilarella sp. Prey: Symmerista albifrons S.
and A., Noctuidae sp.
Ammophila violaceipennis Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 370. 9.
Ammophila cementaria Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 224. °.

Biology: Parker, 1915. Ent. Soc. Wash., Proc. 17: 70-77 (nest, prey, parasite, life cycle).
—Balduf, 1936. Canad. Ent. 68: 137-188 (prey).

Genus EREMNOPHILA Menke

Ammophila subg. Eremnophila Menke, 1964. Canad. Ent. 96: 875.
Type-species: Ammophila opulenta Guerin. Orig. desig.
aureonotata (Cameron). Southern Canada and U.S. east of 100th meridian, south to El
Salvador. Ecology: Nests in sand or hard-packed loam, the burrow vertical, terminating
in a horizontal cell provisioned with a single caterpillar. Parasite: Pseudoxenos lugubris
(Pierce). Prey: Heterocampa guttivitta (Wlkr.), H. sp.; Hesperiidae sp.
Ammophila aureonotata Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 7. 2, 3.

Superfamily SPHECOIDEA 1589

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 25-28 (prey
transport, nest; misdet. as gracilis Lep.). —Peckham and Peckham, 1905. Wasps, social and
solitary, pp. 43-46 (prey transport; misdet. as gracilis Lep.). —Rau, 1922. Acad. Sci. St.
Louis, Trans. 24: 24 (mating, prey). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 104 (nest,
prey transport). —Evans, 1959. Amer. Midland Nat. 62: 464-465 (nest, prey transport).

Genus AMMOPHILA Kirby

Ammophila Kirby, 1798. Linn. Soc. London, Trans. 4: 199.
Type-species: Sphex sabulosa Linnaeus. Desig. by Internat]. Comn. Zool. Nomencl.
Op. 180, 1946. Op. and Decl. by Internatl. Comn. Zool. Nomencel. 2: 569-585.
Ammophylus Latreille, 1802-1803. Hist. Nat. Crust. Ins. 3: 332. Emend. or lapsus.
Miscus Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 130. No species included.
Type-species: Ammophila campestris Latreille. Desig. by Shuckard, 1837.
Ammophilus Latreille, 1829. In Cuvier, Regn. Anim., v. 4, p. 322. Emend. or lapsus.
Coloptera Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 387.
Type-species: Coloptera barbara Lepeletier. Monotypic.
Argyrammophila Gussakovskij, 1928. Leningrad, Inst. Zool. Appl. Phytopath., Bul. 4: 7.
Type-species: Ammophila induta Kohl. Orig. desig.
Apycnemia Leclercq, 1961. Eos 37: 211.
Type-species: Ammophila fallax Kohl. Orig. desig.

The behavior of these slender, elongate wasps has been studied by a host of observers. All
species are fossorial, usually solitary in nesting habits, although a few extralimital species are
gregarious nesters. Typically, the nest consists of a short, oblique to perpendicular burrow ter-
minating in a single cell, although 2-celled nests have been reported in two species. Usually, the
nest is dug before prey is obtained, but two species have been reported as capturing prey before
constructing the nest.

The prey used by Ammophila consists usually of hairless larvae although sparsely to densely
haired larvae may be used occasionally. Lepidopterous larvae are frequently provided, but occa-
sionally hymenopterous (sawfly) larvae are used. There is one record of weevil larvae being
preyed upon by azteca Cam. which also uses lepidopterous and sawfly larvae. It is probable that
caterpillars are the preferred prey, and that sawfly or weevil larvae are used only when there is
not a ready supply of caterpillars. Wasps using larger larvae for prey, store only one per nest;
those using smaller larvae may provide as many as 11 per cell. Most species are mass provi-
sioners, but several practice progressive provisioning. Three species, including our native azteca,
are known to maintain several nests simultaneously which are provisioned progressively.

The Peckhams observed one specimen of wrnaria Dahlb. using a pebble to tamp the earth in
the nest closure. They contended that this constituted improvisation of a tool and intelligent use
of it. Subsequent observations on other species demonstrate conclusively that the supposed
tool-using behavior is not an intelligent act but the culmination of a succession of instinctive
behavioral traits.

Revision: Melander, 1903. Psyche 10: 156-164. —Fernald, 1934. No. Amer. and W. Indies
Sphex, 167 pp. These are not reliable for identification of many North American species.

Biology: Evans, 1959. Amer. Midland Nat. 62: 449-473 (nesting behavior). —Powell, 1964.
Kans. Ent. Soc., Jour. 37: 240-258 (nesting behavior). —Menke, 1965. Ent. News 76:
257-261 (identity of spp. studied by Hicks and Evans).

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99, no. 14: 50, pl. 23, figs. K, M-R
(male genitalia).
aberti Haldeman. West. Provinces and States east to Iowa; Mexico. Ecology: Nests in firm
sand or hard soil, the vertical or oblique burrow terminating in a cell in which up to 10
small caterpillars are stored. Parasite: Pseudoxenos lugubris (Pierce); Hilarella
hilarella (Zett.), Metopia argyrocephala (Meig.), Opsidia sp.; ? Exoprosopa capucina
(F.); Ceratochrysis trachypleura Boh. Prey: Euchlaena sp., Sterrhinae sp., Geometridae
spp.; Hesperiidae sp.
Ammophila aberti Haldeman, 1852. Stansburys Explor. Survey Salt Lake, App. C, Ins., p.
368. 2.
Ammophila urnaria Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 381. 2. Preoce.

1590 Hymenoptera in America North of Mexico

Ammophila tarsata Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 219.

Ammophila yarrowi Cresson, 1875. Rpt. Geog. Geol. Explor. and Survey west of 100th
meridian, v. 5, p. 713. 6. ;

Sphex transversus Fernald, 1934. No. Amer. and W. Indies Sphex, p. 141. d.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 148, figs. 31-33 (larva).

Biology: Williston, 1892. Ent. News 3: 85-86 (nest, prey). —Hicks, 1932. Canad. Ent. 64:
145-151 (nest, prey, parasites; Chrysis perpulchra Cr. misdet.). —Evans, 1959. Amer.
Midland Nat. 62: 454-456 (nest, prey, parasite). —Powell, 1964. Kans. Ent. Soc., Jour. 37:
244-251 (nest, prey, parasite, mating, life cycle).

acuta (Fernald). Western U. S.
Sphex acutus Fernald, 1934. No. Amer. and W. Indies Sphez, p. 150. 6.
aphrodite Menke. Ariz., Nev., Calif.
Ammophila aphrodite Menke, 1964. Acta Hym. 2: 8, fig. 2. 5, 2.
azteca azteca Cameron. Transcont. in Canada and U. S., ranging as far north as N. W. T. and
Yukon; Mexico. Ecology: Nest is a vertical burrow ending in a single cell; a female can
maintain several nests simultaneously, provisioning each progressively with up to seven
prey specimens. Prey: Lycaenidae spp.; Gecmetridae spp.; Gelechiidae sp.;
Pterophoridae sp.; Smerinthus sp.; Nematus sp., Amauronematus sp.; Hypera postica
(Gyll.). Predator: Philanthus zebratus nitens (Bks.).
Ammophila azteca Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 17. 2.
Sphex pilosus Fernald, 1934. No. Amer. and W. Indies Sphex, p. 120. °, 6.
Sphex aculeatus Fernald, 1934. No. Amer. and W. Indies Sphex, p. 145. 2, 6.
Sphex pilosus nudus Murray, 1938. Ent. Soc. Amer., Ann. 31: 28. d, 2. Preoce.
Ammophila pilosa brevisericea Murray, 1951. U. S. Dept. Agr., Agr. Monog. 2: 976. N.
name.

Biology: Hicks, 1933. Canad. Ent. 65: 49-51 (nest, prey, parasite; misdet. as breviceps Sm.).
—Hicks, 1935. Pan-Pacifie Ent. 11: 99-101 (nest; misdet. as aculeatus Fern.). —Evans,
1963. Ent. News 74: 238, fig. 5 (nest, prey). —Powell, 1964. Kans. Ent. Soc., Jour. 37: 244
(prey). —Evans, 1965. Psyche 72: 8-23, 9 figs. (nesting behavior, prey, life cycle).

azteca clemente Menke. Calif. (San Clemente Is.).
Ammophila azteca clemente Menke, 1967. Los Angeles Co. Mus., Contrib. Sci. 123: 7. 6, @.
bella Menke. Ariz.; Mexico (Sonora, Sinaloa, Guerrero, Puebla).
Ammophila bella Menke, 1966. Biol. Soc. Wash., Proc. 79: 27. d, 2.
bellula Menke. Ariz., N. Mex.; Mexico.
Ammophila bellula Menke, 1964. Acta Hym. 2: 17, fig. 10. 3, °.
boharti Menke. Calif., Nev.
Ammophila boharti Menke, 1964. Acta Hym. 2: 9, fig. 7. d, 9.
breviceps Smith. Southwestern U. S.; Mexico. Parasite: Pseudoxenos lugubris (Pierce);
Spintharosoma mesillae (Ckll.).
Ammophila breviceps Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 221. °.
californica Menke. Western U. S.
Ammophila californica Menke, 1964. Acta Hym. 2: 18, fig. 16. 6, @.
centralis Cameron. Tex. (Cameron Co.) south to Guatemala.
Ammophila centralis Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 6, pl. 1, fig. 12. ¢.
Ammophila consors Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 12, pl. 2, fig. 3. d, @.
Ammophila nigro-caerulea Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 12, pl. 1, fig. 8.
Os
cleopatra Menke. Transcont. in U. S.; Mexico. Ecology: Nests in sand, the vertical burrow
terminating in a horizontal cell containing one or two prey. Parasite: Senotainia litoralis
Allen. Prey: Macrurocampa marthesia (Cram.).
Ammophila cleopatra Menke, 1964. Acta Hym. 2: 19, fig. 12. 3, 2.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soe., Trans. 81: 147, figs. 60-62 (larva; misdet.
as juncea Cr.).

Superfamily SPHECOIDEA 1591

Biology: Evans, 1959. Amer. Midland Nat. 62: 461-462 (nest, prey, parasite; misdet. as juncea
Cr.).
coachella Menke. Southern Calif., deserts.
Ammophila coachella Menke, 1966. Biol. Soc. Wash., Proc. 79: 38, figs. 2, 5,6. d, 2.
conditor Smith. Fla.
Ammophila conditor Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 223. 2.
dejecta Cameron. Ariz.; Mexico (Sonora).
Ammophila dejecta Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 14.
dysmica Menke. Calif., Nev., Oreg., Wyo. Ecology: Nest a vertical burrow ending in a single
cell. Prey: Noctuidae sp. Predator: Philanthus zebratus nitens (Bks.).
Ammophila dysmica Menke, 1966. Biol. Soc. Wash., Proc. 79: 30. 6, °.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 485 (nest, prey, predator).
evansi Menke. Eastern United States. Parasite: Pseudoxenos lugubris (Pierce). Replaces
arvensis of American authors, not Dahlbom.
Ammophila evansi Menke, 1964. Acta Hym. 2: 20, fig. 17. d, 9.
extremitata Cresson. Western U.S.
Ammophila extremitata Cresson, 1865. Ent. Soc. Phila., Proc. 4: 457. 2.
femurrubra Fox. Southwestern U.S. Parasite: Spintharosoma mesillae (Ckll.).
Ammophila femur-rubra Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 102. 9.
fernaldi (Murray). Eastern U.S. west to Man. and Ariz.; Mexico. Ecology: Nests in sandy soil,
the cell provisioned with a single larva. Prey: Noctuidae sp.
Sphex fernaldi Murray, 1938. Ent. Soc. Amer., Ann. 31: 19. 2, d.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 241-242, figs. 6, 7 (larva).

Biology: Evans, 1964. Amer. Ent. Soc., Trans. 99: 242 (nest, prey, life cycle).
ferruginosa Cresson. Western U.S. except Coastal States.
Ammophila ferruginosa Cresson, 1865. Ent. Soc. Phila., Proc. 4: 455. 2.
Ammophila collaris Cresson, 1865. Ent. Soc. Phila., Proc. 4: 456. ¢.
Sphex cressoni Smith, 1908. Nebr. Univ., Studies 8: 329. 3.
formicoides Menke. Ariz., N. Mex., Tex.; Mexico (Sonora, Durango).
Ammophila formicoides Menke, 1964. Acta Hym. 2: 10, fig. 3. d, 9.
harti (Fernald). Vt. to Alta. to Utah, Tex. Ecology: Nests in sand, the burrow oblique; wasp
larva is progressively provisioned. Prey: Ennominae spp., Sterrhinae spp., Geometridae
spp.; Noctuidae, probably Acontiinae sp.
Ammophila argentata Hart, 1907. Ill. State Lab. Nat. Hist., Bul. 7: 266. 2, ¢. Preoce.
Sphex harti Fernald, 1931. Ent. Soc. Amer., Ann. 24: 450. N. name.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 146, figs, 57-59 (larva).

Biology: Peckham and Peckham, 1900. Wis. Nat. Hist. Soc., Bul. 1: 90-91 (nest, prey; misdet.
as polita Cr.). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 24-26 (nest, prey). —Evans,
1959. Amer. Midland Nat. 62: 459-461 (nest, prey).

hermosa Menke. Colo., Nev., Ariz., Calif.; Mexico (Chihuahua).
Ammophila hermosa Menke, 1966. Biol. Soc. Wash., Proc. 79: 31. gd, @.
hurdi Menke. Southwestern U. S.
Ammophila hurdi Menke, 1964. Acta Hym. 2: 12, fig. 1. d, 9.
imitator Menke. Ariz. (Cochise Co.); Mexico (Sonora).
Ammophila imitator Menke, 1966. Biol. Soc. Wash., Proc. 79: 38, figs. 1, 3. 3.
juncea Cresson. Transcont. in U. 8.; Mexico (Sonora).
Ammophila juncea Cresson, 1865. Ent. Soe. Phila., Proc. 4: 460. ¢.
Ammophila montezuma Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 13. ¢.
karenae Menke. Calif., Ariz., N. Mex., Nev., Idaho; Mexico (Baja California).
Ammophila karenae Menke, 1964. Acta Hym. 2: 21, fig. 9. d, 2.
kennedyi (Murray). Entire U. S. and South. Canada. Parasite: Pseudoxenos lugubris (Pierce).
Ammophila vulgaris Cresson, 1865. Ent. Soc. Phila., Proc. 4: 458. 2, ¢. Preoce.
Sphex kennedyi Murray, 1938. Ent. Soc. Amer., Ann. 31: 36. N. name.

1592 Hymenoptera in America North of Mexico

leoparda (Fernald). Minn. southwest to Tex., east to Ga., north to Mich.
Sphex urnarius leopardus Fernald, 1934. No. Amer. and W. Indies Sphex, P. 125. 2, d.
macra Cresson. Western U. S. Ecology: Nests in firm sandy loam, the burrow vertical with a
horizontal cell. Prey: Smerinthus geminatus Say.
Ammophila macra Cresson, 1865. Ent. Soc. Phila., Proc. 4: 460. d.

Biology: Evans, 1965. Psyche 72: 21, fig. 9 (nest, prey).
marshi Menke. Calif., Nev.
Ammophila marshi Menke, 1964. Acta Hym. 2: 13, fig. 5. d, 2.
meclayi Menke. Calif., Nev.; Mexico (Baja California).
Ammophila mcclayi Menke, 1964. Acta Hym. 2: 24, fig. 13. d, 2.
mediata Cresson. Canada and western U. S. as far north as N. W. T. and Yukon. Predator:
Philanthus zebratus nitens (Bks.).
Ammophila mediata Cresson, 1865. Ent. Soe. Phila., Proc. 4: 459. 2, d.
mescalero Menke. Tex., Colo., Ariz.; Mexico (Zacatecas, Queretaro, Puebla, Oaxaca).
Ammophila mescalero Menke, 1966. Biol. Soc. Wash., Proc. 79: 33, fig. 7. d, ?.°
mimica Menke. Southern Calif., Ariz.
Ammophila mimica Menke, 1966. Biol. Soc. Wash., Proc. 79: 36, fig. 4. d, 2.
moenkopi Menke. Northern and central Ariz.
Ammophila moenkopi Menke, 1967. Los Angeles Co. Mus., Contrib. Sci. 123: 3, figs. 1, 3.
ore
monachi Menke. Nev.
Ammophila monachi Menke, 1966. Biol. Soc. Wash., Proc. 79: 34. 6, ?.
murrayi Menke. Calif.
Ammophila murrayi Menke, 1964. Acta Hym. 2: 14, fig. 4. 3, °.
nasalis Provancher. Calif. Ecology: Nests in loose sand of river bed or bank, digging a vertical
burrow, the cell provisioned with up to seven caterpillars. Parasite: Pseudoxenos
lugubris (Pierce). Prey: Geometridae sp.
Ammophila nasalis Provancher, 1895. Nat. Canad. 22: 111. d.
Sphex craspedotus Fernald, 1934. No. Amer. and W. Indies Sphex, p. 96. °.

Biology: Hicks, 1935. Pan-Pacific Ent. 11: 97-99 (nest, prey).

nearctica Kohl. Western U. S.
Ammophila nearctica Kohl, 1889. Zool.-Bot. Gesell. Wien, Verh. 39: 18. 6.
nefertiti Menke. Wash., Oreg., Calif., Ariz., Nev., Idaho, a Great Basin sp.
Ammophila nefertiti Menke, 1964. Acta Hym. 2: 16, figs. 8, 11. d, °.
nigricans Dahlbom. Eastern U. S. Ecology: Nests in sandy-clay soil, the burrow oblique or
vertical, terminating in a horizontal cell provisioned with one caterpillar. Prey: Catocala
sp., Zale sp., Euparthenos nubilis Hbn.
Ammophila nigricans Dahlbom, 1843. Hym. Europaea, v. 1, p. 14. d.
Ammophila intercepta Lepeletier, 1845. Hist. Nat. Ins., Hym. v. 3, p. 378. °.

Biology: Rau, 1934. Canad. Ent. 66: 259 (nest, prey). —Strandtmann, 1945. Ent. Soc. Amer.,
Ann. 38: 310, fig. 6 (nest, prey, life cycle). —Evans, 1959. Amer. Midland Nat. 62: 465 (prey
transport).
novita (Fernald). Southwestern U. S.; Mexico.

Sphex novitus Fernald, 1934. No. Amer. and W. Indies Sphex, p. 147. 2, d.
parapolita (Fernald). Wash., Oreg., Idaho, Nev., Utah, Calif.

Sphex parapolitus Fernald, 1934. No. Amer. and W. Indies Sphea, p. 51. 2, d.
parkeri Menke. Calif., Nev. Prey: Geometridae sp.

Ammophila parkeri Menke, 1964. Acta Hym. 2: 23, fig. 14. d, @.

Biology: Powell, 1964. Kans. Ent. Soc., Jour. 37: 243-244 (prey).

peckhami (Fernald). Colo. to Ariz.; Mexico.
Sphex willistoni Fernald, 1934. No. Amer. and W. Indies Sphex, p. 91, fig. 37. 2.
Sphex peckhami Fernald, 1934. No. Amer. and W. Indies Sphex, p. 93. d.

picipes Cameron. Ariz. south to Panama.
Ammophila alticola Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 10, pl. 1, fig. 9a. d.

Superfamily SPHECOIDEA 1593

Ammophila picipes Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 11, pl. 2, fig. 4. d.
Ammophila volcanica Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 17. 9.
Ammophila chiriquensis Cameron, 1888. Biol. Cent-Amer., Hym., v. 2, p. 18. 9.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 156, fig. 3 (sleeping aggregation).
pictipennis Walsh. Eastern U. S.; Mexico. Ecology: Nests in sand, providing a single larva per
nest. Parasite: Pseudoxenos lugubris (Pierce); Hilarella hilarella (Zett.). Prey:
Noctuidae spp., Leucania unipuncta (Haw.), Agrotis C-nigrum (L.), Prodenia
ornithogalli Guen., Heliothis zeae (Bod.); Pholisora catullus F.
Ammophila pictipennis Walsh, 1869. Amer. Ent. 1: 128, 164. 2, 6.
Ammophila anomala Taschenberg, 1869. Ztschr. Gesam. Naturw. Halle 34: 434. 2, ¢.
Sphex nigropilosus Rohwer, 1912. U. S. Natl. Mus., Proce. 41: 465. 2. This is a questionable
synonym.
Biology: Walsh and Riley, 1868. Amer. Ent. 1: 128 (nest, prey). —Rau and Rau, 1918. Wasp
studies afield, pp. 207-2387, figs. 45-49 (prey transport, nest, life cycle). —Rau, 1922. Acad.
Sci. St. Louis, Trans. 24: 23 (prey). —Strandtmann, 1945. Ent. Soc. Amer., Ann. 38:
310-311, fig. 7 (nest, prey; misdet. as placida Sm.).
placida Smith. Western U.S.; Mexico. Ecology: Nests in damp sand, the vertical burrow
ending in a horizontal cell. Parasite: Opsidia sp. Prey: Zale lwnata (Dru.), Noctuidae sp.;
Hesperiidae sp.
Ammophila placida Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 221. 6.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 148, fig. 34 (larva).

Biology: Hicks, 1932. Canad. Ent. 64: 193-198 (nest, prey, life cycle; misdet. as xanthoptera
Cam.). —Hicks, 1934. South. Calif. Acad. Sci., Bul. 33: 39-41 (prey capture). —Evans, 1959.
Amer. Midland Nat. 62: 456-458 (nest, prey transport, life cycle, parasite).

polita Cresson. Western U. S.
Ammophila polita Cresson, 1865. Ent. Soc. Phila., Proc. 4: 458. °.
procera Dahlbom. Transcont. in U. S., south to Guatemala. Ecology: Nests in compact sand, the
burrow oblique to vertical, terminating in a cell containing one caterpillar. Parasite:
Senotainia vigilans Allen, Metopia lateralis Macq., Miltogrammini sp. Prey: Nadata
gibbosa (Abbot), Heterocampa manteo (Dbldy.), H. astarte Dbldy., Datana sp., Schizura
ipomoeae (Dbldy.), Symmerista sp.; Smerinthus cerisyi Kby., Sphingidae sp.; Noctuidae
sp.
Ammophila procera Dahlbom, 1843. Hym. Europaea, v. 1, p. 15.
Ammophila saeva Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 222. 9.
Ammophila gryphus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 460. ¢.
Ammophila barbata Smith, 1873. Ann. and Mag. Nat. Hist. (4) 12: 260. 2.
Ammophila ceres Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 8. 6.
Ammophila championi Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 9. °.
Ammophila striolata Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 10. 2.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 81: 145, figs. 39-46 (larva).

Biology: Pergande, 1892. Ent. Soc. Wash., Proc. 2: 256-258 (nest, prey). —Hartman, 1905.
Tex. Univ. Bul. 65, Sci. Ser. 6: 11-20, figs. 6, 8, 9, 18, 16-18, 22 (nest, prey). —Rau and Rau,
1918. Wasp studies afield, pp. 237-243, fig. 50 (nest, prey, sleeping aggregation). —Rau,
1922. Acad. Sci. St. Louis, Trans. 24: 24 (nest). —Criddle, 1924. Canad. Field Nat. 38:
121-123 (nest, prey). —Wheeler and Wheeler, 1924. Science 59: 486 (nest closure). —Rau,
1926. Acad. Sci. St. Louis, Trans. 25: 211 (prey). —Hicks, 1935. Pan-Pacifie Ent. 11: 101-102
(nest closure). —Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 283-286 (nest, prey,
parasite). —Bohart and Knowlton, 1953. Ent. Soc. Wash., Proc. 55: 100-101 (nest, prey,
mating). —Krombein, 1953. Ent. Soc. Wash., Proc. 55: 118 (nest, prey, parasite). —Tilden,
1953. Pan-Pacifie Ent. 29: 211-218 (nest, prey). —Krombein, 1955. Ent. Soc. Wash., Proc.
57: 151-152 (cocoon). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 104-105 (nest, prey,
parasite, life cycle). —Evans, 1959. Amer. Midland Nat. 62: 451-454 (nest, prey transport,
parasite). —Linsley, 1962. Ent. Soc. Amer., Ann. 55: 156, fig. 4 (sleeping aggregation).

1594 Hymenoptera in America North of Mexico

pruinosa Cresson. Western U. S. Ecology: Nests in sand, the burrow mostly vertical, the cell
progressively provisioned with several caterpillars. Parasite: Pseudoxenos lugubris
(Pierce). Prey: Geometridae spp.; Noctuidae spp.; Phycitinae sp.
Ammophila pruinosa Cresson, 1865. Ent. Soc. Phila., Proc. 4: 455. 2, d.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 149, fig. 35 (larva).

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 156 (sleeping aggregation). —Powell, 1964.
Kans. Ent. Soe., Jour. 37: 251-253, 256 (nest, prey).
regina Menke. Calif., Oreg.
Ammophila regina Menke, 1964. Acta Hym. 2: 25, fig. 15. d, °.
shoshone Menke. Wyo., Utah.
Ammophila shoshone Menke, 1967. Los Angeles Co. Mus., Contrib. Sei. 123: 5, fig. 5. d, @.
stangei Menke. Calif., Nev.
Ammophila stangei Menke, 1964. Acta Hym. 2: 15, fig. 6. d, °.
strenua Cresson. Western Canada and U. S.; Mexico.
Ammophila strenua Cresson, 1865. Ent. Soe. Phila., Proc. 4: 459. °.
Sphex dubius Fernald, 1934. No. Amer. and W. Indies Sphex, p. 189. Preoce.
Ammophila denningi Murray, 1951. U. S. Dept. Agr., Agr. Monog. 2: 975. N. name.
unita Menke. Colo., Wyo., Utah, Nev., Ariz., a Great Basin sp.
Ammophila unita Menke, 1966. Biol. Soc. Wash., Proc. 79: 35. d, °.
urnaria Dahlbom. Eastern U. S. Ecology: Nests in fairly firm soil, the burrow vertical or
oblique, the cell provisioned with 1-6 caterpillars. Parasite: Pseudoxenos lugubris
(Pierce). Prey: Scoliopteryx libatrix L., Autographa sp., Panopoda sp., ? Polia adjuncta
Bdvl.; Ennominae spp., Geometridae spp. The earlier records of wrnaria listed under
Biology may refer to one or more species of Ammophila.
Ammophila urnaria Dahlbom, 1843. Hym. Europaea, v. 1, p. 14.
Ammophila inepta Cresson, 1872. Amer. Ent. Soc., Trans. 4: 209. 2, d.
Sphex arvensis floridensis Fernald, 1933. Ent. News 44: 236. Nom. nud.
Sphex floridensis Fernald, 1934. No. Amer. and W. Indies Sphea, p. 126. °, d.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soe., Trans. 81: 146, figs. 47-49 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 6-28, pl. 3, figs.
5-8, pl. 8, figs. 1-5 (nest, use of “tool”, prey, life cycle). —Peckham and Peckham, 1905.
Wasps, Social and Solitary, pp. 18-39, 46-50 (nest, use of “tool”, prey, life cycle). —Parker,
1915. Ent. Soc. Wash., Proc. 17: 75-76, fig. 8 (nest, prey, life cycle). —Fernald, 1933. Ent.
News 44: 236-238 (prey transport, nest). —Frisch, 1940. Amer. Midland Nat. 24: 345-350
(nest closure). —Evans, 1959. Amer. Midland Nat. 62: 462-464 (nest, prey).

varipes Cresson. Central U. S.; Mexico (Sonora).
Ammophila varipes Cresson, 1865. Ent. Soc. Phila., Proc. 4: 457. 2, d.
Ammophila comanche Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 19, pl. 1, fig. 14.
wrightii (Cresson). Western U. S. Ecology: Digs vertical burrow in soil after capturing prey.
Prey: Geometridae sp.
Coloptera wrightii Cresson, 1865. Amer. Ent. Soc., Trans. 1: 378. 2.
Biology: Hicks, 1934. Psyche 41: 150-157, 2 figs. (nesting behavior, prey).
zanthoptera Cameron. Ariz.; Mexico to Guatemala.
Ammophila zanthoptera Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 8. 2.
Ammophila trichiosoma Cameron, 1888. Biol. Cent.-Amer., Hym., v. 2, p. 11. d.
Ammophila xanthoptera Cameron, 1900. Biol. Cent.-Amer., Hym., v. 2, p. xi. Emend.

UNPLACED TAXON OF AMMOPHILINAE

Ammophila arvensis Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 384. 2, ¢. Amer. Sept.
Preocc. by Dahlbom.

Superfamily SPHECOIDEA 1595

Family PEMPHREDONIDAE

Members of this family are small for sphecoid wasps and include the smallest species (2 mm)
in the superfamily. The majority of species prey on Homoptera, especially aphids, but some of
the smallest species use Thysanoptera or Collembola. The family includes both ground-nesting
species and those which use pre-existing cavities in twigs, galls, etc., as nesting sites.

SUBFAMILY PSENINAE

The revisions listed under the subfamily heading are neither adequate nor reliable for identifi-
cation of most members of the subfamily.

Revision: Fox, 1898. Amer. Ent. Soc., Trans. 25: 1-18 (N. Amer. spp.). —Malloch, 1933. U.S.
Natl. Mus., Proc. 82 (26): 1-60, 2 pls. (N. Amer. spp.).

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 139-145, figs. 1-22 (larvae). —Gittins,
1969. Amer. Ent. Soc., Trans. 95: 49-76, 21 figs. (generic reclassification).

TRIBE PSENINI

Genus AMMOPSEN Krombein

Ammopsen Krombein, 1959. Brooklyn Ent. Soc., Bul. 54: 18.
Type-species: Ammopsen masoni Krombein. Monotypic.

Nothing is known of the biology except that the sole included species is frequently collected
while visiting the flowers of mat Euphorbia in the desert. The presence in the female of a
foretarsal comb and genal ammochaetae suggests that masoni is a ground-nesting species.

masoni Krombein. Southern Calif., Ariz., N. Mex., Utah, Nev.
Ammopsen masoni Krombein, 1959. Brooklyn Ent. Soe., Bul. 54: 19. d, °.

Genus MIMESA Shuckard

Mimesa Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 228.

Type-species: Trypoxylon equestre Fabricius. Orig. desig.
Aporia Wesmael, 1852. Acad. Roy. Sci. Belg., Bul. 19: 272. Preoce.

Type-species: Trypoxylon equestre Fabricius. Desig. by Kohl, 1896.
Aporina Gussakovskij, 1937. Trav. Inst. Zool. Acad. Sci. 4: 665. N. name. Preoce.

Wasps of this genus nest in the ground with several cells off the main burrow. So far as
known they prey on leafhoppers, both nymphs and adults.

agalena Gittins. Calif.
Mimesa agalena Gittins, 1966. Ent. News 77: 251. 2, d.
arizonensis (Malloch). Ariz. (Tucson, Roosevelt Lake).
Psen (Mimesa) arizonensis Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 36. 6.
barri Gittins. Idaho, Oreg., Calif.
Mimesa barri Gittins, 1966. Ent. News 77: 249. 9, 6.
basirufa Packard. Transcont. in Transit. and U. Austr. Zones. Ecology: Nests in flat compacted
soil or vertical sand banks. Prey: Jdiocerus sp. nymphs, Macropsis viridis (Fitch) adults,
Oncopsis variabilis (Fitch) adults, O. sorbrius (Wlkr.) adults.
Mimesa basirufa Packard, 1867. Ent. Soc. Phila., Proc. 6: 406. °.
Mimesa nebrascensis Smith, 1908. Nebr. Univ., Studies 8: 390. 2.

Biology: Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 64 (nest, prey). —Kurezewski and
Lane, 1974. Ent. Soc. Wash., Proc. 76: 377-379, figs. 1, 3 (nest, prey transport, egg).

coquilletti (Rohwer). Calif., Nev.
Psen (Mimesa) coquilletti Rohwer, 1910. U. S. Natl. Mus., Proc. 12: 103. 2 (¢ misdet.).

cressonii atriventris (Malloch). Ont. (Toronto).
Psen (Mimesa) cressoni (!) var. atriventris Malloch, 1938. U. S. Natl. Mus., Proe. 82 (26):

3l. 3.

1596 Hymenoptera in America North of Mexico

cressonii cressonii Packard. Transcont. in Transit. and U. Austr. Zones. Ecology: Nests in

level sand. Parasite: Senotainia trilineata (Wulp). Prey: Doratura stylata (Boh.) adults,
Diplocolenus configuratus (Uhler) adult and nymphs, Athysanella longicauda Beirne
adults, Polyamia compacta (Osborn and Ball) adults, Laevicephalus melsheimeri (Fitch)
adults, Scaphytopius sp. ? nymphs; Delphacodes campestris Van Duzee adult, D. sp.
nymph, Laccocera vittipennis Van Duzee adult, Liburniella ornata (Stal) adult;
Craspedolepta sp. adult.

Mimesa Cressonii Packard, 1867. Ent. Soe. Phila., Proc. 6: 405. @.

Mimesa denticulata Packard, 1867. Ent. Soc. Phila., Proc. 6: 406. ¢.

Mimesa conica Smith, 1908. Nebr. Univ., Studies 8: 389. “2” = 6.

Biology: Kurezewski and Lane, 1974. Ent. Soc. Wash., Proc. 76: 379-382, figs. 2, 4-6 (nest,
prey transport, egg, parasite).
dawsoni Mickel. Nebr. (Harrison).
Mimesa dawsoni Mickel, 1916. Amer. Ent. Soc., Trans. 42: 420. 6.
edentata (Malloch). Calif. (San Diego Co.).
Psen (Mimesa) edentatus Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 37. 6.
ezra (Pate). Maine, Pa., Md., Wis., Kans., Colo., Wash. Ecology: Nests in sandy soil. Prey:
Exitianus exitiosus (Uhler).
Mimesa argentifrons Cresson, 1865. Ent. Soc. Phila., Proc. 4: 487. 2, d. Preoce.
Psen (Mimesa) ezra Pate, 1944. Canad. Ent. 76: 133. N. name.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 226, pl. 33, figs. 1, 3 (nest, prey).

granulosa (Fox). S. Dak., Mont., Idaho, N. Mex.

Psen granulosus Fox, 1898. Amer. Ent. Soc., Trans. 25: 15. 6.
gregaria gregaria (Fox). Wyo., Colo., N. Mex.

Psen gregarius Fox, 1898. Amer. Ent. Soc., Trans. 25: 16. 6.
gregaria simplex (Malloch). Idaho, Utah, Colo.

Psen (Mimesa) gregarius var. simplex Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 42. °,

3.

impressifrons (Malloch). Wash. (Perry).

Psen (Mimesa) impressifrons Malloch, 1933. U.S. Natl. Mus., Proc. 82 (26): 38. ¢.
maculipes Fox. Ont. to Fla.

Mimesa maculipes Fox, 1893. Canad. Ent. 25: 117. ¢.

Psen (Mimesa) nigrescens Rohwer, 1910. Ent. News 21: 168. ¢.

Psen (Mimesa) perplexa Rohwer, 1910. Ent. News 21: 169. 9.
pauper Packard. Transit. Zone east of Rockies.

Mimesa pauper Packard, 1867. Ent. Soc. Phila., Proc. 6: 409. ¢.

Mimesa cingulata Packard, 1867. Ent. Soc. Phila., Proc. 6: 410. d.
polita (Malloch). Maine, Md., Nebr., N. Mex.

Psen (Mimesa) politus Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 35. 2, ¢.
proxima Cresson. Nebr., Colo., N. Mex., Wash.

Mimesa proxima Cresson, 1865. Ent. Soc. Phila., Proc. 4: 188. @.
punctifrons (Malloch). Calif. (Redlands).

Psen (Mimesa) punctifrons Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 36. 3.
pygidialis (Malloch). Mich., Colo., Alta.

Psen (Mimesa) pygidialis Malloch, 1933. U.S. Natl. Mus., Proc. 82 (26): 39. 2, 6.
sabina Gittins. Calif.

Mimesa sabina Gittins, 1966. Ent. News 77: 247. 9, 6.
unicincta Cresson. Colo., Calif., B. C. Predator: Philanthus pulcher D. T.

Mimesa unicincta Cresson, 1865. Ent. Soe. Phila., Proc. 4: 488. 9.

Genus MIMUMESA Malloch

Psen subg. Mimumesa Malloch, 1933. U. S. Natl. Mus., Proe. 82 (26): 16.
Type-species: Psen niger Packard. Orig. desig.

Superfamily SPHECOIDEA 1597

The North American nigra (Pack.) nests in decaying timber and preys upon adult Agallia leaf-
hoppers. Several species in the Old World have similar nesting habits and one species nests in
clay banks. The Old World species prey upon both Cicadellidae and Delphacidae.
canadensis (Malloch). Ont., N. Y., Mich., N. Dak., Colo., Alaska.

Psen (Mimumesa) canadensis Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 22. 2, d.
clypeata (Fox). Colo., Utah, Nev., Idaho, Calif., Wash., Alaska, N. W. T.

Psen clypeatus Fox, 1898. Amer. Ent. Soc., Trans. 25: 15. 2, d.
coloradoensis (Cameron). Colo. (Berkeley Co.).

Psen (Mimesa) coloradoensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 232. 2.
cylindrica (Fox). Ariz.

Psen cylindricus Fox, 1898. Amer. Ent. Soc., Trans. 25: 5. 2, d.
fuscipes (Packard). Mass.

Psen fuscipes Packard, 1867. Ent. Soc. Phila., Proc. 6: 402. °.
interstitialis (Cameron). N. Mex.

Psen (Mimesa) interstitialis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 233. 3.
johnsoni Viereck. N. Y. to Va., W. Va.

Mimesa johnsoni Viereck, 1901. Amer. Ent. Soc., Trans. 27: 340. 9, d.
leucopus (Say). N. H., Md., D. C., Va., Ind., Ill., Alta.

Psen leucopus Say, 1837. Boston Jour. Nat. Hist. 1: 370. 2, d.

Psen elongatus Packard, 1867. Ent. Soc., Phila., Proc. 6: 400. “2” = ¢.
longicornis (Fox). R. I., N. Y. to Fla., La., lowa; Cuba, Cent. Amer.

Psen longicornis Fox, 1898. Amer. Ent. Soc., Trans. 25: 8. d.

Mimesa striatus Viereck, 1901. Amer. Ent. Soc., Trans. 27: 339. 2.

Psen (Mimesa) floridana Rohwer, 1910. Ent. Soc. Wash., Proce. 12: 102. 9.

mandibularis Smith. Nebr.
Mimesa mandibularis Smith, 1908. Nebr. Univ., Studies 8: 392. °.
mellipes (Say). N. Y., Md., D. C., Ohio, Ind., Ill, Iowa, Nebr.
Psen mellipes Say, 1837. Boston Jour. Nat. Hist. 1: 369. °.
Psen chalcifrons Packard, 1867. Ent. Soc. Phila., Proc. 6: 401. °.
mixta (Fox). Transcont. in Transit. Zone. Predator: Philanthus pulcher D. T.
Psen mixtus Fox, 1898. Amer. Ent. Soc., Trans. 25: 7. 2, d.
Mimesa alticola Viereck, 1903. Amer. Ent. Soc., Trans. 29: 65. 2.
Psen (Mimesa) similis Rohwer, 1910. Ent. Soc. Wash., Proc. 12 OESe
nigra (Packard). Transcont. in Transit. Zone. Ecology: Nests in decaying wood. Prey: Agallia
sp. adults.
Psen niger Packard, 1867. Ent. Soc. Phila., Proc. 6: 399. 2 (6 misdet.).
Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 143-144, figs. 21-22 (larva).

Biology: Gurney, 1951. Ent. Soc. Wash., Proc. 53: 280 (nest, prey).
propinqua Kincaid. Alaska.
Mimesa rropinqua Kincaid, 1900. Wash. Acad. Sci., Proc. 2: 508. d.

psychrus (Pate). Canada (Hudson Bay).
Mimesa borealis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 431. 6. Preoce.
Psen (Mimesa) psychrus Pate, 1944. Canad. Ent. 76: 183. N. name.

regularis (Fox). N. J., Pa.
Psen regularis Fox, 1898. Amer. Ent. Soc., Trans. 25: 6. °.

Taxonomy: Viereck, 1901. Amer. Ent. Soc., Trans. 27: 339. 2.
Genus PSENEO Malloch

Psen subg. Pseneo Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 7.
Type-species: Psen kohlii Fox. Orig. desig.

Meager records indicate that these wasps nest either in the soil or in decaying wood, and prey
upon leafhoppers, both nymphs and adults.

1598 Hymenoptera in America North of Mexico

Revision: Krombein, 1950. Ent. Soc. Wash., Proc. 52: 277-287 (N. Amer. spp.). —van Lith,
1975. Tijdschr. v. Ent. 118: 15-39, figs. 16-37 (New World spp.).
longiventris kohlii (Fox). N. Y. south to Ga., W. Va., Ala., Ind., Kans.
Psen Kohlii Fox, 1898. Amer. Ent. Soc., Trans. 25: 9. 2 (¢ misdet.).
Psen (Pseneo) fulvipes Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 11, fig. 3. °.
Psen (Pseneo) angulatus Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 12. ¢.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 22, figs. E-J (male
genitalia).
longiventris longiventris (Cameron). N. Mex., Ariz. south to Colombia and Surinam.
Mimesa longiventris Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 187, pl. 8, fig. 18. “2?”
= 6.
Mimesa montezuma Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 188. 9.
Psen (Pseneo) spicatus Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 12. 6.
punctatus carolina (Rohwer). Va. to north. Fla., Ala., Miss., Ark. Ecology: Nests in soil in
flowerpot. Prey: Homalodisca triquetra (F.), Graphocephala coccinea (Foerster).
Psen (Mimesa) punctata var. carolina Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 103. 2.

Biology: Krombein, 1950. Ent. Soc. Wash., Proc. 52: 283 (nest, prey).
punctatus ferrugineus (Viereck). South. Fla.
Mimesa ferruginea Viereck, 1901. Amer. Ent. Soc., Trans. 27: 341. ¢.
punctatus punctatus (Fox). S. Dak., Nebr., Colo.; Mexico (Morelos).
Psen punctatus Fox, 1898. Amer. Ent. Soc., Trans. 25: 9. 9°.
simplicicornis (Fox). Que., N. J. and Pa. south to N. C. Ecology: Nests in decaying wood;
cocoons of reared series bear fragments of wood pulp. Prey: Graphocephala sp.,
Cicadellinae sp.
Psen simplicicornis Fox, 1898. Amer. Ent. Soc., Trans. 25: 10. 2 (¢ misdet.).

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 142 (larva).

Biology: Krombein, 1950. Ent. Soc. Wash., Proc. 52: 286 (nest). —Krombein, 1951. U. S. Dept.
Agr., Agr. Monog. 2: 959 (prey).

Genus PSEN Latreille

Psen Latreille, 1796. Precis Caract. Gen. Ins., p. 122. No species.
Type-species: Sphex atra Fabricius. First included species.
Psenia Stephens, 1829. System. Cat. Brit. Ins., p. 361.
Type-species: Sphex atra Fabricius. Desig. by Pate, 1937.
Dahlbomia Wissman, 1849. Stettin. Ent. Ztg. 10: 9.
Type-species: Sphex atra Fabricius. Monotypic.
Mesopora Wesmael, 1852. Acad. Roy. Sci. Belg., Bul. 19: 279.
Type-species: Sphex atra Fabricius. Monotypic.
Caenopsen Cameron, 1899. Ann. and Mag. Nat. Hist. (7) 4: 55.
Type-species: Caenopsen fuscinervis Cameron. Monotypic.
Punctipsen van Lith, 1968. Tijdschr. Ent. 111: 125.
Type-species: Mimesa exarata Eversmann. Orig. desig.

The North American barthi Vier. nests in dead wood and preys upon Membracidae, and
erythropoda Roh. preys upon Cercopidae. Extralimital species have been reported as nesting in
dead wood or in sandy or clay soil, and as preying upon Cicadellidae, Cercopidae and Mem-
bracidae.

Revision: van Lith, 1975. Tijdschr. v. Ent. 118: 2-15, figs. 1-15 (New World spp.).
barthi Viereck. Que., Conn., Pa., Md., Ga., Wis. Ecology: Nests in wood. Prey: Cyrtolobus
fenestratus (Fitch); Atymna inornata (Say); Micrutalis calva (Say); cocoons from which
type series of myersiana was reared bear attached fragments of Membracidae, perhaps
Enchenopa binotata (Say).
Psen (Mimesa) barthi Viereck, 1907. Wis. Nat. Hist. Soe., Bul. 5: 251. °.
Mimesa myersiana Rohwer, 1909. Ent. News 20: 324. ?, 6.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 142-143, fig. 20 (larva).

Superfamily SPHECOIDEA 1599

Biology: Barth, 1907. Wis. Nat. Hist. Soc., Bul. 5: 251-257 (nest, prey).

erythropoda Rohwer. Ont., Maine to Ga., Wis., Ind. Prey: Aphrophora quadrinotata Say.
Psen (Mimesa) erythopoda (!) Rohwer, 1910. Ent. Soe. Wash., Proce. 12: 102. 9.
Psen (Mimesa) erythropoda Rohwer, 1910. Ent. Soc. Wash., Proce. 12: 176. Emend.

Biology: Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 15 (prey).
monticola (Packard). Ont., N. H. to Ga., W. Va., Mich., Ala.
Mimesa monticola Packard, 1867. Ent. Soe. Phila., Proc. 6: 407. ¢.
unifasciculatus Malloch. N. Mex. (Beulah). Possibly the opposite sex of and a synonym of

montivagus D. T.
Psen (Psen) unifasciculatus Malloch, 1933. U. S. Natl. Mus. Proce. 82 (26): 15. 3.

TRIBE PSENULINI

Genus PLUTO Pate

Psenia Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 44. Preoce.
Type-species: Mimesa tibialis Cresson. Orig. desig.
Pluto Pate, 1937. Amer. Ent. Soc., Mem. 9: 51. N. name.

Biological notes are available for only one species which nests in large aggregations in sandy
soil and preys upon nymphal and adult leafhoppers. The presence of a foretarsal pecten in the
female suggests that members of the genus are all ground-nesting.
aerofacies (Malloch). Tex.; Mexico.

Psenia aerofacies Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 49. 2, 6.
albifacies (Malloch). Iowa, Tex. Ecology: Nests gregariously in flat clay-sand, each nest with

several cells and 5-12 prey per cell. Prey: Opsius stactogalus Fieb., nymphs and adults.
Psenia albifacies Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 50. 9.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 140-141, figs. 1-6 (larva).

Biology: Evans, 1968. Ent. Soc. Amer., Ann. 61: 1344 (nest, prey).

angulicornis (Malloch). Tex.

Psenia angulicornis Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 58. 2, 6.
arenivagus Krombein. N. C., Ga., Fla.

Psenia angulicornis var. Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 48 (in key).

Pluto arenivagus Krombein, 1950 (1949). Elisha Mitchell Sci. Soc., Jour. 65: 268. 2, 6.
brevipetiolatus (Rohwer). Calif.

Psenulus (Neofoxia) brevipetiolatus Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 100. 2.
clavicornis (Malloch). Ariz.; Mexico.

Psenia clavicornis Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 50. 6.
littoralis (Malloch). Md., Fla.

Psenia littoralis Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 56. , d.
longiventris (Malloch). Ariz., Calif.

Psenia longiventris Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 52. 2, 6.
marginatus (Malloch). S. C., La.

Psenia marginata Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 54. 2, 6.
minutus (Malloch). Tex. (San Diego).

Psenia minuta Malloch, 1938. U. S. Natl. Mus., Proc. 82 (26): 59. °.
pallidistigma (Malloch). Ariz., Tex.

Psenia pallidistigma Malloch, 1933. U. S. Natl. Mus., Proce. 82 (26): 52. 2, ¢.
rufibasis (Malloch). Ga.

Psenia rufibasis Malloch, 1933. U.S. Natl. Mus., Proc. 82 (26): 58. 2, d.
sayi (Rohwer). Austr. Zone.

Psenulus (Neofoxia) sayi Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 100. °.

suffusus (Fox). N. Mex., Nev., Calif.
Psen suffusus Fox, 1898. Amer. Ent. Soc., Trans. 25: 18. 2 (misdet. in part).

1600 Hymenoptera in America North of Mexico

texanus (Malloch). Tex. (Brownsville).

Psenia texana Malloch. 1933. U. S. Natl. Mus., Proc. 82 (26): 56. 2, d.
tibialis (Cresson). D. C., Va., S. C., Tenn., Ala., La., Mo., Tex.

Mimesa tibialis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 227. 2, 6.

Genus PSENULUS Kohl

Psenulus Kohl, 1896. K. K. Naturhist. Hofmus., Ann. 11: 254, 293.

Type-species: Psen fuscipennis Dahlbom. Desig. by Ashmead, 1899.
Neofoxia Viereck, 1901. Amer. Ent. Soc., Trans. 27: 338.

Type-species: Psen atrata (Fabricius) of Panzer. Orig. desig.
Stenomellinus Schulz, 1911. Zool. Ann. 4: 142.

Type-species: Psen dilectus Saussure. Monotypic.
Psenulus subg. Eopsenulus Gussakovskij, 1934. Mushi 7: 84.

Type-species: Psenulus (Eopsenulus) iwatai Gussakovskij. Orig. desig.
Nipponopsen Yasumatsu, 1938. Mushi 11: 84.

Type-species: Nipponopsen anomoneurae Yasumatsu. Orig. desig.

These wasps nest in pre-existing cavities in twigs, stems and grass, and also in abandoned
beetle borings in wood. Prey records of North American species are Aphididae and Psyllidae.
Some extralimital species prey upon Delphacidae and Cicadellidae.

Revision: Krombein, 1950. Brooklyn Ent. Soe., Bul. 45: 35-40 (N. Amer. spp.).
alienus (Krombein). Calif. Ecology: Nests in borings in Sambucus stems. Prey: Psyllidae sp.

adults.
Diodontus (Diodontus) alienus Krombein, 1950. Brooklyn Ent. Soc., Bul. 45: 38. 9.

Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 94 (nest, prey).
frontalis (Fox). Colo., N. Mex., Ariz., Utah, Calif., Wash.
Psen frontalis Fox, 1898. Amer. Ent. Soc., Trans. 25: 4. 9.
Diodontus occidentalis Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 5. 2. Preoce.
Diodontus hesperus Pate, 1944. Canad. Ent. 76: 133. N. name.
pallipes parenosas (Pate). N. H. to Ga., Utah, Calif. Ecology: Nests in borings in wood, in old
anobiid borings, and in canes of Rubus. Parasite: Pyemotes ventricosus (Newp.);
Lackerbaueria krombeini Baker; Chrysididae sp., possibly Omalus. Prey: Drepanaphis
acerifoliae (Thos.) adults, D. sp. nymphs, Therioaphis sp. ? nymphs, Macrosiphum sp.
nymphs. Typical pallipes (Panz.) and other subspecies occur in the Palaearctic Region.
Diodontus parenosas Pate, 1944. Canad. Ent. 76: 133. N. name for Malloch’s misdet. of
trisulcus (Fox).

Taxonomy: Krombein, 1950. Brooklyn Ent. Soc., Bul. 45: 37 (lectotype desig.). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 144-145, figs. 13-19 (larva).

Biology: Krombein, 1951. U. S. Dept. Agr., Agr. Monog. 2: 958 (nest). —Krombein, 1955.

Brooklyn Ent. Soc., Bul. 50: 15-16 (nest, prey). —Krombein, 1958. Biol. Soc. Wash., Proc.
71: 22 (nest, prey). —Krombein, 1967. Trap-nesting wasps and bees, pp. 282-234 (nest,
prey, life cycle, cocoon, parasites).

trisuleus (Fox). N. H. to Tenn., Mo., Kans. Ecology: Nests in elder stems. Parasite: Omalus

iridescens (Nort.).

Psen trisulcus Fox, 1898. Amer. Ent. Soc., Trans. 25: 5. 2.
Diodontus corusanigrens Rohwer, 1920. U.S. Natl. Mus., Proc. 57: 228. “2” = 6.
Diodontus sulcatus Malloch, 1933. U. S. Natl. Mus., Proc. 82 (26): 6. @.

Taxonomy: Pate, 1944. Canad. Ent. 76: 133.
Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 21-22 (nest).
SUBFAMILY PEMPHREDONINAE

Revision: Fox, 1892. Amer. Ent. Soc., Trans. 19: 307-326 (N. Amer. spp.; not reliable for
identification).

Superfamily SPHECOIDEA 1601

Taxonomy: Pate, 1937. Amer. Ent. Soc., Trans. 63: 90-93 (key to gen.). —Evans, 1958. Amer.
Ent. Soe., Trans. 84: 126-136, figs. 49-57, 60-69 (larvae). —Evans, 1964. Amer. Ent. Soc.,
Trans. 90: 246-253, figs. 17-25 (larvae).

TRIBE PEMPHREDONINI
Genus DIODONTUS Curtis

Diodontus Curtis, 1834. Brit. Ent., v. 11, text for pl. 496.
Type-species: Pemphredon tristis Vander Linden. Desig. by Internat]. Comn. Zool.
Nomencl., 1968. Op. 844, Bul. Zool. Nomencel. 25: 10.
Xylocelia Rohwer, 1915. U.S. Natl. Mus., Proce. 49: 243.
Type-species: Diodontus occidentalis Fox. Orig. desig.

Members of this genus nest in the ground. Aphids are the normal prey, but there is one record
of a leafhopper being used.

Taxonomy: Mickel, 1916. Ent. Soc. Amer., Ann. 9: 344-352 (N. Amer. spp.). —Krombein, 1939.
Brooklyn Ent. Soc., Bul. 34: 143 (key to N. Y. spp.). —Bohart and Menke, 1965. Bul. Zool.
Nomencl. 22: 257-258 (request to place Diodontus on Official List of Generic Names with
tristis as type-species).

adamsi Titus. Mich. (Isle Royale).
Diodontus adamsi Titus, 1909 (1908). Mich. Board Geol. Survey Rpt. for 1908, p. 319. °.
americanus Packard. Maine, Wis. Ecology: Nests in soil. Prey: Choke-cherry aphids.
Diodontus americanus Packard, 1867. Ent. Soe. Phila., Proc. 6: 393. 2.

Biology: Peckham and Peckham, 1898. Wis. ‘Geol. Nat. Hist. Survey, Bul. 2: 99-107, pl. 10, fig.
5, pl. 14, fig. 3 (nest, prey, life cycle).
antennatus (Mickel). Nebr. (Omaha).
Xylocelia antennatus Mickel, 1916. Ent. Soc. Amer., Ann. 9: 348. °.
argentinae Rohwer. Colo., Wyo. Ecology: Nests gregariously in flat sand and makes up to 5
cells per nest. Parasite: Diptera sp. Prey: Aphididae sp. or spp. Predator: Philanthus

pulcher D. T.
Diodontus argentinae Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 104. 6.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 486 (nest, prey, parasite, predator).
ater (Mickel). Nebr., Kans., Pa. Ecology: Nests in vertical sand or clay bank, and in sloping
gravel bank. Prey: Aphididae sp. or spp. Predator: Philanthus pacificus Cr.
Xylocelia ater Mickel, 1916. Ent. Soc. Amer., Ann. 9: 351. 9.
Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 21 (nest). —Evans, 1970. Mus. Compar.
Zool., Bul. 140: 486-487 (nest, prey, predator).
beulahensis (Rohwer). N. Mex. (Beulah).
Xylocelia beulahensis Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 174. 9.
bidentatus Rohwer. N. B., N. Y., Mich.
Diodontus bidentatus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 560. 6.
brunneicornis Viereck. Kans. (Sedgwick Co.).
Diodontus brunneicornis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 212. d.
cockerelli Rohwer. Colo. (Florissant).
Diodontus cockerelli Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 105. 6.
crassicornus Viereck. Oreg. (Corvallis).
Diodontus crassicornus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 243. ¢.
flavitarsis Fox. Colo.
Diodontus flavitarsis Fox, 1892. Amer. Ent. Soc., Trans. 19: 316. ¢.
florissantensis Rohwer. Colo. (Florissant).
Diodontus florissantensis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 107. 2, d.
franclemonti (Krombein). N. Y., Pa., Md., D. C., Va., Mich. Ecology: Nests gregariously in

sand.
Xylocelia franclemonti Krombein, 1939. Brooklyn Ent. Soc., Bul. 34: 141. 2, 6.

1602 Hymenoptera in America North of Mexico

Taxonomy: Evans, 1958. Amer. Ent. Soe., Trans. 84: 133, figs. 65, 66 (larva).

Biology: Lin, 1967. Ent. Soc. Wash., Proc. 69: 343-346 (linear copulation).
fraternus Rohwer. Colo. (Florissant).
Diodontus fraternus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 106. ¢.
gillettei Fox. Nebr., Colo., Idaho. Predator: Philanthus pulcher D. T., P. pacificus Cr.
Diodontus Gillette: Fox, 1892. Amer. Ent. Soc., Trans. 19: 316. 2.
leguminiferus Cockerell. N. Mex. (Santa Fe).
Diodontus leguminiferus Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila.,
Proc. 49: 141. 6.
maestus Mickel. Nebr., Colo.
Xylocelia maestus Mickel, 1916. Ent. Soc. Amer., Ann. 9: 347. 6.
metathoracicus (Mickel). Ala., Mo., Nebr. Ecology: Nests in clay bank, the cell with up to 48
aphids. Prey: Hysteroneura setariae (Thos.).
Xylocelia metathoracicus Mickel, 1916. Ent. Soc. Amer., Ann. 9: 349. °.
Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 102-108, figs. 27, 28 (nest, prey,
mating).
neomexicanus Rohwer. N. Mex. (Rowe).
Diodontus neomexicanus Rohwer, 1909. Amer. Ent. Soe., Trans. 19: 317. 9.
nigritus Fox. Colo.
Diodontus nigritus Fox, 1892. Amer. Ent. Soc., Trans. 19: 317. °.
occidentalis Fox. Nebr., Ariz., Calif. Ecology: Nests gregariously in level sand, the cells
provisioned with 23-30 aphids. Parasite: Omalus cressoni (Aar.). Prey: Aphis sp., Myzus
persicae Sulz. (?), Macrosiphum (Acyrthosiphon) sp., Rhopalosiphum sp.
Diodontus occidentalis Fox, 1892. Amer. Ent. Soc., Trans. 19: 315. 9, 6.
Biology: Powell, 1964. Wasmann Jour. Biol. 21: 155-176 (nest, prey, parasites, mating, life
cycle, cocoon).
rugosus Fox. Ill., Nebr., Colo., Mont.
Diodontus rugosus Fox, 1892. Amer. Ent. Soc., Trans. 19: 315. ¢.
siouxensis (Mickel). Nebr. (Sioux Co.).
Xylocelia siouxensis Mickel, 1916. Ent. Soc. Amer., Ann. 9: 350. °.
spiniferus (Mickel). Nebr. (Omaha).
Xylocelia spiniferus Mickel, 1916. Ent. Soc. Amer., Ann. 9: 348. 9.
striatus (Mickel). N. Dak., Wyo.
Xylocelia striatus Mickel, 1916. Ent. Soc. Amer., Ann. 9: 350. 9.
vallicolae Rohwer. Colo. (Florissant, Boulder).
Diodontus vallicolae Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 104. 3.
Diodontus vallicolae salicis Rohwer, 1909. Amer. Ent. Soe., Trans. 35: 105. 6.
virginianus (Rohwer). N. Y., Md., Va. Ecology: Nests gregariously in flat soil, or in pockets of
soil between roots, the cell provisioned with up to 5 aphids. Parasite: Omalus
intermedius (Aar.). Prey: Prociphilus tesselatus (Fitch) nymphs; Typhlocyba sp. adult.
Xylocelia virginiana Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 175. d.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 133, fig. 67 (larva).

Biology: Krombein, 1951. U. S. Dept. Agr., Agr. Monog. 2: 964 (prey). —Krombein, 1958. Biol.
Soc. Wash., Proc. 71: 22-24 (nest, prey). —Krombein, 1963. Ent. Soc. Wash., Proc. 65: 264
(parasite).

Genus PEMPHREDON Latreiile

The species nest in twigs, deserted galls, abandoned beetle burrows, or in rotten wood and
provision the cells with Aphididae.

Genus PEMPHREDON Subgenus PEMPHREDON Latreille

Pemphredon Latreille, 1796. Precis. Caract. Gen. Ins., p. 128. No species.
Type-species: Crabro lugubris Fabricius. Desig. by Shuckard, 1837.

Superfamily SPHECOIDEA 1603

Revision: Rohwer, 1917. Brooklyn Ent. Soc., Bul. 11: 97-102 (N. Amer. spp.).
concolor Say. Transcont. in Transit. Zone. Ecology: Nests in beetle borings in stump. Parasite:

Omalus janus (Hald.), O. sp.; Perithous mediator pleuralis (Cr.); Phalacrotophora
longifrons (Brues). Prey: Longistigma caryae (Harr.).

Pemphredon concolor Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 339.

Pemphredon Morio Cresson, 1865. Ent. Soc. Phila., Proc. 4: 486. ?. Preoce.

Pemphredon concolor Provancher, 1882. Nat. Canad. 13: 78. ?. Preoce.

Pemphredon Cressonii Dalla Torre, 1897. Cat. Hym., v. 8, p. 359. N. name.

Pemphredon Provancheri Dalla Torre, 1897. Cat. Hym., v. 8, p. 359. N. name.

Pemphredon shawii Rohwer, 1917. Brooklyn Ent. Soc., Bul. 12: 100. 6.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 128-129, figs. 49-55 (larva).

Biology: Reinhard, 1929. Nature Mag. 13: 154-157, 7 figs. (nest, prey, cocoon, life cycle,
parasites). —Reinhard, 1929. The Witchery of Wasps, pp. 192-214, 3 pls. (nest, prey,
cocoon, life cycle, parasites).

confertim Fox. Wash., Oreg., Calif. Ecology: Nests in oak gall and in borings in stems.
Parasite: Anthrax irroratus Say; Senotainia trilineata (Wulp); Omalus janus (Hald.), O.
purpuratus (Prov.); Perithous mediator neomexicanus (Vier.).
Pemphredon confertim Fox, 1892. Amer. Ent. Soc., Trans. 19: 311. d.
Pemphredon errans Rohwer, 1917. Brooklyn Ent. Soc., Bul. 12: 99. 2, ¢.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest, parasites).

foxii Rohwer. Pa., N. J.
Pemphredon foxii Rohwer, 1917. Brooklyn Ent. Soc., Bul. 12: 101. 9.
montanus Dahlbom. Transit. Zone east of Rocky Mts., B. C., N. W. T.; Europe.
Pemphredon lugubris var. b. Zetterstedt, 1838. Ins. Lapponica, p. 441.
Pemphredon montanus Dahlbom, 1844. Hym. Europaea, v. 1, p. 262. 2, d.
Pemphredon angularis Fox, 1892. Amer. Ent. Soc., Trans. 19: 310. 2, 6.
nearcticus Kohl. Nev., Colo.
Pemphredon (Cemonus) nearcticus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 55. 2, ¢.
Pemphredon cockerelli Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 103. “2” = 6.
rileyi Fox. Calif. (Placer Co.).
Pemphredon Rileyi Fox, 1892. Amer. Ent. Soc., Trans. 19: 310. 2 (¢ in part, d misdet.).
tinctipennis Cameron. Ariz.
Pemphredon tinctipennis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 234. 2.
virginianus Rohwer. N. Y., Pa., Va., W. Va., Wis. Ecology: Nests in rotten stump of tuliptree.
Prey: Cinara sp.
Pemphredon virginiana Rohwer, 1917. Brooklyn Ent. Soc., Bul. 12: 101. 2, 6.

Genus PEMPHREDON Subgenus CEMONUS Panzer

Cemonus Jurine, 1801. Intell. Blatt. Litt. Ztg. Erlangen, v. 1, p. 164. Name suppressed by
Internatl. Comn. Zool. Nomencl., Op. 135, 1939.
Cemonus Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 186.
Type-species: Sphex unicolor Panzer. Monotypic.
Cemonus Jurine, 1807. Nouv. Meth. Class., Hym. Dipt., p. 213. Preoce.
Type-species: Sphex unicolor Panzer. Desig. by Shuckard, 1837.
Dineurus Westwood, 1837. Mag. Nat. Hist. (n. s.) 1: 173.
Type-species: Pemphredon unicolor of Latreille. Orig. desig.
Cenomus Gimmerthal, 1836. Soc. Imp. Nat. Moscou 9: 436. Emend. or lapsus.
Diphlebus Westwood, 1840. Introd. Mod. Classif. Ins. 2, Gen. Synop., p. 81.
Type-species: Pelopoeus unicolor Fabricius of Panzer. Orig. desig.
Chevrieria Kohl, 1883. Schweiz. Ent. Gesell., Mitt. 6: 658. Preoce.
Type-species: Pelopoeus unicolor Fabricius of Panzer. Orig. desig.
bipartior Fox. N. Y. to Va., W. Va., Mo., Tex., N. W. T. Ecology: Nests in borings in twigs of
sumac and elder. Parasite: Omalus purpuratus (Prov.) ?; Perithous mediator pleuralis
(Cr.) ? Prey: Eriosoma lanigerum (Hausm.), Rhopalosiphum rhois Mon., Aphididae sp.
Pemphredon bipartior Fox, 1892. Amer. Ent. Soc., Trans. 19: 313. 2.

1604 Hymenoptera in America North of Mexico

Pemphredon bipartitor (!) Dalla Torre, 1897. Cat. Hym., v. 8, p. 356.
Pemphredon biparitor (!) Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 369.
Pemphredon (Cereonus (!)) harbecki Rohwer, 1910. Ent. News 21: 170. °.

Biology: Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 369-378, figs. 46, 47 (nest, prey, life cycle).
—Rau, 1934. Canad. Ent. 66: 259 (nest, prey, life history). —Krombein, 1960. Ent. News
71: 34 (nest, prey).
grinnelli (Rohwer). B. C., Calif., Ariz., Utah, Colo. Ecology: Nests in borings in Sambucus

stems. Parasite: Anthrax irroratus Say; Omalus trilobatus Boh. and Camp., O. cressoni
(Aar.); Eurytoma stigma Ashm.; Habrocytus analis (Ashm.).

Ceratophorus gennelli (!) Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 104. ?. This name was
intended as a patronymic for F. Grinnelli, Jr., who collected the holotype.

Ceratophorus grinnelli utahensis Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 562. °.

Cemonus giffardi Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 246. ?.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 129, fig. 57 (larva).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest, parasites).
inornatus Say. Transcont. in Transit. and U. Austr. Zones; Europe. Ecology: Nests in cavities

in twigs, in rotten wood, and in exposed tree roots. Parasite: Perithows mediator
pleuralis Cr. Prey: Macrosiphum sp.

Pemphredon inornatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 339.

Cemonus Shuckardi Morawitz, 1864. Acad. Imp. des Sci. St. Petersburg, Bul. 7: 460. 2, ¢.

Cemonus dentatus Puton, 1871. Soe. Ent. de France, Ann. (5) 1: 94. °.

Pemphredon tenax Fox, 1892. Amer. Ent. Soc., Trans. 19: 313. 2, ¢.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 128-129, figs. 49-55 (larva).

Biology: Rau and Rau, 1918. Wasp Studies Afield, p. 108 (nest, prey, life cycle). —Rau, 1928.
Acad. Sci. St. Louis, Trans. 25: 373-375 (nest, prey). —Rau, 1946. Brooklyn Ent. Soc., Bul.
41: 10. —Evans, 1958. Amer. Ent. Soc., Trans. 84: 129 (nest).

lethifer (Shuckard). Ont. south to Fla., Pa. west to IIl., Colo., Utah, Wash., Calif.; Europe.

Ecology: Nests in North America in soft pith of roses, berries, sumac, hibiscus. Possibly
adventive in North America. Parasite: Perithows divinator (Rossi); Omalus auratus (L.),
O. purpuratus (Prov.). Prey: Aphis gossypii Glov., A. sp. nymphs, Chaitophorus
populicola patchae H. R. L. nymphs and adults.

Cemonus lethifer Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 201. 9, 6.

Cemonus strigatus Chevrier, 1870. Schweiz. Ent. Gesell., Mitt. 3: 269. 2, ¢.

Cemonus Fabricii Mueller, 1911. Ent. Rundschau 28: 107. N. name for wnicolor Auctt.

Pemphredon (Diphlebus) littoralis Wagner, 1918. Deut. Ent. Ztschr., p. 148. 9, d.

Pemphredon (Diphlebus) fuscatus Wagner, 1918. Deut. Ent. Ztschr., p. 143. 3.

Pemphredon (Diphlebus) neglectus Wagner, 1918. Deut. Ent. Ztschr., p. 148. 6.

Pemphredon (Diphlebus) minutus Wagner, 1918. Deut. Ent. Ztschr., p. 143. 6.

Pemphredon (Dineurus) lethifer form confusa Wagner, 1932 (1931). Deut. Ent. Ztschr., p.
2319

Pemphredon (Dineurus) brevipetiolatus Wagner, 1932 (1931). Deut. Ent. Ztschr., p. 232,
fig. 14. 3.

Taxonomy: Krombein. 1959. Brooklyn Ent. Soc., Bul. 54: 95-96 (occurrence of lethifer in N.
Amer. and differentiation from N. Amer. spp.). —Evans, 1964. Amer. Ent. Soc., Trans. 90:
246-248, figs. 23-25 (larva).

Biology: Rau, 1948. Ent. Soc. Amer., Ann. 41: 326 (nest, prey; misdet. as inornatus).
—Krombein, 1959. Brooklyn Ent. Soc., Bul. 54: 96 (parasite). —Krombein, 1960. Ent. News
71: 34-35 (nest, prey, cocoon, life cycle, parasite). —Krombein, 1964. Biol. Soc. Wash., Proc.
77: 99-100, fig. 17 (nest, prey, life cycle). —Thomas, 1964. Mich. Acad. Sci., Arts and
Letters, Papers 49: 199-201 (nest, parasite, life history). —Parker and Bohart, 1966.
Pan-Pacific Ent. 42: 94 (nest).

Genus PASSALOECUS Shuckard
Xyloecus Shuckard, 1837. Essay Indig. Fossor. Hym., Conspect. Gen., No. 25. Preoce.

Superfamily SPHECOIDEA 1605

Passaloecus Shuckard, 1837. Essay Indig. Fossor. Hym., p. 188. N. name.
Type-species: Pemphredon insignis Vander Linden. Orig. desig.
Coeloecus Verhoeff, 1890. Ent. Nachr. 16: 383.
Type-species: Diodontus gracilis Curtis. Desig. by Pate, 1937.
Heroecus Verhoeff, 1890. Ent. Nachr. 16: 383. No species.
Type-species: Pemphredon insignis Vander Linden. Included and desig. by Pate,
1937.

Most Passaloecus nest in pre-existing borings, cavities in wood or twigs and stems, in decay-
ing wood and in abandoned galls, although one extralimital species has been recorded as nesting
in sandy soil. Two North American species, annulatus and cuspidatus, and the Holarctic insig-
nis, have been reported in North America as making a series of linear cells with the partitions
and closing plugs made from resin. The Peckhams’ report (1905) of annulatus closing its nest
with pellets of mud is questionable and may have been based on a misidentification. However,
two species in Europe and Japan are known to form the cell partitions from grains of earth, in-
sect feces and other debris, so perhaps annulatus is not so restricted in its choice of nesting
materials as cuspidatus and insignis. Aphids are the preferred prey, but rarely, and perhaps in-
advertently, a few psyllids may be included in a cell.

Taxonomy: Krombein, 1938. Brooklyn Ent. Soc., Bul. 33: 122-127 (key to N. Y. spp.).

annulatus annulatus (Say), n. status. Ont., transcont. in U. S. Ecology: Nests in deserted
beetle burrows in wood or pith, and in logs. Prey: Drepanaphis acerifoliae (Thos.) ?
nymph, D. sp. nymph, Macrosiphum sp. nymph, Neothomasia populicola (Thos.),
Aphididae sp. Another subsp. occurs in Korea.
Pemphredon annulatus Say, 1837. Boston Jour. Nat. Hist. 1: 379. 2, ¢.
Passalaccus (!) rivertonensis Viereck, 1904. Amer. Ent. Soc., Trans. 30: 243. 6.
Passalaecus (!) equalis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 212. 9.

Biology: Peckham and Peckham, 1905. Wasps, Social and Solitary, pp. 87-89 (nest, prey, life
cycle). —Krombein, 1955. Brooklyn Ent. Soc., Bul. 50: 16 (prey). —Krombein, 1958. Biol.
Soc. Wash., Proc. 71: 24 (nest, prey). —Krombein, 1960. Ent. News 71: 35-36 (nest, prey).
—Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 65 (nest, prey).

armeniacae Cockerell. N. Mex., Ariz., Colo., Idaho, Calif., B. C.
Passaloecus armeniacae Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila.,
Proc. 49: 141. 2.

borealis Dahlbom. Alaska, western Canada, Rocky Mts. south to Utah, Colo.; north. Europe
and at higher altitudes in south.
Passaloecus borealis Dahlbom, 1844. Hym. Europaea, v. 1, p. 247. 2, d.
cuspidatus Smith. Transcont. chiefly in Transit. and U. Austr. Zones. Ecology: Nests in
borings in wood, and provisions each cell with 11-52 aphids. Parasite: Anthrax irroratus
Say; Poemenia a. americana (Cr.), P. a. nebulosa Hab. and Tow.; Chalcididae sp.;
Omalus aeneus (F.), O. purpuratus (Prov.), O. cressoni (Aar.). Prey: Cinara abieticola
(Chol.), C. formacula Hottes, Pterocomma bicolor (Oest.), Macrosiphum euphorbiae
(Thos.), M. rosae (L.), M. spp., Myzus porosus (Sand.), Masonaphis sp., Rhopalosiphum
sp., Huceraphis betulae (Koch). Predator: Philanthus pulcher D. T., P. pacificus Cr.
Passaloecus cuspidatus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 427. °.
Pemphredon mandibularis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 487. °.
Passaloecus distinctus Fox, 1892. Amer. Ent. Soc., Trans. 19: 319.
Passaloecus dispar Fox, 1892. Amer. Ent. Soc., Trans. 19: 320. d.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 131-132, figs. 60-64 (larva). —Evans,
1959. Amer. Ent. Soc., Trans. 85: 167 (larva). —Evans, 1964. Amer. Ent. Soc., Trans. 90:
250-251 (larva).

Biology: Packard, 1869. Guide Study Ins., p. 161 (nest, prey, parasite). —Krombein, 1956.
Brooklyn Ent. Soc., Bul. 51: 42-43 (nest, prey). —Krombein, 1958. Biol. Soc. Wash., Proc.
71: 24-25 (nest, prey). —Fye, 1965. Canad. Ent. 97: 740, 742 (nest, prey, parasite, life cycle).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 236-239, figs. 66, 67 (nest, prey, life
cycle, parasites). —Evans, 1973. Great Basin Nat. 33: 154-155 (nest, prey, parasites,
predators).

1606 Hymenoptera in America North of Mexico

gracilis (Curtis). Pa. and N. J. to Tex. along coast, Ohio, Ind., Mich.; west. Europe. Ecology:
Nests in borings in twigs. Adventive from Europe. Parasite: Perithous divinator
(Rossi); Omalus awratus (L.).
Diodontus gracilis Curtis, 1834. Brit. Ent., v. 11, pl. 496. d.
Passaloecus turionwm Dahlbom, 1844. Hym. Kur. 1: 246. @, d.
Passaloecus brevicornis Morawitz, 1864. Acad. Sci. St. Petersburg, Bul. 7: 462. 2, 6.

Taxonomy: Krombein, 1961. Ent. News 72: 258-259 (adventive in U. S. and separation from
N. Amer. spp.).
Biology: Krombein, 1961. Ent. News 72: 258 (nest, parasite).
marginatus (Say). Pa.
Pemphredon marginatus Say, 1837. Boston Jour. Nat. Hist. 1: 379.
melanocrus Rohwer. Tex., Colo., Ariz. Ecology: Nests in abandoned burrows of Dendroctonus
barberi Hopkins.
Passaloecus melanocrus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 561. ¢.
melanognathus Rohwer. Oreg., Calif.
Passaloecus melanognathus Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 104. 9.
monilicornis ithacae Krombein, n. status (D. Vincent). Mass., Conn., N. Y., Va., Ont., Mich.,
Minn., Alta., Alaska. Ecology: Nests in borings in twigs and wood. Parasite: Omalus
aeneus (F.), O. sp.; Poemenia albipes (Cr.), P. sp. Prey: Cinara hottesi (Gill. and Palm.),
C. abieticola (Chol.), C. fornacula Hottes, C. palmerae (Gill.), C. braggii (Gill.), C. sp.,
Neosymdobius americanus (Baker), Pterocomma smithiae Mon. ?, Amphorophora sp.,
Anuraphis rosea Baker, Rhopalosiphum fitchii (Sand.), Euceraphis betulae (Koch).
Typical monilicornis Dahlbom occurs in the Palaearctic Region.
Passaloecus ithacae Krombein, 1938. Brooklyn Ent. Soc., Bul. 33: 126. @, 6.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 249-250, fig. 22 (larva).

Biology: Fye, 1965. Canad. Ent. 97: 737-740, fig. 9 (nest, prey, parasites, life cycle).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 234-236. (nest, prey, life cycle).
relativus Fox. Wyo., Colo., Utah, Nev., Ariz. Parasite: Poemenia americana nebulosa Hab. and
Tow. Predator: Philanthus pulcher D. T., P. pacificus Cr.
Passaloecus relativus Fox, 1892. Amer. Ent. Soc., Trans. 19: 319. ¢.

Taxonomy: Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 100. @.
singularis singularis Dahlbom. Mass., N. Y., Pa., Ohio, Mich., Utah, Colo.; Europe. Ecology:
Nests in abandoned burrows of Pissodes strobi (Peck). Possibly adventive in N. Amer.
Another subsp. occurs in Mongolia.
Passaloecus singularis Dahlbom, 1844. Hym. Europaea, v. 1, p. 2438. ¢.
Passaloecus tenuis Morawitz, 1864. Acad. Imp. des Sci. St. Petersburg, Bul. 7: 462. 2, 3.
Passaloecus gertrudis Krombein, 1938. Brooklyn Ent. Soc., Bul. 33: 124. 2, d.

Genus POLEMISTUS Saussure

Polemistus Saussure, 1892. In Grandidier, Hist. Nat. Madagascar, v. 20, p. 565.
Type-species: Polemistus macilentws Saussure. Desig. by Pate, 1937.
Polymistus Ashmead, 1899. Canad. Ent. 31: 222. Emend. or lapsus.
pusillus Saussure. Utah, Ariz.; Mexico. Ecology: Nests in abandoned Trypargilwm mud nests

and makes cell partitions and nest closure from a transparent glass-like substance (resin
?), and stores 6-8 aphids per cell. Parasite: Monodontomerus sp.; Omalus sp., Chrysis
sp. Prey: Aphis gossypii Glov., A. sp., Macrosiphum sp.

Polemistus pusillus Saussure, 1892. In Grandidier, Hist. Nat. Madagascar, v. 20, p. 565. °.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 647 (nest, prey, parasites).
TRIBE STIGMINI

Genus STIGMUS Panzer

Taxonomy: Krombein, 1973. Biol. Soc. Wash., Proc. 86: 211-230, 16 figs. (synonymy,
distribution and key to N. Amer. spp.).

Superfamily SPHECOIDEA 1607

Genus STIGMUS Subgenus STIGMUS Panzer

Stigmus Panzer, 1804. Faunae Ins. German., heft 86, No. 7.
Type-species: Stigmus pendulus Panzer. Monotypic.

Gonostigmus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 559.
Type-species: Gonostigmus typicus Rohwer. Orig. desig.

Most species nest in pre-existing cavities, often beetle borings, in twigs, wood, structural
timber, shelf fungi and galls, but some tunnel in soft pith. The cells are usually placed in a linear
series except when old galls are used.

So far as known, only aphids are preyed upon.

americanus Packard. Transcont. in Canada from Labrador and N. S. to B. C. and N. W. T., in

U. S. east of 100th meridian and Wash. Ecology: Nests most commonly in old beetle
borings in dead trees or structural timber, occasionally in twigs; cells in a linear series
are separated by partitions of wood or pith particles, but there is an authentic record of
two wasps developing in a single brood chamber. Parasite: Omalus janus (Hald.), O.
iridescens (Nort.), O. purpuratus (Prov.). Prey: Drepanaphis acerifoliae (Thos.) nymphs,
D. sp. nymphs and adult, Rhopalosiphum sp. nymphs, Aphis sp. nymph, Anuraphis sp.
nymphs, Myzocallis ? sp. nymph, Chaitophorus ? sp. nymph, Therioaphis sp. nymphs
and adults, Aphidini sp., Panaphidini sp. nymphs and adult, Aphidinae sp. nymph.

Stigmus americanus Packard, 1867. Ent. Soc. Phila., Proc. 6: 386. 2, d.

Stigmus lucidus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 102. ¢.

Stigmus fraternus coloradensis Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 559. °.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 44-45 (nest, prey,
parasite). —Krombein, 1954. Brooklyn Ent. Soc., Bul. 49: 6 (nest, prey). —Krombein, 1955.
Brooklyn Ent. Soc., Bul. 50: 16 (nest, prey). —Krombein, 1956. Brooklyn Ent. Soc., Bul. 51:
42. (prey). —Krombein, 1958. Biol. Soc. Wash., Proc. 71: 24 (nest, prey). —Krombein, 1961.
Brooklyn Ent. Soc., Bul. 56: 64-65 (nest, prey, life cycle).

aphidiperda Rohwer. Pa. to N. C. Ecology: Nests in peach twig. Prey: Aphis persicaeniger Sm.
Stigmus aphidiperda Rohwer, 1911. U.S. Natl. Mus., Proce. 40: 558. 2, ¢.
fraternus Say. Mass. and N. H. to S. C., W. Va., Tenn., Mo. Ecology: Nests in twigs, old galls
and beetle borings in structural timber. Prey: Monellia sp. nymphs, Therioaphis sp.
nymphs, Aphididae spp.
Stigmus fraternus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 340.
Stigmus conestogorum Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 557. 9.
Stigmus fraternus raui Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 100. 2, 6.

Biology: Packard, 1869. Guide Study Ins., p. 161 (nest). —Rau, 1928. Acad. Sci. St. Louis,
Trans. 25: 379-381 (nest, prey, life cycle). —Krombein, 1958. Biol. Soc. Wash., Proc. 71: 24
(nest, prey).

fulvicornis Rohwer. Miss. (Starksville). Ecology: Nesting in floors of porch and in house. Prey:
Aphididae sp.
Stigmus fulvicornis Rohwer, 1923. Wash. Acad. Sci., Jour. 13: 370. 2.
Biology: Smith, 1923. Jour. Econ. Ent. 16: 553-554 (nest, prey).
hubbardi Rohwer. Colo., Utah, N. Mex., Ariz., Calif., Wash. Ecology: Nests in the shelf fungus

Polyporus.
Stigmus inordinatus hubbardi Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 559. 9, 6.

Biology: Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 559 (nest).
inordinatus inordinatus Fox. Mont., Colo. and N. Mex. west to B. C. and Calif. Ecology: Nests

in cavities or borings in twigs or stems of mulberry, peony, peach, raspberry and
elderberry, constructing a linear series of cells stored with 12-30 aphids per cell.
Parasite: Omalus variatus (Aar.), O. glomeratus (Buyss.), O. cressoni (Aar.). Prey:
Aphis frangulae Kalt. ?, Aphididae sp.

Stigmus inordinatus Fox, 1892. Amer. Ent. Soc., Trans. 19: 322. 9, ¢.

Stigmus fulvipes var. coquilletti Rohwer, 1911. U. 8S. Natl. Mus., Proce. 40: 559. 2, 6.

Stigmus reticulatus Mickel, 1918 (1917). Nebr. Univ., Studies 17: 330. 9.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 134-135, figs. 68, 69 (larva).

1608 Hymenoptera in America North of Mexico

Biology: Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 102 (prey). —Wasbauer and Simonds,
1964. Pan-Pacifie Ent. 40: 114-116, 1 fig. (mest, prey, cocoon). —Parker and Bohart, 1966.
Pan-Pacifie Ent. 42: 95 (nest, parasites).

inordinatus universitatis Rohwer. Conn. to Va., W. Va., Ill., Colo. Ecology: Nests in old insect
galls on oak. Prey: Aphididae sp.
Stigmus inordinatus universitatis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 102. 2.

Biology: Richardson, 1915. Psyche 22: 104-105 (nest, prey). —Krombein, 1973. Biol. Soc.
Wash., Proc. 86: 224 (nest).
podagricus podagricus Kohl. South. Tex. and Ariz.; Mexico (Veracruz, Morelos).
Stigmus podagricus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 65. d.
podagricus tarsalis Krombein. Ga., Fla., Ala., centr. and north. Tex.
Stigmus (Stigmus) podagricus tarsalis Krombein, 1973. Biol. Soc. Wash., Proc. 86: 225,
figs. 6, 16. d, 9.

Genus STIGMUS Subgenus ATOPOSTIGMUS Krombein

Stigmus subg. Atopostigmus Krombein, 1973. Biol. Soc. Wash., Proc. 86: 218.
Type-species: Stigmus fulvipes Fox. Orig. desig.
The single known species apparently nests in abandoned burrows of other insects in the soil.
The preferred prey is unknown.

fulvipes Fox. B. C. transcont. in U. S.; Mexico (Morelos). Ecology: Nests in sand or mud banks
in abandoned burrows of other insects.
Stigmus fulvipes Fox, 1892. Amer. Ent. Soc., Trans. 19: 324. °.

Taxonomy: Krombein, 1952. Amer. Ent. Soc., Trans. 78: 94 (separation from eastern spp.).
Biology: Krombein, 1973. Biol. Soc. Wash., Proc. 86: 218 (nesting site).

NOMEN NUDUM IN STIGMUS PANZER

coloradensis Ashmead. Colo.
Stigmus coloradensis Ashmead, 1889. In Cockerell, Colo. Biol. Assn., 10th Rept., p. 2.
Probably published originally in Custer County Courant newspaper.

Genus SPILOMENA Shuckard

Celia Shuckard, 1837. Essay Indig. Fossor. Hym., p. 182. Preocce.
Type-species: Stigmus troglodytes Vander Linden. Monotypic.
Spilomena Shuckard, 1838. Ent. Soc. London, Trans. 2: 79 (footnote). N. name.
Microglossa Rayment, 1930. Roy. Soc. Victoria, Proc. (n. s.) 42: 212. Preoce.
Type-species: Microglossa longifrons Rayment. Orig. desig.
Microglossella Rayment, 1935. A Cluster of Bees, p. 634. N. name.
Taialia Tsuneki, 1971. Life Study (Fukui) 15: 10.
Type-species: Taialia formosana Tsuneki. Orig. desig.

Biological notes have been recorded for only three North American species. All of them nest
in abandoned borings of anobiid beetles in structural timber or logs. They prey upon Thysanop-
tera, usually immatures although occasionally adults are captured. Some extralimital species
nest in twigs or decayed wood; in addition to Thysanoptera these species also prey upon imma-
ture Psyllidae, Coccidae and Aphididae. About a dozen undescribed species occur in the
southwestern deserts.

Taxonomy: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 54, 60-61, figs. 2-6 (key to eastern
Spp.).
alboclypeata Bradley. B. C., Oreg., Calif., Idaho, Mont., Utah, Ariz., N. Mex., Colo., Kans., W.
Va., Va. Ecology: Nests in abandoned borings of anobiid beetles. Prey: Immature
Thysanoptera.
Spilomena alboclypeata Bradley, 1906. Canad. Ent. 38: 380. 3.

Taxonomy: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 60, figs. 4, 4a, 6. 2, 6.
Biology: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 62 (prey, nest).

Superfamily SPHECOIDEA 1609

ampliceps Krombein. W. Va. (Lost River St. Pk.). Ecology: Presumably nests in old beetle
borings in logs.
Spilomena ampliceps Krombein, 1952. Ent. Soc. Wash., Proce. 54: 178, figs. 1-3. d, 2.
Taxonomy: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 60, figs. 3, 3a. 2, d.
barberi Krombein. Mass., N. Y. and Ont. south to Ga. and Tenn., Iowa, Kans., Colo., N. Mex.,
Ariz., Utah, Calif. Ecology: Nests in abandoned anobiid borings in wood. Prey:
Sericothrips sp. larva, Frankliniella or Thrips sp. larva, Thripidae sp. larva.
Spilomena barberi Krombein, 1962. Biol. Soc. Wash., Proc. 75: 12. d, 2.
foxii Cockerell. Colo., N. Mex., Ariz., Calif., Wash.; Mexico (Baja California, Puebla).
Spilomena foxti Cockerell, 1897. Entomologist 30: 136. 2.
pusilla (Say). Conn. to N. C., west to Iowa, Kans. and Tenn. Ecology: Nests in abandoned
anobiid burrows in structural timber and in logs. Prey: Sericothrips sp. larva in
variabilis (Beach) section, Thripidae spp. larvae and adult.
Stigmus pusillus Say, 1837. Boston Jour. Nat. Hist. 1: 378. 2, d.
Taxonomy: Krombein, 1952. Ent. Soc. Wash., Proc. 54: 178-179, figs. 4-6. 2, ¢. —Krombein,
1958. Ent. Soc. Wash., Proc. 60: 53, 61, figs. 2, 2a, 5. 2, 6.

Biology: Krombein, 1952. Ent. Soc. Wash., Proc. 54: 181 (nest). —Krombein, 1956. Ent. Soe.
Wash., Proc. 58: 155 (nest, prey). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 53 (nest,

prey).
Genus XYSMA Pate

Xysma Pate, 1937. Amer. Ent. Soc., Trans. 63: 94.
Type-species: Ammoplanus ceanothae Viereck. Orig. desig.

Telexysma Leclercq, 1959. Pare Natl. Upemba, I. Mission G. Witte, fasc. 53 (2), p. 2.
Type-species: Telexysma africana Leclercq. Orig. desig.

The North American ceanothae nests in abandoned anobiid borings in structural timber and
preys upon immature Thysanoptera. Nothing is known of the biology of the only other Xysma,
the South African africana.
ceanothae (Viereck). Pa., Md., D. C., Va., Ga. Ecology: Nests in abandoned anobiid borings in

structural timber. Prey: Thripinae spp. larvae.
Ammoplanus ceanothae Viereck, 1904. Psyche 11: 72. 2 (2 in part, 6, misdet.).

Taxonomy: Krombein, 1958. Biol. Soc. Wash., Proc. 71: 25. d.

Biology: Krombein, 1958. Biol. Soc. Wash., Proc. 71: 25 (nest, prey).
TRIBE AMMOPLANINI
Genus PULVERRO Pate

Pulverro Pate, 1937. Amer. Ent. Soc., Trans. 63: 107.
Type-species: Pulverro mescalero Pate. Orig. desig.
This genus has been recorded only from the Nearctic Region. One species has been recorded
as nesting in soil and preying upon Thysanoptera, mostly adults but a few immatures.
Revision: Pate, 1937. Amer. Ent. Soc., Trans. 63: 107-119, figs. 4, 6, 7, 9-12, 16, 20.

californicus Eighme. Calif. in North Coast Ranges.
Pulverro californica Eighme, 1973. Pan-Pacific Ent. 49: 49. 3, 9.
chumashano Pate. West. Calif.
Pulverro chumashano Pate, 1937. Amer. Ent. Soce., Trans. 63: 118. 6.
Pulverro costano Pate, 1937. Amer. Ent. Soc., Trans. 63: 116. @.

Taxonomy: Pate, 1939. Amer. Ent. Soc., Trans. 64: 417, fig. 24 (female chumashano).

columbianus (Kohl). Colo., Utah, Idaho, Oreg., B. C.
Ammoplanus (?) columbianus Kohl, 1890. K. K. Naturhist. Hofmus., Ann. 5: 61. “2” = 6.
Ammoplanus eriogoni Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 101. d.
Pulverro colorado Pate, 1937. Amer. Ent. Soc., Trans. 63: 115. 2.

1610 Hymenoptera in America North of Mexico

constrictus:(Provancher). South. Calif.
Anacrabro constrictus Provancher, 1895. Nat. Canad. 22: 141. @.
Pulverro serrano Pate, 1937. Amer. Ent. Soc., Trans. 63: 114, fig. 9. 3.

Taxonomy: Pate, 1939. Amer. Ent. Soc., Trans. 64: 418, fig. 6 (female serrano).
laevis (Provancher). Calif. (Los Angeles).
Anacrabro laevis Provancher, 1895. Nat. Canad. 32: 142. °.

mescalero Pate. Tex., N. Mex., Ariz.; n. and cent. Mexico.
Ammoplanus mandibularis Ashmead, 1898. In Cockerell, Davenport Acad. Nat. Sci., Proc.
7: 148. Nom. nud.
Pulverro mescalero Pate, 1937. Amer. Ent. Soc., Trans. 63: 111. 2, d.

Taxonomy: Pate, 1939. Amer. Ent. Soc., Trans. 64: 411.

monticola Eighme. Calif. in North Coast Ranges. Ecology: Nests in sloping banks, constructs
several cells off side burrows, and provisions each with up to 21 thrips. Prey:
Frankliniella moultoni Hood, Aeolothrips fasciatus (L.); mostly adults and some larvae.
Pulverro monticola Eighme, 1968. Pan-Pacifie Ent. 44: 263, figs. 1-7. 6, ?.

Taxonomy: Bohart and Grissell, 1972. Pan-Pacific Ent. 48: 148-149, figs. 1-4 (larva).
Biology: Bohart and Grissell, 1972. Pan-Pacific Ent. 48: 145-148, fig. 5 (nest, prey, life cycle).

Genus AMMOPLANOPS Gussakovskij

Ammoplanops Gussakovskij, 1931. Soc. Espan. Hist. Nat., Bol. 31: 457.
Type-species: Ammoplanops carinatus Gussakovskij. Orig. desig.

Nothing is known of the nesting habits or prey. The presence of a tarsal pecten and pygidium
in the female suggests that the species are ground-nesting.

Revision: Pate, 1939. Amer. Ent. Soc., Trans. 64: 392-411, figs. 3, 10-12, 17-23, 25.

ashmeadi Pate. Calif. (Palm Springs).

Ammoplanops ashmeadi Pate, 1939. Amer. Ent. Soc., Trans. 64: 397. 2, 6.
cockerelli (Ashmead). Tex., N. Mex., Colo., Utah, Ariz., Calif.

Ammoplanus cockerelli Ashmead, 1903. Ent. News 14: 295. 2.
cressoni Pate. Ariz., Nev., Calif.

Ammoplanops cressoni Pate, 1939. Amer. Ent. Soc., Trans. 64: 395. ¢.
foxi Pate. Calif. (Clark Mt. in San Bernardino Co.).

Ammoplanops foxi Pate, 1939. Amer. Ent. Soc., Trans. 64: 409. ¢.
moenkopi Pate. N. Mex., Ariz., Utah.

Ammoplanops moenkopi Pate, 1939. Amer. Ent. Soc., Trans. 64: 402. ¢.
timberlakei Pate. Calif. (Andreas Canyon in Riverside Co.).

Ammoplanops timberlakei Pate, 1939. Amer. Ent. Soe., Trans. 64: 404. ¢.
vierecki Pate. N. Mex. (Alamogordo).

Ammoplanops vierecki Pate, 1939. Amer. Ent. Soc., Trans. 64: 406. 2, 6.

Genus AMMOPLANUS Giraud

Ammoplanus Giraud, 1869. Soc. Ent. France, Ann. (4) 9: 469.
Type-species: Ammoplanus Perrisi Giraud. Desig. by Pate, 1937.
Hoplocrabron De Stefani, 1886. Nat. Sicil. 6: 60.
Type-species: Hoplocrabron Marathroicus De Stefani. Monotypic.
Ammoplanus subg. Ceballosia Giner Mari, 1943. Eos 19: 293. Preoce.
Type-species: Ammoplanus rjabovi Gussakovskij. Orig. desig.

Nothing is known definitely as to the ethology of our species. In Europe and Africa species of
this genus nest in pre-existing cracks in stone, wood or soil and provision the cells with imma-
ture Thysanoptera.

Revision: Pate, 1943 (1942). South. Calif. Acad. Sci., Bul. 41: 144-157, figs. 1, 5, 6, 8-12 (N.
Amer. spp.).

Superfamily SPHECOIDEA 1611

chemehuevi Pate. Calif. (Riverside Co.).
Ammoplanus (Ammoplanus) chemehuevi Pate, 1943 (1942). South. Calif. Acad. Sci., Bul.

ANSI So 10):

loti Pate. Calif. (Riverside).
Ammoplanus (Ammoplanus) loti Pate, 1943 (1942). South. Calif. Acad. Sci.,.Bul. 41: 146.

2.

quabajai Pate. Calif.
Ammoplanus (Ammoplanus) quabajai Pate, 1943 (1942). South. Calif. Acad. Sci., Bul. 41:

156. ?.

sechi Pate. Calif. (Riverside Co.).
Ammoplanus (Ammoplanus) sechi Pate, 1943 (1942). South. Calif. Acad. Sci., Bul. 41: 149.
3

tetli Pate. Calif. (Tetly Park, San Bernardino Mts.).
Ammoplanus (Ammoplanus) tetli Pate, 1943 (1942). South Calif. Acad. Sci., Bul. 41: 147.
é.

unami Pate. Pa., W. Va. Ecology: Presumably nests in abandoned beetle borings in logs.
Ammoplanus (Ammoplanus) unami Pate, 1937. Amer. Ent. Soc., Trans. 63: 101. 9.

Taxonomy: Krombein, 1956. Ent. Soc. Wash., Proc. 58: 159, figs. 2, 2a (male).

Biology: Krombein, 1956. Ent. Soc. Wash., Proc. 58: 160 (putative nest site).

vanyumi Pate. Calif., Idaho.
Ammoplanus (Ammoplanus) vanyumi Pate, 1943 (1942). South. Calif. Acad. Sci., Bul. 41:
154. 9.

Genus AMMOPLANELLUS Gussakovskij

Revision: Pate, 1943 (1942). South. Calif. Acad. Sci., Bul. 41: 158-162, figs. 2-4, 7 (N. Amer.
spp.).
Genus AMMOPLANELLUS Subgenus AMMOPLANELLUS Gussakovskij

Ammoplanus subg. Ammoplanellus Gussakovskij, 1931. Soc. Espan. Hist. Nat., Bol. 31:

442.
Type-species: Ammoplanus (Ammoplanellus) chorasmius Gussakovskij. Orig.

desig.
The species are thought to nest in pre-existing holes or crannies in wood, and possibly to
provision the cells with Thysanoptera.
umatilla (Pate). Wash.
Ammoplanus (Ammoplanellus) umatilla Pate, 1945. Pan-Pacific Ent. 21: 82. 9.

xila (Pate). Ariz. (Phoenix). Ecology: Possibly nesting in chair containing burrows of
Hesperorhipis mirabilis Knull.
Ammoplanus (Ammoplanellus) xila Pate, 1945. Pan-Pacific Ent. 21: 84. 9.

Genus AMMOPLANELLUS Subgenus PARAMMOPLANUS Pate

Ammoplanus subg. Parammoplanus Pate, 1939. Amer. Ent. Soc., Trans. 64: 391.
Type-species: Ammoplanus (Ammoplanellus) apache Pate. Orig. desig.

apache (Pate). N. Mex., Ariz., Calif.
Ammoplanus (Ammoplanellus) apache Pate, 1937. Amer. Ent. Soc., Trans. 63: 106. 3.

lenape lenape (Pate). Pa. (Lehigh Gap).
Ammoplanus (Ammoplanellus) lenape Pate, 1937. Amer. Ent. Soc., Trans. 63: 104. d.

lenape olamentke (Pate). Calif., Mont.
Ammoplanus (Parammoplanus) lenape olamentke Pate, 1943 (1942). South. Calif. Acad.

Sci., Bul. 41: 160. 2, 6.
Genus TIMBERLAKENA Pate
Timberlakena subg. Riparena Pate, 1939. Amer. Ent. Soc., Trans. 64: 378.

1612 Hymenoptera in America North of Mexico

Type-species: Timberlakena (Riparena) cahuilla Pate. Orig. desig.
Timberlakena subg. Mojavena Pate, 1939. Amer. Ent. Soc., Trans. 64: 381.

Type-species: Timberlakena (Mojavena) yucaipa Pate. Orig. desig.
Timberlakena subg. Timberlakena Pate, 1939. Amer. Ent. Soc., Trans. 64: 383.

Type-species: Timberlakena (Timberlakena) nolcha Pate. Orig. desig.

Females lack a tarsal pecten and pygidium so it is presumed that they may nest in
pre-existing cavities, such as abandoned beetle borings, in twigs or structural lumber.

Revision: Pate, 1939. Amer. Ent. Soc., Trans. 64: 374-390, figs. 1, 2, 4, 5, 7-9, 13-16.
cahuilla Pate. Calif. (Riverside Co.).
Timberlakena (Riparena) cahuilla Pate, 1939. Amer. Ent. Soc., Trans. 64: 378. 2, d.
hualga Pate. Calif.
Timberlakena (Timberlakena) nolcha hualga Pate, 1939. Amer. Ent. Soc., Trans. 64: 388.
OPE
nolcha Pate. Ariz., Calif., Idaho.
Timberlakena (Timberlakena) nolcha nolcha Pate, 1939. Amer. Ent. Soc., Trans. 64: 385.
Ge
ocha Pate. Calif. (Riverside Co.), Ariz.
Timberlakena (Timberlakena) ocha Pate, 1939. Amer. Ent. Soc., Trans. 64: 389. 9.
yucaipa Pate. Calif. (San Bernardino Co.).
Timberlakena (Mojavena) yucaipa Pate, 1939. Amer. Ent. Soc., Trans. 64: 381. 2, d.

Family ASTATIDAE

So far as is known all members of this family are ground-nesting forms, making one or more
cells per nest, and preying principally upon nymphal Hemiptera. The only group that occurs in
the New World is the typical subfamily Astatinae.

SUBFAMILY ASTATINAE

Taxonomy: Maidl and Klima, 1939. Hym. Cat., Pars 8, Sphecidae, pp. 7-29 (world catalog).
—Evans, 1958. Amer. Ent. Soc. Trans. 84: 109-113, Figs. 1-8, 46-48 (larvae). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 165-166 (larvae). —Parker, 1962. Ent. Soc. Amer., Ann. 55:
643-644 (key to genera). —Parker, 1966. Ent. Soc. Amer., Ann. 59: 765-766, 10 figs. (key to
N. Amer. genera).

Biology: Evans, 1958 (1957). N. Y. Ent. Soc., Jour. 65: 159-185, 8 figs. (ethology of world
Spp.).

Genus DIPLOPLECTRON Fox

Diploplectron Fox, 1893. Amer. Ent. Soc., Trans. 20: 38.
Type-species: Liris ? brunneipes Cresson. Monotypic.

Revision: Parker, 1972. Ent. Soc. Amer., Ann. 65: 1192-1203, 37 figs. (N. Amer. spp.).

Taxonomy: Ashmead, 1899. Ent. News 10: 55-56 (key to N. Amer. spp.). —Rohwer, 1909.
Amer. Ent. Soc., Trans. 35: 123-124 (key to N. Amer. spp.). —Krombein, 1939. Brooklyn
Ent. Soc., Bul. 34: 138-1389 (key to N. Amer. spp.).

beccum Parker. Calif., Nev., Utah, Ariz., southwest. Tex.
Diploplectron beccwm Parker, 1972. Ent. Soc. Amer., Ann. 65: 1195, figs. 5, 19, 32. 6, 9.

brunneipes (Cresson). Idaho, Wyo., Colo., Calif. Ecology: Makes a nest of 2-3 cells in

hard-packed sandy clay, stores 6 prey per cell. Prey: Uhleriola floralis (Uhler), nymphs.
Predator: Philanthus pacificus Cr.

Liris ? brunneipes Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proc. p. iii. 2 (misdet. in
part), ¢.

Diploplectron bidentatus Ashmead, 1899. Ent. News 10: 56. “2” = 6.

Diploplectron foxii Ashmead, 1899. Ent. News 10: 56. @.

Diploplectron bidentatiformis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 121. 3.

Superfamily SPHECOIDEA 1613

Taxonomy: Krombein, 1939. Brooklyn Ent. Soc., Bul. 34: 138 (synonymy). —Evans, 1975.
Great Basin Nat. 35: 123-125, 5 figs. (larva).

Biology: Evans, 1975. Great Basin Nat. 35: 123 (nest, prey).
californicum Parker. Calif., Oreg., Idaho. Prey: Rhyparochromus californicus Van D. adults,
Emblethis vicarius Horv. ? nymphs, Megalonotus chiragrus (F.) adult.
Diploplectron californicum Parker, 1972. Ent. Soc. Amer., Ann. 65: 1199, figs. 1, 13, 25. d,
0

Biology: Williams, 1946. Hawaii. Ent. Soc., Proc. 12: 648 (prey).

diablense Williams. Coast Range and Sierra Nevada Mts. of Calif.
Diploplectron diablensis Williams, 1951 (1950). Wasmann Jour. Biol. 8: 363, fig. 1. d.

Taxonomy: Williams, 1959. Wasmann Jour. Biol. 17: 303, fig. 5. 2.

ferrugineum Ashmead. Idaho, Colo., N. Mex., Ariz., Calif.
Diploplectron ferrugineus Ashmead, 1899. Ent. News 10: 56. “2” = 6.
Diploplectron ashmeadi Rohwer, 1909. Amer. Ent. Soc. Trans. 35: 122. 3.
Diploplectron cressoni Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 123. 9.
Diploplectron relativus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 123. 9.

fossor Rohwer. B. C., Wyo., Colo., Utah, Nev., Calif. Prey: Labops sp. nymphs. Predator:

Philanthus pacificus Cr.

Diploplectron fossor Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 120. 2.
Diploplectron rufoantennatus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 120. ¢.

Biology: Parker, 1972. Ent. Soc. Amer., Ann. 65: 1193 (prey).
irwini Parker. South Ariz.
Diploplectron irwini Parker, 1972. Ent. Soc. Amer., Ann. 65: 1200, figs. 7, 12. 6.
kantsi Pate. Southwest. Tex., south. N. Mex.; Mexico (Tamaulipas).
Diploplectron kantsi Pate, 1941. Ent. News 52: 6. 6d.
peglowi Krombein. N. Y., N. W. T., Wyo., Colo., Utah, Idaho, Nev., Calif. Ecology: Nests in
sand, making either a unicellular nest or 3-4 cells per nest and storing 2-6 prey per cell.
Parasite: Hedychridium sp. ? Prey: Sphaerobius insignis (Uhl.), Lygaeus sp., Emblethis
vicarius Horv., Sphragisticus nebulosus (Fall.); Aufeius impressicollis Stal. All prey
specimens were nymphs.
Diploplectron peglowi Krombein, 1939. Brooklyn Ent. Soc., Bul. 34: 136. 2, 6.

Biology: Williams, 1946. Hawaii. Ent. Soc., Proc. 12: 648 (prey, nest). —Parker, 1972. Ent.
Soc. Amer., Ann. 65: 1193 (nest, prey). —Kurezewski, 1972. Ent. Soc. Wash., Proc. 74:
385-397, 4 figs. (nest, prey hunting and transport). —Kurezewski, 1975. Ent. Soc. Wash.,
Proc. 77: 97-99 (nest, prey, egg).

reticulatum Williams. South. Calif. and Ariz.
Diploplectron reticulatus Williams, 1946. Hawaii. Ent. Soc., Proc. 12: 648. d.

Taxonomy: Williams, 1951 (1950). Wasmann Jour. Biol. 8: 365, fig. 2. 9.
secoense Parker. Calif. in Coast Range and Sierra Nevada Mts.
Diploplectron secoense Parker, 1972. Ent. Soc. Amer., Ann. 65: 1201, figs. 7, 20, 29. gd, 9.
sierrense Parker. Sierra Nevada Mts. of Calif. and Nev.
Diploplectron sierrense Parker, 1972. Ent. Soc. Amer., Ann. 65: 1199, figs. 5, 15, 30, 36, 37.
Crm.
vierecki Pate. South. Ariz. and N. Mex., southwest. Tex. Prey: Microporus obliquus Uhl.
nymph.
Diploplectron vierecki Pate, 1941. Ent. News 52: 4. 6.

Genus ASTATA Latreille

Astatus Latreille, 1796. Precis Caract. Gen. Ins., p. 114. No species.
Astata Latreille, 1796. Precis Caract. Gen. Ins., p. xiii. Emend.
Type-species: Tiphia abdominalis Panzer. Desig. by Latreille. First included
species. Astata placed on Official List of Generic Names, Op. 139, Op. and Declar.
Internatl. Comn. Zool. Nomencl., v. 2, sect. A, p. 37, 1943.

1614 Hymenoptera in America North of Mexico

Dimorpha Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, p. 126.
Type-species: Tiphia abdominalis Panzer. Monotypic.

The species of Astata build multicellular nests in soil of varying types ranging from sandy to
hard-packed with included gravel. The cells may be in short series in some species separated by
earthen partitions. They prey upon Hemiptera, principally Pentatomidae and Lygaeidae, and
usually the nymphal stages.

Revision: Fox, 1892. Canad. Ent. 24: 232-235 (N. Amer. spp.). —Fox, 1894 (1893). Acad. Nat.
Sci. Phila., Proc. 45: 539-548 (N. Amer. spp.). —Parker, 1962. Ent. Soc. Amer., Ann. 55:
643-659, 42 figs. (spp. north of Mexico). —Parker, 1964. Ent. Soc. Amer., Ann. 57: 552-559,
21 figs. (Mexican and Cent. Amer. spp.).

bakeri Parker. Ont., Sask. to B. C., south to Tex. and Calif.; Mexico (Chihuahua, Jalisco,
Guanajuato, Morelos, Puebla, Chiapas). Prey: Nysius raphanus How. adult and nymph,
Lygaeus bicrucis Say ? nymph. Predator: Philanthus crabroniformis Sm.
Astata bakeri Parker, 1962. Ent. Soc. Amer., Ann. 55: 647, text fig. 1, figs. 11, 21. d, .

Biology: Williams, 1946. Hawaiian Ent. Soc., Proc. 12: 647 (prey; rec. as Astata sp.).
bechteli Parker. Desert areas, Wash. and Colo., south to Calif. and Tex.; Mexico (Sonora).
Astata nubecula bechteli Parker, 1962. Ent. Soc. Amer., Ann. 55: 649. d, °.
bicolor Say. Southern Canada and U. S. east of Rocky Mts.; Mexico (Chihuahua, Sonora,
Durango, San Luis Potosi, Jalisco, Guanajuato, Mexico). Ecology: Nests in hard-packed
sand, sometimes beneath overhanging vegetation, provides up to 5 cells per nest with
6-8 nymphs per cell. Prey: Pentatomidae spp.
Astata bicolor Say, 1823. West. Quart. Rptr. 2: 78. d, 9.
Astata terminata Cresson, 1872. Amer. Ent. Soc., Trans. 4: 218. d.
Astatus pygidialis Fox, 1892. Canad. Ent. 24: 234. °.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 165-166 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 92-97, 217, pls. 1,
4, 9 (nest, prey). —Peckham and Peckham, 1905. Wasps Social and Solitary, pp. 287-289
(orientation flight). —Mickel, 1918 (1917). Nebr. Univ., Studies 17: 108 (prey). —Evans,
1962. N. Y. Ent. Soc., Jour. 70: 30-31 (nest, prey).

bigeloviae Cockerell. Western Tex. to Calif.; Mexico (Jalisco).
Astatus bigeloviae Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila., Proc. 49:
138. 9.
boharti Parker. N. Mex., Ariz.
Astata boharti Parker, 1962. Ent. Soc. Amer., Ann. 55: 650, fig. 25. d, 9.
clypeata Parker. East slope of Rocky Mts. in U. S.; Mexico (western mts.).
Astata clypeata Parker, 1962. Ent. Soc. Amer., Ann. 55: 651, figs. 15, 26. 6, °.
femorata Parker. Ariz. (Pena Blanea in Santa Cruz Co.).
Astata femorata Parker, 1963. Pan-Pacific Ent. 39: 185, figs. 1-3, 3.
leuthstromi Ashmead. Alaska, Canada, northern U. S., south in mts. to Ariz. Ecology: Nests in
soil, sometimes beneath overhanging vegetation, provides 2 cells per nest with up to 5
nymphal prey per cell. Prey: Acrosternum hilare Say, Cosmopepla bimaculata Thom.
Astata leuthstromi Ashmead, 1897. Psyche 8: 129. °.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 98 (nest, prey).
—Evans, 1958 (1957). N. Y. Ent. Soc., Jour. 65: 176-177 (nest, prey transport). —Evans,
1962. N. Y. Ent. Soc., Jour. 70: 31-32 (nest, prey).

mexicana Cresson. Canada and U. S. west of Rocky Mts.; Mexico south to El Salvador.
Astata mexicana Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proc. p. v. d.
nevadica Cresson. Western Canada and U. S.; Mexico (Chihuahua, Sonora, Durango, Zacatecas,
Guanajuato, Mexico).
Astata nevadica Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proce. p. v. °.
nubecula Cresson. N. W. T., northern U. S. south to Va. in east and along mts. in Great Basin
and in Calif. to San Diego. Ecology: Nests in hard, bare stony soil or in sand, provides
up to 8 cells per nest, each cell provisioned with 2-4 nymphs. Parasite: Senotainia

Superfamily SPHECOIDEA 1615

trilineata Wulp, Hilarella hilarella Zett. ? Prey: Chlorochroa uhleri Stal ?, Thyanta sp.,
prob. casta Stal or pallidovirens Stal, Pentatomidae sp. Predator: Philanthus zebratus
nitens (Bks.), P. crabroniformis Sm.

Astata nubecula Cresson, 1865. Ent. Soe. Phila., Proc. 4: 466. 6.

Astata nigropilosa Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proce. p. iv. d, 2.

Biology: Parker, 1962. Ent. Soc. Amer., Ann. 55: 649 (nest, prey). —Evans, 1970. Mus.
Compar. Zool., Bul. 140: 487-488 (male territoriality, nest, prey, parasites).
occidentalis Cresson. South. Canada, U. S.; Mexico (Chihuahua, Baja California, Guanajuato,
Morelos, Michoacan, Puebla). Ecology: Nests in hard-packed clay devoid of vegetation,
makes up to 14 cells per nest, and stores 3-9 usually adult prey per cell. Parasite:
Senotainia trilineata Wulp; Diptera sp. Prey: Hymenarcys nervosa Say, Thyanta
calceata (Say), T. p. pallidovirens Stal, T. p. accerra McAtee, T. brevis Van D., T.
custator F., T. punctiventris Van D., Euschistus variolarius (Beauv.), Holcostethus
limbolarius (Stal), Banasa calva (Say), Perillus bioculatus (F.), Trichopepla sp.
Astata occidentalis Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proce. p. iii. dé.
Astata apicipennis Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 66., pl. 5, fig. 1. 9°.
Astata tinctipennis Cameron, 1890. Biol. Cent.-Amer., Hym. v. 2, p. 67. ¢.
Astatus sayi Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 542. 9.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 110-111, figs. 1-8 (larva). —Parker, 1964.
Ent. Soe. Amer., Ann. 57: 559 (synonymy).

Biology: Evans, 1958 (1957). N. Y. Ent. Soc., Jour. 65: 168-176, figs. 3-8 (nest, prey transport,
parasite). —Powell and Burdick, 1960. Pan-Pacific Ent. 36: 25-30 (nest, prey transport,
parasite).

unicolor Say. Southern Canada, U. S.; Mexico to El Salvador, Cuba, Jamaica. Ecology: Nests

usually beneath overhanging vegetation in hard-packed sand or heavier soil containing
gravel, makes 12-14 cells in a complete nest, and stores 2-4 nymphal prey per cell.
Parasite: Chrysis sp. Prey: Podisus modestus F., P. maculiventris (Say), Euschistus
tristigmus Say, E. euschistoides (Voll.) ?

Astata unicolor Say, 1824. In Keating, Narr. Long’s 2nd Exped. v. 2, p. 337. 9.

Astata insularis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 140. @.

Astata rufiventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 218. 2.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 111-112, figs. 46-48 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 88-92, pls. 9, 11,
12 (nest, prey, parasite). —Peckham and Peckham, 1905. Wasps Social and Solitary, pp.
289-290 (orientation flight). —Evans, 1958 (1957). N. Y. Ent. Soc., Jour. 65: 160-168, figs. 1,
2 (nest, prey hunting and transport, male behavior, life cycle).

Morphology: Snodgrass, 1941. Smithsn. Inst., Misc. Collect. 99 (14): pl. 20, figs. L, M (male
genitalia).
williamsi Parker. Oreg., Idaho, Utah, Nev., Calif.
Astata williamsi Parker, 1962. Ent. Soc. Amer., Ann. 55: 653, figs. 14, 29, 35, 36. d, °.

Genus DRYUDELLA Spinola

Dryudella Spinola, 1843. Soc. Ent. France, Ann. (2) 1: 185.
Type-species: Dryudella ghilianii Spinola. Desig. by Verhoeff, 1951.

Members of this genus have a wider prey range than other genera of the subfamily. In addi-
tion to Pentatomidae and Lygaeidae, they have been reported to prey upon Scutelleridae, Redu-
viidae, Cydnidae, Alydidae and Rhopalidae; both nymphs and adults are preyed upon.

Revision: Parker, 1969. Ent. Soc. Amer., Ann. 62: 963-976, 67 figs. (New World spp.).
bella (Cresson). Idaho, Nev., Calif., Ariz., N. Mex.; Mexico.
Astata bella Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proce. p. vi. d.
caerulea (Cresson). Wash., Idaho, Colo., Nev., Calif., Ariz., N. Mex.; Mexico south to Puebla.
Ecology: Nests in sand under vegetation, 3 cells per nest, with 4-6 prey per cell. Prey:
Hyalmenus tarsatus (F.) nymphs; Euschistus conspersus Uhl. nymphs.
Astata caerulea Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proce. p. iv. ¢.

1616 Hymenoptera in America North of Mexico

Biology: Alcock, 1973. Wasmann Jour. Biol. 31: 328-329, fig. 4 (nest, prey).
elegans (Cresson). Idaho, Wyo., Utah, Nev., Calif., Ariz.
Astata elegans Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proe. p. vi. 6, 2.
immigrans (Williams). Wash., Idaho and Wyo. south to Calif. and southwest. Tex.; Hawaii;
Mexico south to Chiapas including Baja California. Prey: Nysius caenosulus Stal
nymphs. Adventive in Hawaii.
Astata immigrans Williams, 1946. Hawaii. Ent. Soc., Proc. 12: 641. d, °.

Biology: Williams, 1946. Hawaii. Ent. Soc., Proc. 12: 642-648 (prey, nesting under lab.
conditions).
millsi Cockerell. Colo. (Canad. Zone of Long’s Peak).
Dryudella millsi Cockerell, 1914. Ent. News 25: 32. 9.
montana (Cresson). Wash. to Wyo. south to Calif. and Colo. Ecology: Nests in flat, bare sand,
makes 2-3 cells per nest. Parasite: Senotainia trilineata (Wulp). Prey: Zelus or
Pselliopus sp.; Eurygaster alternatus Say.; Corimelaena montana Van D.; Alydus sp.;
Aufeius impressicollis Stal; Euschistus conspersus Uhl. All observed prey were
nymphs.
Astata montana Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proe. p. v. @.
Diploplectron florissantensis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 122. 2.

Biology: Evans, 1963. Ent. News 74: 236 (nest, prey). —Parker, 1969. Ent. Soc. Amer., Ann.
62: 965 (prey). —Aleock, 1973. Wasmann Jour. Biol. 31: 324-326, figs. 2, 3, 7, 8 (nest, prey
transport, parasite).

pernix Parker. South. Calif. to southwest. Tex.; Mexico (Hidalgo).
Dryudella pernix Parker, 1969. Ent. Soc. Amer., Ann. 62: 972, figs. 18, 22, 25, 40, 54, 66. 6,
oF
picta (Kohl). N. W. T., Oreg., Calif., Idaho, Nev., Utah, Colo., Ariz.; Mexico south to Morelos.
Prey: Liorhyssus hyalinus (F.) adult; Lygus sp. nymphs in laboratory.
Astatus pictus Kohl, 1888. Zool.-Bot. Gesell. Wien, Verhandl. 38: 146. ¢.
Astata Kohli Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 68, pl. 5, figs. 2, 2a. 9.
Astatus asper Fox, 1894 (1898). Acad. Nat. Sci. Phila., Proc. 45: 546. 3, 9.
Astata aspera Dalla Torre, 1897. Cat. Hym., v. 8, p. 651. Emend.
Astata asperiformis Rohwer, 1909. Ent. News 20: 371. 6.

Biology: Parker, 1969. Ent. Soc. Amer., Ann. 62: 965 (prey). —Steiner, 1973. Quaestiones Ent.
9: 23 (mating flight).
pinguis (Dahlbom). Alaska, N. W. T., Colo.; northern Europe and Asia. Prey: Trapezonotus
arenarius (L.), Drymus sylvaticus (F.), Nysius thymi (Wolff) ? in Europe.
Larra pinguis Dahlbom, 1832. Exercit. Hym., fase. 4, p. 50. 2.
Astata jaculator Smith, 1845. Zoologist 3: 1157. 2.

Biology: Verhoeff, 1951. Zool. Meded. 31: 162 (prey in Europe).

~ rhimpa Parker. B. C. to Calif., and Idaho and Wyo. to N. Mex.; Mexico south to Queretaro.
Ecology: Nests in compact soil containing gravel. Prey: Thyanta punctiventris Van D.
nymphs and adult; Lygaeus sp. nymph; Leptocoris trivittatus (Say) nymph.

Dryudella rhimpa Parker, 1969. Ent. Soc. Amer., Ann. 62: 970, figs. 5, 8, 17, 30, 44, 55, 62.
On or

Biology: Parker, 1969. Ent. Soc. Amer., Ann. 62: 965 (nest, prey).

1617
Family LARRIDAE

The contributions cited below treat only the subfamilies Larrinae and Miscophinae because
the Trypoxyloninae have been associated with the former two groups only since 1964. The revi-
sions are quite inadequate for identification in most genera for many additional species have
been described in intervening years.

Revision: Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc., 45: 467-551 (U. S. spp.). —Williams,
1914 (1913). Kans. Univ. Sci. Bul. 8: 121-213, pls. 22-30 (Kans. spp.).

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 113-126, figs. 9-45, 58, 59 (larvae).
—Evans, 1959. Amer. Ent. Soc., Trans. 85: 166-167, figs. 54-59 (larvae).

SUBFAMILY LARRINAE

All of the American genera are ground-nesting but one extralimital genus is known to nest in
twigs. The North American species are reported to prey upon various kinds of Orthoptera. How-
ever, no prey preference has been reported for our single species of Prosopigastra; in Europe
some species of the genus prey upon Hemiptera and Homoptera. One extralimital genus preys
upon lepidopterous larvae.

TRIBE LARRINI
Genus LARRA Fabricius

Genus LARRA Subgenus LARRA Fabricius

Larra Fabricius, 1793. Ent. System., v. 2, p. 220.
Type-species: Larra ichneumoniformis Fabricius. Desig. by Latreille, 1810.
Larrana Rafinesque, 1815. Analyse Nature ou Tabl. Univers, Palmero, p. 124. Emend. or
n. name.
Lara Drapiez, 1819. Bruxelles Gen. Sci. Phys., Ann., v. 1, p. 54. Lapsus or emend.
Monomatium Shuckard, 1840. In Swainson and Shuckard, Cabinet Cyclopedia of Lardner,
v. 129, p. 181. No species.
Type-species: Larraxena princeps Smith. Desig. by Pate, 1935. First included
species.
Tachytes subg. Lyrops Dahlbom, 1843. Hym. Europaea, v. 1, p. 132. Preoce.
Type-species: Tachytes (Lyrops) pagana Dahlbom. Monotypic.
Larraxena Smith, 1851. Ann. and Mag. Nat. Hist. (2) 7: 30.
Type-species: Larraxena princeps Smith. Monotypic.
Larrada Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 337.
Type-species: Sphex anathema Rossi. Orig. desig.

Only the typical subgenus occurs in North America. Larra preys upon mole crickets and

several extralimital species have been introduced into other areas for control of these pests.

analis Fabricius. East. states north to N. Y., south to Fla., La., west to Tex., Nebr. Prey:
Gryllotalpa hexadactyla Perty.
Larra analis Fabricius, 1804. Systema Piezatorum, p. 220. 9.
Larrada canescens Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 292. °.
Larrada americana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 214. ¢.
Larra cressonti Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 482. N. name.

Taxonomy: Cushman, 1935. Ent. Soe. Wash., Proc. 37: 82-87 (larva).
Biology: Smith, 1935. Ent. Soe. Wash., Proc. 37: 65-82.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 20, figs. N, P, Q (male
genitalia).

1618 Hymenoptera in America North of Mexico
Genus LIRIS Fabricius
Genus LIRIS Subgenus LIRIS Fabricius

Liris Fabricius, 1804. Systema Piezatortim, p. 227.
Type-species: Sphex aurata Fabricius. Desig. by Patton, 1881.
Lirisis Rafinesque, 1815. Analyse Nature ou Tabl. Univers, Palermo, p. 124. Emend. or n.
name.

The typical subgenus does not occur in the New World.
Genus LIRIS Subgenus LEPTOLARRA Cameron

Notogonia Costa, 1867. Mus. Zool. Univ. Napoli, Ann. 4: 83. Preoce.
Type-species: Tachytes nigra Fabricius of Vander Linden. Monotypic.
Caenolarra Cameron, 1900. Ann and Mag. Nat. Hist. (7) 5: 28.
Type-species: Caenolarra appendiculata Cameron. Monotypic.
Leptolarra Cameron, 1900. Ann. and Mag. Nat. Hist. (7) 5: 29.
Type-species: Leptolarra reticulata Cameron. Desig. by Richards, 1935.
Spanolarra Cameron, 1900. Ann. and Mag. Nat. Hist. (7) 5: 32.
Type-species: Spanolarra rufitarsis Cameron. Monotypic.
Notogonius Howard(!), 1901. Insect Book, pl. 6, fig. 1. Lapsus.
Chrysolarra Cameron, 1901. Ann. and Mag. Nat. Hist. (7) 8: 118.
Type-species: Chrysolarra appendiculata Cameron. Desig. by Pate, 1937.
Notogonidea Rohwer, 1911. Ent. Soc. Wash., Proc. 13: 234. N. name.
Dociliris Tsuneki, 1967. Etizenia 20: 26.
Type-species: Larrada subtessellata Smith. Orig. desig.
Nigliris Tsuneki, 1967. Etizenia 20: 27.
Type-species: Notogonia japonica Kohl. Orig. desig.
Liris subg. Colloliris Tsuneki, 1974. Polskie Pismo Ent. 44: 612.
Type-species: Notogonidea negrosensis Williams. Orig. desig.

The species of Leptolarra nest in soil, making one or several cells per nest at the end of a
rather short burrow. The usual prey are crickets, although one extralimital species has been re-
ported to use Gryllacrididae. Both nymphal and adult stages are used as prey, but in North
America the nymphs are used more frequently.

Taxonomy: Krombein, 1954. Amer. Ent. Soc., Trans. 80: 15-17 (key to U.S. spp.).
apicipennis (Cameron). South. Tex. to Panama.

Notogonia apicipennis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 58, pl. 4, figs. 16,
16anoeG
argentata (Beauvois). South. Ont. west to Nev., south to Fla., Ariz. and south. Calif.; south in
Mexico to Chiapas, Venezuela ?; Cuba, Bahamas; adventive in Hawaii. Ecology: Nests in
sand or compacted soil, makes 1-3 cells per nest and provides 1-6 prey per cell. Prey:
Gryllus pennsylvanicus Burm., G. rubens (Seudd.)?, G. sp., Miogryllus verticalis (Serv.),
Acheta assimilis F., Nemobius fasciatus DeG., N. sp., Orocharis saltator Uhl.; most
prey records are of nymphs, but adults are used occasionally.
Larra Pensylvanica Beauvois, 1811. Ins. Afr., Amer., p. 118.
Larra argentata Beauvois, 1811. Ins. Afr., Amer., p. 119. 9.
Tachytes murina Dahlbom, 1843. Hym. Europaea, v. 1, p. 182. ?. N. syn.
Larrada Pennsylvanica Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 292. Emend.
Larra nuda Taschenberg, 1870. Ztschr. Gesam. Naturw. Halle 34: 5. °.

Taxonomy: Krombein, 1976. Ent. Soc. Wash., Proe. 78: 333 (syn. of murina).

Biology: Ashmead, 1894. Psyche 7: 63 (nest, prey). —Williams, 1914 (1913). Kans. Univ. Sci.
Bul. 8: 189-192, fig. 118 (nest, prey hunting and transport). —Rau and Rau, 1918. Wasp
studies afield, pp. 152-158, fig. 35 (nest, prey). —Rau, 1922. Acad. Sci. St. Louis. Trans. 24
(7): 26 (prey, nest). —Reinhard, 1929. Witchery of wasps, pp. 67-71 (nest, prey).
—Krombein and Evans, 1955. Ent. Soe. Wash., Proc. 57: 233 (prey). —Krombein, 1958. Ent.
Soc. Wash., Proc. 60: 103 (prey). —Kurezewski and Kurczewski, 1971. Kans. Ent. Soc.,
Jour. 44: 134 (prey).

Superfamily SPHECOIDEA 1619

argenticauda (Cameron). South. Tex. to Costa Rica.
Notogonia chrysura Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 57. pl. 4, fig. 14. d.
Notogonia argenticauda Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 58, pl. 4, fig. 15.
3.

beata (Cameron). Transcont. in U. S. mostly in Austr. Zone south to Panama. Prey: Acheta
assimilis F. nymph. ,
Notogonia violaceipennis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 51, pl. 4, fig. 12.

9. N. syn.
Notogonia montezuma Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 52. 9. N. syn.

Notogonia truncata Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 54. 2. Preoce. N. syn.

Notogonia beata Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 56, pl. 4, fig. 18. d.

Notogonia nigripennis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 485. 2. Preoce. N.
syn.

Notogonia aequalis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 486. 2. N. syn.

Larra nigripennata Dalla Torre, 1897. Cat. Hym., v. 8, p. 670. N. name for nigripennis
Fox. N. syn.

Larra truncatula Dalla Torre, 1897. Cat. Hym., v. 8, p. 675. N. name for truncata
Cameron. N. syn.

Notogonia nigripennis var. occidentalis Viereck, 1903 (1902). Acad. Nat. Sci. Phila., Proc.
54: 731. 2. N. syn.

Notogonia subaequalis Rohwer, 1909. Ent. News. 20: 370. 2. N. syn.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 188 (prey).
fuliginosa muspa (Pate). South. Fla. Typical fuliginosa (Dahlbom) occurs in Cuba and Puerto

Rico.
Motes muspa Pate, 1943. Canad. Ent. 75: 201. d.

Taxonomy: Krombein, 1976. Ent. Soc. Wash., Proc. 78: 333-334 (lectotype of fwliginosa).
luctuosa dahlbomi (Cresson). South. Fla.; Cuba, Puerto Rico, Virgin Islands, Dominica.
Typical luctwosa (Smith) occurs in Santo Domingo, Mexico and Brazil.
Larrada Dahlbomi Cresson, 1865. Ent. Soc. Phila., Proc. 4: 138. 2. A provisional name to
be used if specimens identified as fuliginosa (Dahlbom) by Cresson were misidentified;
they were.

Taxonomy: Krombein, 1976. Ent. Soc. Wash., Proc. 78: 334-335 (lectotype of dahlbomi).

mescalero (Pate). South. Tex. to south. Ariz., south to Honduras.

Motes mescalero Pate, 1943. Canad. Ent. 75: 200. d.
panamensis muesebecki (Krombein), n. status. South. Fla.

Motes muesebecki Krombein, 1954. Amer. Ent. Soc., Trans. 80: 15. d, °.
panamensis panamensis (Cameron), n. status. South. Tex. to Panama.

Notogonia panamensis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 59, pl. 4, fig. 17. ¢.

Notogonidea sternalis Rohwer, 1914. U.S. Natl. Mus., Proc. 47: 519. ¢. N. syn.

UNPLACED TAXON OF LARRINI

Tachytes dives Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 247. “2” = 6. Carolina. Bohart
and Menke (1975) treat this as a questionable species of Liris. Presumably the locality
is in error, for no species of Larrini in America north of Mexico has the coloration of
vestiture, antenna and legs ascribed to dives.

TRIBE TACHYTINI

Genus LARROPSIS Patton

Larropsis Patton, 1892. Ent. News 3: 90.
Type-species: Larrada tenuicornis Smith. Orig. desig.

So far as known this genus is restricted to the Nearctic Region. The scanty biological records
indicate that the species are ground-nesting and that they prey upon cave and camel crickets
(Gryllacrididae).

1620 Hymenoptera in America North of Mexico

Revision: Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 653-685, 23 figs.

arizonensis Bohart and Bohart. Ariz., N. Mex., Utah.
Larropsis arizonensis Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 660, fig. 5. 2,
é.

atra Williams. Nebr., Kans., west. Tex., N. Mex., Ariz.
Larropsis ater Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 146. ¢.

chilopsidis (Cockerell). Kans., Colo., west. Tex., N. Mex.; Mexico (Chihuahua).
Ancistromma chilopsidis Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila.,
Proc. 49: 137. 2.
Ancistromma zerbeii Viereck, 1906. Amer. Ent. Soc., Trans. 32: 208. 2.
Ancistromma tachysphecoides Viereck, 1906. Amer. Ent. Soc., Trans. 32: 209. ¢.

conferta (Fox). Mont., Nebr., Iowa, Kans., Colo., Tex.
Ancistromma conferta Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 494. @.
Ancistromma paenerugosa Viereck, 1906. Amer. Ent. Soc., Trans. 32: 210. ¢.
Ancistromma bruneri Smith, 1906. Ent. News 17: 248. do.
Larropsis minor Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 146. 3.
Larropsis gracilis Rohwer, 1915. U.S. Natl. Mus., Proc. 49: 244. 2, 6.

consimilis (Fox). Kans., Okla., Tex., N. Mex.
Ancistromma consimilis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 496. °.
Ancistromma vegetoides Viereck, 1906. Amer. Ent. Soc., Trans. 32: 208. 9.
deserta Bohart and Bohart. South. Calif. deserts.
Larropsis deserta Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 665, figs. 4, 23. @,
Vo
discreta (Fox). Ga., Fla.
Ancistromma discreta Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 493. °.
divisa (Patton). Mont., Wyo., Nebr., Kans., north. Tex., N. Mex. Ecology: Nests in burrow off

crack in hard-packed soil. Prey: Ceuthophilus sp. nymphs.
Larra divisa Patton, 1879. U. 8. Geol. Geog. Survey, Bul. 5: 368. 2.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 192-193 (nest, prey).

elegans Bohart and Bohart. South. Tex., southeast N. Mex.
Larropsis elegans Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 667, figs. 1, 16. 2,
3.

filicornis Rohwer. West. Nebr., south. Okla., southeast. Tex. Prey: Ammobaenetes sp. nymph.
Larropsis filicornis Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 582. 9.
Larropsis yatesi Mickel, 1918 (1917). Nebr. Univ., Studies 17: 412. ¢.

Biology: Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 655 (prey).

greenei Rohwer. N. J., S. C., Fla., Kans.
Larropsis greenei Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 178. 9.

interocularis Bohart and Bohart. Kans., Ariz.
Larropsis interocularis Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 670, figs. 4, 8.
ee
lucida Bohart and Bohart. Ariz. (Mohave Co.).
Larropsis lucida Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 671, fig. 7. @.
rugosa (Fox). Mont., N. Dak. and Iowa south to Kans. and Ariz.; Mexico (Chihuahua, Durango,
Zacatecas).
Ancistromma rugosa Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 496. ¢.
sericea Bohart and Bohart. Nebr., Kans., Tex.
Larropsis sericea Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 672, fig. 6. 2, 3.
snowi Bohart and Bohart. Ariz. in U. and L. Sonoran Zones.
Larropsis snowi Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 673, figs. 6, 18. 2, d.

Superfamily SPHECOIDEA 1621

sparsa Bohart and Bohart. West. Tex., N. Mex., Ariz.; Mexico (Coahuila, Durango).
Larropsis sparsa Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 675, fig. 6. d.

striata Bohart and Bohart. Calif. (San Bernardino Co.).
Larropsis striata Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 676, fig. 5. d.

tenuicornis (Smith). Wash., Idaho, Oreg., Nev., Calif.; Mexico (Baja California).
Larrada tenuicornis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 293. °.

testacea Bohart and Bohart. Kans. (Pottawatomie Co.).
Larropsis testacea Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 677, figs. 5, 14. .

texensis Bohart and Bohart. Cent. and south. Tex.
Larropsis texensis Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 678, fig. 7. 3.

uniformis Bohart and Bohart. Idaho, Nev. to Colo. south to Calif. and N. Mex.; Mexico
(Chihuahua, Coahuila, Durango).
Larropsis uniformis Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 678, figs. 2, 12,
om CenGr
vegeta (Fox). Wyo., Nebr., Colo., west. Tex., north. Ariz.
Ancistromma vegeta Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 497. 2, d.

washoensis Bohart and Bohart. Nev. (Washoe Co.).
Larropsis washoensis Bohart and Bohart, 1966. Amer. Ent. Soc., Trans. 92: 681, figs. 3, 10,
belie onor

Genus ANCISTROMMA Fox

Ancistromma Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 487.
Type-species: Larrada distincta Smith. Desig. by Rohwer, 1911.

These ground-nesting wasps apparently construct multicelled nests at the bottom of
pre-existing burrows or cavities. The recorded prey includes both typical crickets (Gryllidae)
and cave crickets (Gryllacrididae); both nymphs and adults are used.

Revision: Bohart and Bohart, 1962. Ent. Soc. Wash., Proc. 64: 21-37, 32 figs. (N. Amer. spp.).

aurantia (Fox). Alta. east to N. Dak., south to Ariz. and Kans. Prey: Ceuthophilus fusiformis
Seudd.
Larra aurantia Fox, 1891. Ent. News 2: 194. °.
Ancistromma aurulenta(!) Fox, 1894 (1898). Acad. Nat. Sci. Phila., Proc. 45: 388.

Biology: Bohart and Bohart, 1962. Ent. Soc. Wash., Proc. 64: 25 (prey).

bradleyi (Bohart and Bohart). Oreg., Calif.
Larropsis (Ancistromma) bradleyi Bohart and Bohart, 1962. Ent. Soc. Wash., Proc. 64: 25,
figs. 20, 21, 31. 2, d.

capax (Fox). B. C., Alta., east to N. Dak., south to Calif, Kans. and N. Mex. Prey:
Ceuthophilus sp., nymph. Predator: Philanthus zebratus Cr.
Ancistromma capax Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 490. °.
Ancistromma dolosa Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 492. °.
Larropsis dolosana Rohwer, 1915. U. S. Natl. Mus., Proc. 49: 245. 9.
Larropsis picina Mickel, 1916. Amer. Ent. Soc., Trans. 42: 418. 2, d.
Biology: Bohart and Bohart, 1962. Ent. Soc. Wash., Proc. 64: 30 (prey).
corrugata (Bohart and Bohart). B. C., Oreg., Calif.
Larropsis (Ancistromma) corrugata Bohart and Bohart, 1962. Ent. Soe. Wash., Proc. 64:
31, figs. 3, 18, 19, 27. 2, ¢.
distincta (Smith). Transcont., B. C. to Maine, south to Calif., Kans., N. C. Ecology: Nests in
natural cavities in coarse soil, making up to 9 cells per nest, each cell provided with 1-3
adult crickets. Parasite: Metopia argyrocephala (Meig.), Senotainia trilineata (Wulp).
Prey: Nemobius fasciatus (DeG.) adults.

Larrada distincta Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 292. 2.
Larropsis distincta var. semirufa Banks, 1921. Ent. Soc. Amer., Ann. 14: 19. °, 3.

1622 Hymenoptera in America North of Mexico

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 119-120, figs. 29-38 (larva).

Biology: Evans, 1958. Ent. News 69: 197-200 (nest, prey, parasites).

granulosa (Bohart and Bohart). Calif., Nev., Idaho, Ariz., N. Mex.
Larropsis (Ancistromma) granulosa Bohart and Bohart, 1962. Ent. Soe. Wash., Proc. 64:
33, figs. 4, 6, 7, 28. 2, do.
hurdi (Bohart and Bohart). Calif.
Larropsis (Ancistromma) hurdi Bohart and Bohart, 1962. Ent. Soc. Wash., Proce. 64: 34,
figs. 5, 8, 9, 30. 2, 6.
platynota (Bohart and Bohart). Ariz. (Tucson).
Larropsis (Ancistromma) platynota Bohart and Bohart, 1962. Ent. Soc. Wash., Proc. 64:
35, figs. 24, 32. 2.

portiana (Rohwer). Western Tex., N. Mex.
Larropsis portianus Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 583. 2, 6.

sericifrons Smith. Nebr., Wyo., Tex., N. Mex., Ariz.
Ancistromma sericifrons Smith, 1906. Ent. News 17: 247. 6.
Larropsis rubens Mickel, 1918 (1917). Nebr. Univ., Studies 17: 329. 9.

shappirioi (Bohart and Bohart). D. C., Va.
Larropsis (Ancistromma) shappirioi Bohart and Bohart, 1962. Ent. Soc. Wash., Proc. 64:
37, figs. 14, 15, 28. 2, ¢.

Genus TACHYTES Panzer

Tachytes Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 129.
Type-species: Sphex tricolor Fabricius. Monotypic.

Lyrops Illiger, 1807. Fauna Etrusca, v. 2, p. 162.
Type-species: Andrena etrusca Rossi. Monotypic.

Tachyptera Dahlbom, 1843. Hym. Europaea, v. 1, p. 133. Preoce.
Type-species: Apis obsoleta Rossi. Desig. by Patton, 1880.

Tachytes subg. Holotachytes Turner, 1917. Ann. and Mag. Nat. Hist. (8) 20: 10.
Type-species: Tachytes dichroa Smith. Orig. desig.

Tachytes subg. Calotachytes Turner, 1917. Ann. and Mag. Nat. Hist. (8) 20: 10.
Type-species: Tachytes marshalli Turner. Orig. desig.

Tachyoides Banks, 1942. Mus. Compar. Zool., Bul. 89: 397.
Type-species: Tachytes mergus Fox. Orig. desig.

Tachytes subg. Tachyplena Banks, 1942. Mus. Compar. Zool., Bul. 89: 397. N. name.

Tachytes subg. Tachynana Banks, 1942. Mus. Compar. Zool., Bul. 89: 398.
Type-species: Tachytes obscurus Cresson. Orig. desig.

Members of this genus are ground-nesting and usually construct multicellular nests. Some
species begin their burrows in pre-existing holes such as rodent burrows or abandoned burrows
of large insects, whereas others excavate their burrows from the ground surface. Members of
the Aurulentus Species Group prey on katydids (Tettigoniidae), those of the Pepticus and
Distinctus Groups use grasshoppers (Acrididae), species of the Abdominalis Group prey upon
grasshoppers, grouse locusts (Tetrigidae) and pygmy mole crickets (Tridactylidae), and the Mer-
gus Group uses only pygmy mole crickets. Members of the extralimital Ambidens Group have
been observed preying upon geometrid caterpillars.

Revision: Fox, 1892. Amer. Ent. Soc., Trans. 19: 234-252 (N. Amer. spp.). —Banks, 1942. Mus.
Compar. Zool., Bul. 89: 395-4386 (N. Amer. spp.).

Biology: Evans and Kurezewski, 1966. Kans. Ent. Soc., Jour. 39: 323-332, 3 figs. (comparative
behavior of a few N. Amer. spp.).

SPECIES GROUP AURULENTUS

This species group is the equivalent of Tachytes sens. str.

auricomans Bradley. Ga.
Tachytes auricomans Bradley, 1919. Ent. News 30: 298. 3.

Superfamily SPHECOIDEA 1628

aurulentus (Fabricius). East. Canada and States south to Fla., north to N. J., west to Tex.,

Kans. Ecology: Nests in sand or clay with 2-4 cells per nest and 6-11 prey per cell. Prey:
Conocephalus saltans (Scudd.), C. spp., Orchelimum fidicinium Rehn and Heb., O. spp.,
Tettigoniidae sp.; all nymphs.

Larra aurulenta Fabricius, 1804. Systema Piezatorum, p. 220. °.

Tachytes mandibularis Patton, 1881. Boston Soe. Nat. Hist., Proc. 20: 394. 2, 6.

Tachytes propinquus Rohwer, 1909. Ent. News 20: 198. 2.

Tachytes duplicatus Rohwer, 1920. Ent. Soc. Wash., Proc. 22: 59. 2.

Taxonomy: Strandtmann, 1945. Ent. Soc. Amer., Ann. 38: 307, figs. 1-2 (larva). —Evans, 1958.
Amer. Ent. Soc., Trans. 84: 117, figs. 18-20 (larva). —van der Vecht, 1961. Zool. Verhandl.
Rijksmus. Natuurlijke Hist. Leiden, No. 48, pp. 11-12 (synonymy).

Biology: Patton, 1892. Ent. News 3: 90 (prey). —Williams, 1914 (1913). Kans. Univ. Sci. Bul.
8: 198 (prey hunting). —Dow, 1930. Psyche 37: 182 (prey transport). —Strandtmann, 1945.
Ent. Soc. Amer., Ann. 38: 305-308, figs. 1-4 (nest, prey, cocoon, life cycle). —Evans and
Kurezewski, 1966. Kans. Ent. Soc., Jour. 329-330 (nest, prey).

badius Banks. Tex. (Comal Co.), Ariz. (Baboquivari Mts.).
Tachytes (Tachyplena) badius Banks, 1942. Mus. Compar. Zool., Bul. 89: 417. 9.

columbiae Fox. N. J., Md., Va., N. C.
Tachytes columbiae Fox, 1892. Amer. Ent. Soce., Trans. 19: 241. 2, 3.

crassus Patton. Conn., Mass., N. Y., Md., Wis., Iowa, Mo., Nebr. Ecology: Nests in sand or in
heavy clay-loam, constructs as many as 9 cells per nest, and stores 5-10 prey per cell.
Prey: Orchelimum gladiator Brun., O. spp., Conocephalus f. fasciatus (DeG.), C.
nigropleurus (Brun.), C. attenuatus (Scudd.); most prey are nymphs but adults are
occasionally stored.
Tachytes crassus Patton, 1881. Boston Soc. Nat. Hist., Proc. 20: 241. 2, 6.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 285-286, figs. 96-98 (larva).

Biology: Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 64 (nest, prey transport). —Evans and
Kurczewski, 1966. Kans. Ent. Soc., Jour. 39: 324-326, figs. 1-2 (nest, prey transport).

ermineus Banks. West. Tex., Ariz., south. Utah and Nev.; Mexico (Baja California).
Tachytes (Tachyplena) ermineus Banks, 1942. Mus. Compar. Zool., Bul. 89: 413. 2, 3.

exornatus Fox. Southwest. Tex., N. Mex., Ariz.; Mexico (Baja California). Parasite: Zanysson
t. tecanus (Cr.) ? Prey: Conalcea sp.? nymph.
Tachytes exornatus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proe. 45: 501. d.

Biology: Cockerell, 1903. Entomologist 36: 100 (parasite ?). —Krombein, 1960. Brooklyn Ent.
Soc., Bul. 55: 75 (prey transport).

floridanus Rohwer. N. C. to Fla., west to Ariz.
Tachytes pepticus floridanus Rohwer, 1920. Ent. Soc. Wash., Proc. 22: 59. 92.
Tachytes (Tachyplena) foxi Banks, 1942. Mus. Compar. Zool., Bul. 89: 416. °.
Tachytes (Tachyplena) comanche Banks, 1942. Mus. Compar. Zool., Bul. 89: 417. 3.

harpax Patton. N. H., Mass., Conn., Pa., Va., N. C., Wis., Mich., Iowa, Nebr. Ecology: Nests in
fine silt and sand, constructs several cells per nest and stores 2-3 prey per cell. Prey:
Conocephalus brevipennis (Seudd.).
Tachytes harpax Patton, 1881. Boston Soc. Nat. Hist., Proc. 20: 395. 2, ¢.
Tachytes dubitatus Rohwer, 1909. Ent. News 20: 202. 2, d.

Biology: Parker, 1921. Ent. Soc. Wash., Proc. 23: 103-104 (nest, prey transport).

praedator Fox. Md. to Fla., west to Iowa, Kans. and Tex. Ecology: Nests in fine-grained,
somewhat moist sand, apparently makes only a single cell stored with up to 5 prey.
Prey: Scudderia sp., Conocephalus f. fasciatus (DeG.), C. sp., Neoconocephalus sp.,
Eremopedes sp.; usually nymphs are stored, but occasionally adults.
Tachytes praedator Fox, 1892. Amer. Ent. Soe., Trans. 19: 240. 2, 6.

1624 Hymenoptera in America North of Mexico

Biology: Evans and Kurezewski, 1966. Kans. Ent. Soc., Jour. 39: 330-331 (nest, prey
transport). —Lin, 1967. Amer. Midland Nat. 77: 241-242, 1 fig. (nest, prey hunt and
transport).

sayi Banks. U.S. west of 100th meridian, Nebr. to Wash. south to Tex. and cent. Calif.
Ecology: Nests in sandy soil. Prey: Melanoplus lakinus (Seudd.) adults.
Tachytes (Tachytes) sayi Banks, 1942. Mus. Compar. Zool., Bul. 89: 421. 6, 9.
Tachytes (Tachytes) brevipilis Banks, 1942. Mus. Compar. Zool., Bul. 89: 422. @.
Tachytes (Tachytes) hesperus Banks, 1942. Mus. Compar. Zool., Bul. 89: 423. 2, ¢.

Biology: Evans and Kurczewski, 1966. Kans. Ent. Soc., Jour. 39: 331 (nest, prey).

validus Cresson. Ont. to Fla., west to Mich., Colo. and Tex. Ecology: Nests in sand, constructs

6 to more than 9 cells per nest and stores 1-5 prey per cell. Parasite: Senotainia
trilineata (Wulp), Miltogrammini sp.; Diptera sp. Prey: Conocephalus brevipennis
(Seudd.), C. nigropleurum (Brun.), C. f. fasciatus (DeG.), C. spp.; adults are preyed upon
more frequently than nymphs.

Tachytes validus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 216. 2, 3.

Tachytes breviventris Fox, 1892. Amer. Ent. Soc., Trans. 19: 239. 2, ¢.

Tachytes calcaratus Fox, 1892. Amer. Ent. Soc., Trans. 19: 239. 9, d.

Tachytes calcaratiformis Rohwer, 1909. Ent. News 20: 204. 2, 6.

Tachytes (Tachyplena) calcaratiformis var. coloradensis Banks, 1942. Mus. Compar. Zool.,
Bul. 89: 411. 2, ¢.

Tachytes (Tachyplena) belfragei Banks, 1942. Mus. Compar. Zool., Bul. 89: 411. 2.

Tachytes quadrifasciatus Dreisbach, 1948. Ent. News 59: 151. 6.

Biology: Parker, 1921. Ent. Soc. Wash., Proc. 23: 104-107 (nest, prey transport, cocoon, life
cycle, parasite). —Evans and Kurczewski, 1966. Kans. Ent. Soc., Jour. 39: 326-329 (nest,
prey transport, parasite). —Kurczewski and Ginsburg, 1971. Kans. Ent. Soc., Jour. 44:
113-131, 9 figs. (nest, prey transport, egg, parasites). —Kurezewski and Kurczewski, 1971.
Kans. Ent. Soc., Jour. 44: 132 (prey).

SPECIES GROUP PEPTICUS

Revision: Bohart, 1962. Pan-Pacific Ent. 38: 117-129, 26 figs.

californicus Bohart. Calif., Oreg., Idaho.
Tachytes californicus Bohart, 1962. Pan-Pacific Ent. 38: 125, figs. 15-17. d, @.

chelatus Bohart. Ariz., Utah.
Tachytes pepticus chelatus Bohart, 1962. Pan-Pacific Ent. 38: 120, figs. 5, 6, 7. 3d, @.

fulviventris fulviventris Cresson. N. Dak. south to Tex., west to Wyo., Utah, N. Mex.; Mexico
(Oaxaca). Ecology: Nests in deserted prairie-dog burrow. Parasite: Diptera sp. Prey:
Cordillacris crenulata (Brun.) adults.
Tachytes fulviventris Cresson, 1865. Ent. Soc. Phila., Proc. 4: 466. 9.
Tachytes caelebs Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 355. 6.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 197-198 (nest, prey, parasite).

fulviventris rossi Bohart. South. Calif.; Mexico (Baja California).

Tachytes fulviventris rossi Bohart, 1962. Pan-Pacific Ent. 38: 124, figs. 18-20. ¢.
nevadensis Bohart. Nev., Calif., Oreg., Wash.

Tachytes nevadensis Bohart, 1962. Pan-Pacific Ent. 38: 124, figs. 21-28. d, 9.
pennsylvanicus Banks. N. Y. to Va. west to Idaho, Colo., N. Mex., Oreg., B. C.

Tachytes pennsylvanicus Banks, 1921. Ent. Soc. Amer., Ann. 14: 18. 6.
pepticus (Say). N. C. to Fla. west to IIl., Kans., Nebr., Colo. and Tex. Prey: Melanoplus sp.

nymph.
Lyrops pepticus Say, 1837. boston Jour. Nat. Hist. 1: 371. 2, 6.
Tachytes sericatus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 216. 2 (male misdet.?).

Superfamily SPHECOIDEA 1625

Tachytes (Tachytes) fulviventris var. inferioris Banks, 1942. Mus. Compar. Zool., Bul. 89:
ADO oe
Tachytes (Tachytes) cressoni Banks, 1942. Mus. Compar. Zool., Bul. 89: 425. °.

Biology: Evans and Kurezewski, 1966. Kans. Ent. Soc., Jour. 39: 331 (prey).
sculleni Bohart. Western Tex. to southern Calif., Nev.
Tachytes sculleni Bohart, 1962. Pan-Pacifie Ent. 38: 126, figs. 8-11. ¢, 2.
spatulatus Fox. Colo., N. Mex., Ariz., Utah, Nev., Wyo., Calif.
Tachytes spatulatus Fox, 1892. Amer. Ent. Soc., Trans. 19: 243. 3.
Tachytes basirufus Rohwer, 1909. Ent. News 20: 197. 3.
Tachytes (Tachytes) utahensis Banks, 1942. Mus. Compar. Zool., Bul. 89: 424. 2, 6.

SPECIES GROUP DISTINCTUS

amazonus Smith. N. C. to Fla., west to Nebr., Oreg. and Calif.; south through Cent. and South

Amer. to Argentina. Prey: Melanoplus spp. adults and nymphs.

Tachytes Amazonum Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 303. 2, ¢.

Tachytes clypeatus Taschenberg, 1870. Ztschr. Gesell. Naturwiss. 36: 10. 2.

Tachytes scalaris Taschenberg, 1870. Ztschr. Gesell. Naturwiss. 36: 11. 3.

Tachytes rufofasciatus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 247. 6.

Tachytes fervens Smith, 1873. Ann. and Mag. Nat. Hist. (4) 12: 57. 6.

Tachytes dives Holmberg, 1884. Soc. Cient. Argentina, An. 18: 220. ¢. Preoce.

Tachysphex rufomaculatus Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 64, pl. 4, fig.
Bp

Tachytes Holmbergii Dalla Torre, 1897. Cat. Hym., v. 8, p. 690. N. name.

Tachytes nigricaudus Brethes, 1909. Mus. Nac. Buenos Aires, An. (8) 12: 241. d.

Tachytes Fiebrigi Brethes, 1909. Mus. Nac. Buenos Aires, An. (3) 12: 242. 6.

Tachytes anisitsi Strand, 1910. Zool. Jahrb., Abt. f. System., v. 29, h. 2, p. 164. 9.

Tachytes rufoannulatus Strand, 1910. Zool. Jahrb., Abt. f. System, v. 29, h. 2, p. 167. ¢.

Tachytes mimeticus Schrottky, 1909. Soc. Cient. Argentina, An. 68: 250.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Studies 8: 199-200 (nest, prey hunt and
transport).
chrysocercus Rohwer. South. Tex. and Ariz.; Mexico (Coahuila, Baja California). Parasite:
Zanysson plesia (Rohwer) ?
Tachytes chrysocercus Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 581. °.
Tachytes (Tachytes) elongatus var. apache Banks, 1942. Mus. Compar. Zool., Bul. 89: 419.
3.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, p. 90 (host ?).
distinctus distinctus Smith. Md. to Fla., west to B. C. and Calif.; Mexico south to South

America. Ecology: Nests in sandy soil and inside lizard holes, makes up to 20 cells per
nest and stores 1-4 mostly immature prey per cell; also nests in abandoned burrows of
the cicada killer, Sphecius speciosus (Dru.). Parasite: Diptera sp.; Zanysson texanus
(Cr.) ? Prey: Melanoplus femurrubrum (DeG.), M. spp., Ageneotettix deorum (Scudd.),
Orphulella sp. near speciosa (Secudd.), Schistocerca sp., Hesperotettix sp., Paraidemona
sp.; most prey is nymphal but occasionally adults are used. Another subsp. occurs in the
Bahamas Islands.

Larrada fulvipes Smith, 1856. Cat. Hym., Brit. Mus., v. 4, p. 288. @.

Tachytes distinctus Smith, 1856. Cat. Hym., Brit. Mus., v. 4, p. 307. °.

Tachytes elongatus Cresson, 1872. Amer. Ent Soc., Trans. 4: 215. d.

Tachytes Yucatanensis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 60, pl. 4, fig. 18. 9.

Tachytes contractus Fox, 1892. Amer. Ent. Soc., Trans. 19: 245. 2. Preoce.

Tachytes (Tachytes) elongatus var. seminole Banks, 1942. Mus. Compar. Zool., Bul. 89: 419.
3

Tachytes (Tachytes) austrinus Banks, 1942. Mus. Compar. Zool., Bul. 89: 419. N. name.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 116-117, figs. 9-17 (larva). —Evans, 1964.
Amer. Ent. Soc., Trans. 90: 286-287 (larva).

1626 Hymenoptera in America North of Mexico

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 194-197, figs. 113-116 (nest, prey hunt
and transport). —Rau, 1934. Canad. Ent. 66: 260 (nest). —Evans, 1964. Amer. Ent. Soc.,
Trans. 90: 287 (nest, prey). —Lin, 1965. Brooklyn Ent. Soc., Bul. 59 and 60: 82-84, 1 fig.
(nest). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 89-90 (host ?).
—Lin, 1972. In Michener and Lin, Quart. Rev. Biol. 47: 140-141 (nest guarding by males,
parasite).

guatemalensis Cameron. East. U. S. to Guatemala.
Tachytes Guatemalensis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 60, pl. 4, fig. 19.

OF
Liris coxalis Patton, 1892. Ent. News 3: 90. 9, 6.
SPECIES GROUP ABDOMINALIS
This is the equivalent of the subgenus Tachynana Banks.

abdominalis (Say). Kans., Tex. to Ariz., Utah; Mexico. Prey: Melanoplus sp.; Tetrigidae sp.; all

prey were nymphs.
Larra abdominalis Say, 1823. Western Quart. Rptr. 2: 77. 2.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 193 (nest, prey).
birkmanni Rohwer. Tex., Ariz.
Tachytes birkmanni Rohwer, 1909. Ent. News 20: 199. °.
Tachytes (Tachynana) atomus Banks, 1942. Mus. Compar. Zool., Bul. 89: 433. 2.
chrysopyga obscurus Cresson. Md., Nebr. and Wash. south to Fla. and Ariz.; north. Mexico.
Prey: Acrididae sp. Typical chrysopyga (Spinola) and other subspp. occur in Mexico,
West Indies, Central and South America.
Tachytes obscurus Cresson, 1872. Amer. Ent. Soce., Trans. 4: 217. @.
Tachytes texcanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 217. ¢.
Tachytes (Tachynana) hirsutifrons Banks, 1942. Mus. Compar. Zool., Bul. 89: 430. 3.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 199 (prey).
intermedius (Viereck). N. Y. to Fla., west to Nebr. and Tex. Ecology: Nests in sand and may

make only one cell per nest with several prey per cell. Prey: Tridactylus apicalis Say,
T. minutus Seudd.; both nymphs and adults.

Tachysphex intermedius Viereck, 1906. Amer. Ent. Soc., Trans. 32: 211. ¢.

Tachytes minutus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 128. ¢.

Tachytes maestus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 417. @.

Tachytes austerus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 417. °.

Tachytes (Tachynana) amiculus Banks, 1942. Mus. Compar. Zool., Bul. 89: 432. ¢.

Biology: Krombein and Kurczewski, 1963. Biol. Soc. Wash., Proc. 76: 146 (prey). —Krombein,
1963. Biol. Soc. Wash., Proc. 76: 273 (prey hunt). —Kurczewski, 1966. Kans. Ent. Soc., Jour.
39: 147-154, figs. 2, 5 (nest, prey hunt and transport). —Kurezewski and Kurczewski, 1971.
Kans. Ent. Soc., Jour. 44: 132 (prey).
obductus Fox. N. Y. to Fla. west to Nebr. and Tex. Ecology: Nests in sand, constructs up to 6
cells per nest, and stores 3-7 prey per cell. Prey: Tetrix o. ornata (Say), Tetrigidae sp.;
nymphs and adults.
Tachytes obductus Fox, 1892. Amer. Ent. Soc., Trans. 19: 250. °.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Studies 8: 198 (prey hunting and transport).
—Kurezewski and Kurczewski, 1971. Kans. Ent. Soc., Jour. 44: 133-134, 2 figs. (nest, prey).

parvus Fox. N. J. to Fla. west to Idaho and Calif. Prey: Neotettix femoratus (Scudd.) nymph.
Tachytes parvus Fox, 1892. Amer. Ent. Soc., Trans. 19: 249. ¢.
Tachytes (Tachynana) pattoni Banks, 1942. Mus. Compar. Zool., Bul. 89: 428. 9.
Tachytes (Tachynana) arizonicus Banks, 1942. Mus. Compar. Zool., Bul. 89: 429. 9.

Biology: Kurezewski and Kurczewski, 1971. Kans. Ent. Soc., Jour. 44: 134 (prey transport).
SPECIES GRoUP MERGUS

This is the equivalent of Tachyoides Banks.

Superfamily SPHECOIDEA 1627

mergus Fox. N. J. to Fla. west to Nebr. and Ariz.; Mexico. Ecology: Nests in pure sand with

high water table, constructs 1-3 cells per nest, and stores 6-13 small prey per cell.
Parasite: Phrosinella fulvicornis (Coq.). Prey: Tridactylus apicalis Say, T. minutus
Scudd.; mostly nymphs but a few adults may be used.

Tachytes mergus Fox, 1892. Amer. Ent. Soc., Trans. 19: 250. 2.

Tachytes minor Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 127. 3.

Tachytes obscuranus Rohwer, 1909. Ent. News 20: 205. 2, (¢ misdet.).

Tachyoides ariella Banks, 1942. Mus. Compar. Zool., Bul. 89: 434. 2.

Taxonomy: Krombein, 1948. Brooklyn Ent. Soc., Bul. 43: 18-20. 6. —Evans, 1964. Amer. Ent.
Soc., Trans. 90: 287-288, figs. 85-87 (larva).

Biology: Krombein and Kurczewski, 1963. Biol. Soc. Wash., Proc. 76: 143-150, figs. 1-3 (nest,
prey hunting and transport, egg, life cycle, parasites). —Kurczewski, 1966. Kans. Ent. Soc.,
Jour. 39: 147-154, figs. 1, 3, 4 (nest, prey hunt and transport).

Genus TACHYSPHEX Kohl

Tachysphex Kohl, 1883. Deut. Ent. Ztschr. 27: 166.

Type-species: Tachysphex filicornis Kohl. Desig. by Bingham, 1897.
Schistosphex Arnold, 1922. Transvaal Mus., Ann. 9: 137.

Type-species: Schistosphex Breijeri Arnold. Orig. desig.
Atelosphex Arnold, 1923. Transvaal Mus., Ann. 9: 177.

Type-species: Atelosphex miscophoides Arnold. Orig. desig.

Members of this genus are ground-nesting and usually construct shallow, multicellular nests in
which one or several prey are stored per cell. All authenticated prey records are for various
kinds of Orthoptera including grasshoppers, mantids, cockroaches, crickets and katydids. In
America north of Mexico members of the Pompiliformis and Terminatus Groups prey mostly
upon Acrididae and occasionally use Tettigoniidae, one member of the Undatus Group uses
Acrididae, two members of the Brullii Group use Blattidae or Tettigoniidae respectively, and
one member of the Julliani Group uses Mantidae.

Revision: Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 504-533 (N. Amer. spp.).
Taxonomy: Kurczewski, 1971. Ent. Soc. Wash., Proc. 73: 113-114 (key to Fla. spp.).

Biology: Kurczewski, 1966. Kans. Ent. Soc., Jour. 39: 436-453, 9 figs. (comparative male
behavior).

SPECIES GROUP POMPILIFORMIS

acutus (Patton). East. States north to N. B., Ont., Man., west to S. Dak., Kans., south to N. C.
Larra acuta Patton, 1881. Boston Soc. Nat. Hist., Proc. 20: 390. °.
Tachysphex bruesi Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 577. 9.
aethiops (Cresson). West. States north to B. C., N. W. T. and Sask., south to Calif. and Colo.
Ecology: Constructs nest in sand off side of burrow of Bembiz, up to 3 cells per nest,
provides 1 prey per nest. Parasite: Diptera sp. Prey: Trimerotropis suffusa Scudd.?, T.
sp., Acrididae sp. Predator: Philanthus pulcher D. T., P. zebratus nitens (Bks.).
Larrada aethiops Cresson, 1865. Ent. Soc. Phila., Proc. 4: 465. 2.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 489-490 (nest, prey, parasite). —Evans,
1973. Great Basin Nat. 33: 149-150 (nest, prey). —Aleock, 1973. Wasmann Jour. Biol. 31:
329, fig. 5 (nest, prey).

amplus Fox. West. States north to Oreg. and Wyo., south to Calif. and Tex.; Mexico
(Zacatecas).
Tachysphex amplus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 522. 2, ¢.
Tachysphex gillettei Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 571. @.
Tachysphex neomexicanus Rohwer, 1911. U. S. Natl. Mus., Proe. 40: 575. @.
angularis Mickel. Nebr. (Sioux Co.).
Tachysphex angularis Mickel, 1916. Amer. Ent. Soc., Trans. 42: 416. ¢.

1628 Hymenoptera in America North of Mexico

antennatus Fox. Transcont., N. H., Mont. and Oreg., south to Va., La., Calif.; Mexico
(Veracruz, Chiapas).
Tachysphex antennatus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 516. °.
Tachysphex sculptiloides Williams, 1914 (1918). Kans. Univ. Sci. Bul. 8: 166. 2°. N. syn. (W.
J. Pulawski).
Tachysphex nigrocaudatus Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 167. 2, 3d. N.
syn. (W. J. Pulawski).
crassiformis Viereck. N. C., Kans., Wyo., Calif. south to Fla., Tex., Ariz.; south through Mexico
and Central America to Colombia and Venezuela. Ecology: Makes 1-celled nest in flat,
loose sand, stores 1-3 prey per cell. Prey: Psinidia fenestralis (Serv.), Scirtetica
marmorata picta Seudd., Tryxalinae sp.; only nymphs have been reported as prey.
Tachysphex crassiformis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 210. @.
Tachysphex wheeler: Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 579. 2. N. syn. (W. J.
Pulawski).
Tachysphex plenoculiformis Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 167. 2. N. syn.
(W. J. Pulawski).
Tachysphex boharti Krombein, 1963. Ent. News 74: 177. 2, 36. N. syn. (W. J. Pulawski).
Tachysphex gibbus Pulawski, 1974. Polskie Pismo Ent. 44: 20, figs. 9-17. N. syn. (W. J.
Pulawski).

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 201-202 (prey). —Krombein, 1963. Ent.
News 74: 179-180 (nest, prey transport).
crenulatus Fox. N. Mex., Ariz., Calif.
Tachysphex crenulatus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 512. 2.
decorus Fox. N. Dak.
Tachysphex decorus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 524. °.
eldoradensis Rohwer. Calif., Oreg., Wyo., Canad. Zone. Ecology: Occurs in openings in forests.
Tachysphex eldoradensis Rohwer, 1917. U.S. Natl. Mus., Proe. 53: 245. @.
erythraeus Mickel. Nebr. (Sioux Co.).
Tachysphex erythraeus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 415. °.
glabrior Williams. Kans., Tex.; Mexico (Puebla, San Luis Potosi), E] Salvador, Costa Rica,
Venezuela.
Tachysphex glabrior Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 170. 2, d.
hurdi Bohart. Oreg., Calif.; Mexico (Baja California).
Tachysphex hurdi Bohart, 1962. Biol. Soc. Wash., Proc. 75: 33, figs. 18-15. 3, 9.
krombeini Kurczewski. Fla., Ga. Ecology: Makes 1-celled nest in flat sand and stores up to 7
prey. Prey: Melanoplus sp. nymphs; Odontoxiphidium apterum Morse nymph.
Tachysphex krombeini Kurezewski, 1971. Ent. Soc. Wash., Proc. 73: 111, 1 fig. d, 9.

Biology: Kurezewski, 1971. Ent. Soc. Wash., Proc. 73: 115-116 (nest, prey hunt and transport,
egg).
laevifrons (Smith). N. C., Fla., Kans., Tex. Ecology: Makes 1-celled nest in flat vegetated sand,
stores 1 or a few larger prey. Prey: Melanoplus sp. nymphs.
Larrada laevifrons Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 291. 9.
Tachysphex leensis Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 578. 2. N. syn. (W. J.
Pulawski).
Tachysphex consimiloides Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 164. 2. N. syn.
(W. J. Pulawski).

Biology: Kurezewski, 1966. Kans. Ent. Soc., Jour. 39: 439 (male behavior). —Krombein, 1967.
U.S. Dept. Agr., Monog. 2, Sup. 2, p. 393 (prey). —Kurcezewski, 1971. Ent. Soc. Wash.,
Proc. 73: 114-116 (nest, prey).

montanus (Cresson). Mont., Wyo., Colo., Utah, Nev., Calif., B. C.
Larrada montana Cresson, 1865. Ent. Soe. Phila., Proc. 4: 465. °.
Tachysphex inusitatus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 524. ¢.
Tachysphex compactus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 528. 6.
Tachysphex triquetrus Fox, 1894 (1898). Acad. Nat. Sci. Phila., Proc. 45: 528. 9. N. syn. (W.
J. Pulawski).

Superfamily SPHECOIDEA 1629

parvulus (Cresson). Colo., Wyo., Idaho, Oreg.; Canad. Zone. Ecology: Nests in sand bank with
dense vegetation, 1 cell per nest, 1-2 prey per cell. Parasite: Miltogrammini sp. Prey:
Acrididae spp.
Larrada parvula Cresson, 1865. Ent. Soe. Phila., Proe. 4: 465. ¢.
Tachysphex consimilis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 526. 2, 3.
Tachysphex agryrotrichus Rohwer, 1911. U. S. Natl. Mus., Proc. 4: 572. 3.

Biology: Alcock, 1973. Wasmann Jour. Biol. 31: 329-331, fig. 9 (nest, prey capture and
transport, parasite).
pauxillus Fox. B. C., Wash., Oreg., Idaho, Colo., Utah, Calif. Ecology: Nests in sand. Parasite:
Senotainia sp. ? Prey: Melanoplus sp. nymph. Predator: Philanthus pulcher D. T.
Tachysphex pauxillus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 530. 2.
Tachysphex nigrior Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 530. 2, d. N. syn.
(W. J. Pulawski).

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 490 (nest, prey, parasite, predator).

pechumani Krombein. N. Y., N. J., Mich. Ecology: Nests in sparsely vegetated pine barrens.
Tachysphex tarsatus pechumani Krombein, 1938. Ent. Soc. Amer., Ann. 31: 468. 2.

Taxonomy: Kurczewski, Elliott and Vasey, 1970. Ent. Soc. Amer., Ann. 63: 1594-1597, 5 figs.
ORO:
powelli Bohart. Calif.
Tachysphex powelli Bohart, 1962. Biol. Soc. Wash., Proc. 75: 35, figs. 1-3. 6, °.
psilocerus Kohl. Colo.; Mexico (Baja California, Durango, Mexico).
Tachysphex psilocerus Kohl, 1884 (1883). Zool.-Bot. Gesell. Wien, Verhandl. 33: 374. 2.
Tachysphex helianthi Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 570. 2. N. syn. (W. J.
Pulawski).
Tachysphex nitelopteroides Williams, 1958. Pan-Pacifie Ent. 34: 207, fig. 1. 2, 6. N. syn.
(W. J. Pulawski).
punctifrons (Fox). U.S. east of Rocky Mts., north to Mich. and N. Y. Prey: Melanoplus sp.
probably bivittatus (Say).
Larra punctifrons Fox, 1891. Ent. News 2: 194. °.
Tachysphex fedorensis Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 576. 2, 6.
quebecensis (Provancher). Que., Ont., Alta., N. W. T., Maine, Mass.
Larra quebecensis Provancher, 1882. Nat. Canad. 13: 150. 2, ¢.
Larra abdominalis Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 266.
Lapsus.
sculptilis Fox. Nebr. (Blaine Co.), Colo., Ariz., Nev., Calif.; montane.
Tachysphex asperatus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 516. ?. N. syn. (W.
J. Pulawski).
Tachysphex sculptilis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 517. 9.
Tachysphex nigrescens Rohwer, 1908. Ent. News 19: 220. ?. N. syn. (W. J. Pulawski).
Tachysphex sphecodoides Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 578. °.
semirufus (Cresson). Yukon to Calif., east to Ont., Wyo., Colo., Ariz. Prey: Melanoplus spretus
(Walsh), young nymphs.
Larrada semirufa Cresson, 1865. Ent. Soc. Phila., Proc. 4: 464. °.
Tachysphex punctulatus Smith, 1906. Ent. News 17: 246. 2. Preoce.
Tachysphex puncticeps Smith, 1908. Nebr. Univ., Studies 8: 381. N. name.
Tachysphex giffardi Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 244. 2.

Biology: Riley, 1878. U. S. Dept. Agr., Ent. Comn. Rpt. 1: 317 (prey).
sonorensis (Cameron). U. S. west of 100 degrees; Mexico (Baja California, Sonora, Chihuahua,

Puebla).

Larra sonorensis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 50. °.

Tachysphex dakotensis Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 98. 2. N. syn. (W. J.
Pulawski).

Tachysphex schlingeri Bohart, 1962. Biol. Soc. Wash., Proc. 75: 36, figs. 4-6. d, 2. N. syn.
(W. J. Pulawski).

1630 Hymenoptera in America North of Mexico

tarsatus (Say). Transcont. in U. S., Canad., Transit., and Austr. Zones.; Mexico (Baja

California, Jalisco). Ecology: Makes unicellular nest in a variety of vegetated, sandy
soils. Parasite: Diptera sp. Prey: Melanoplus spp., Trimerotropis sp. ?; only nymphs
have been reported as prey.

Larra tarsata Say, 1823. Western Quart. Rptr. 2: 78. °.

Tachysphex dubius Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proe. 45: 515. d. Preoce.

Tachysphex dubiosus Dalla Torre, 1897. Cat. Hym., v. 8, p. 679. N. name.

Tachysphex hitei Rohwer, 1908. Ent. News 19: 221. °.

Tachysphex sanguinosus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 414. 2. N. syn. (W. J.
Pulawski).

Tachysphex zimmeri Mickel, 1916. Amer. Ent. Soe., Trans. 42: 415. °.

Biology: Riley, 1880. U. S. Dept. Agr., Ent. Comn. Rpt. 2: 270-271 (prey). —Peckham and
Peckham, 1900. Wis. Nat. Hist. Soe., Bul. 1: 89-90 (nest, prey transport). — Williams, 1914
(1913). Kans. Univ. Sci. Bul. 17: 203-206, fig. 117 (mest, prey hunt and transport).
—Kurezewski, 1966. Kans. Ent. Soe., Jour. 39: 439 (nest). —Evans, 1970. Mus. Compar.
Zool., Bul. 140: 490 (prey transport, nest). —Aleock and Gamboa, 1975. Ariz. Acad. Sci.,
Jour. 10: 164 (nest, prey).

tenuipunctus Fox. West. Canada and U. S. in mountains south to Ariz., N. Mex. Ecology:
Makes unicellular nest in sandy rangeland, stores 1-2 prey. Parasite: Taxigramma
heteroneura (Meig.); Sphaeropthalma orestes (Fox). Prey: Oedaleonotus enigma
(Seudd.), Aulocara elliotti (Thom.), Melanoplus sp.; only nymphs have been recorded as
prey.
Tachysphex tenuipunctus Fox, 1894 (1893). Acad. Nat. Sci., Phila., Proc. 45: 525. 9.
Tachysphex granulosus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 413. 9.

Biology: Newton, 1956. Jour. Econ. Ent. 49: 615-619 (nest, prey hunt and transport, life cycle,
egg, larva, cocoon, parasite).
texanus (Cresson). Transcont., north to N. Y., Mich., Mont., south to Fla., Tex., Ariz., Calif.;
Mexico (Jalisco). Prey: Oedipodinae sp. nymph.
Larrada texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 214. 9, d.
Tachysphex sepulcralis Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 169. 2, ¢. N. syn.
(W. J. Pulawski).
Tachysphex maneei Banks, 1921. Ent. Soc. Amer., Ann. 14: 19. 2. N. syn. (W. J. Pulawski).

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 206 (prey).
williamsi Bohart. Calif. (San Francisco Co.).
Tachsphex williamsi Bohart, 1962. Biol. Soc. Wash., Proc. 75: 38, figs. 10-12. d, °.

SPECIES GROUP TERMINATUS

alpestris Rohwer. N. W. T., Alta., B. C. to Calif., eastwards to Wyo., Nebr., Colo. and N. Mex.,
south to Costa Rica.
Tachysphex foxti var. alpestris Rohwer, 1908. Ent. News 19: 233. °.
apicalis apicalis Fox, n. status (W. J. Pulawski). D. C., N. C., Ga., Fla. Ecology: Makes
multicellular nests with up to 7 prey per cell in open sand, frequently in sand cliffs.
Prey: Melanoplus sp. possibly pwer (Scudd.) nymph, Acrididae sp.
Tachysphex apicalis Fox, 1893. N. Y. Ent. Soc., Jour. 1: 538. 9, d.
Tachysphex fumipennis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 518. °.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 288-289, figs. 99-103 (larva).

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 15 (nest, prey transport, egg).
—Kurezewski, 1966. Kans. Ent. Soe., Jour. 39: 436-453, figs. 2, 3, 5, 6b, 8, 9 (nest, male
behavior). —Kurezewski and Snyder, 1968. Conservationist 23 (2): 30-31, 6 figs. (nest,
prey).

apicalis fusus Fox, n. status (W. J. Pulawski). Transcont. except southeast. U. S., north to Md.,
Ky., N. Dak., Wash., south to Central America; introduced into Hawaii. Ecology: Nests
in mortar between foundation rocks, in abandoned insect burrows in ground, in earthen
bank and sand cliffs, stores several prey per cell in multicellular nests. Prey:

Superfamily SPHECOIDEA 1631

Melanoplus sp., Chortophaga sp., Oxya sp. ?, Tryxalinae sp.; only nymphs are used so
far as recorded.
Tachysphex fusus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 519. 2, d.
Tachysphex foxii Rohwer, 1908. Ent. News 19: 222. 2.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 201 (prey, nest). —Rau and Rau, 1918.
Wasp Studies Afield, pp. 149-150 (nest). —Kurezewski and Snyder, 1968. Conservationist
23 (2): 30-31 (nest, prey).
clarconis Viereck. Western States north to B. C., Wash. and Wyo., south to Calif. and N. Mex.
Tachysphex clarconis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 211. 9.
Tachysphex plesia Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 245. 9. N. syn. (W. J.
Pulawski).
linsleyi Bohart. Western States north to Idaho and Wyo., eastwards to Colo. and Tex., south
to Calif.; Mexico. Prey: Acrididae sp. nymph.
Tachysphex linsleyi Bohart, 1962. Biol. Soc. Wash., Proc. 75: 35, figs. 7-9. 5, 2.

Biology: Krombein, 1967. U. S. Dept. Agr., Monog. 2, Sup. 2, p. 393 (prey).
similis Rohwer. North to N. B. and Sask., west to Alta., Utah, N. Mex.; Mexico. Ecology:
Nests in open sand, makes multicellular nest, stores 4-10 prey per cell. Parasite:
Hedychridium fletcheri Bod. Prey: Radinotatum sp., Atenopedes sp., Schistocerca sp.,
Melanoplus sp.; only nymphs have been recorded as prey.
Tachysphex similis Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 51. 9, d.
Tachysphex similans Rohwer, 1910. Ent. Soc. Wash., Proce. 12: 52. °.

Taxonomy: Elliott and Kurezewski, 1974. Ent. Soc. Amer., Ann. 67: 725-727, 2 figs. (character
displacement).

Biology: Krombein and Evans, 1955. Ent. Soc. Wash., Proce. 57: 231 (prey transport).
—Krombein, 1964. Amer. Mus. Novitates 2201: 15-17, fig. 1 (nest, prey transport, egg).
—Kurczewski, 1966. Kans. Ent. Soc., Jour. 39: 436-453, fig. 4 (nest, male behavior).
—Kurczewski, 1967. Kans. Ent. Soc., Jour. 40: 278-284 (parasite).

terminatus (Smith). Transcont. except Fla. and Pacific States, north to P. EK. I., Que., Ont.,

Man., Sask., N. W. T., south to Colombia and north. brazil. Parasite: Anthrax a.
albofasciatus Macq.; Phrosinella fulvicornis (Coq.), Senotainia trilineata (Wulp). Prey:
Phaneroptera spp.; Chortophaga viridifasciata DeG., Chloealtis conspersa Harr.,
Chorthippus curtipennis (Harr.), Dissosteira carolina (L.), Pardalophora apiculata
Harr. ?, Melanoplus bivittatus (Say), M. femurrubrum (DeG.), M. keeleri luridus
(Dodge), M. spp., Tryxalus spp., Tryxalinae sp., Syrbula admirabilis Uhl.,
Dichromorpha viridis Seudd.; all recorded prey were nymphs.

Larrada terminata Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 291. 6.

Larra minor Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 268. 2, ¢.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 118, figs. 21-28 (larva). —Elliott and
Kurezewski, 1974. Ent. Soc. Amer., Ann. 67: 725-727, 2 figs. (character displacement).

Biology: Ashmead, 1894. Psyche 7: 63 (prey). — Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8:
201 (prey). —Rau and Rau, 1918. Wasp Studies Afield, pp. 144-149 (nest, prey transport).
—Rau, 1927. Acad. Sci. St. Louis, Trans. 25: 188-190 (nest, prey, life cycle). —Rau, 1946.
Brooklyn Ent. Soc., Bul. 41: 10. —Strandtmann, 1953. Kans. Ent. Soe., Jour. 26: 49-51, fig. 2
(nest, prey transport, life cycle). —Kurczewski, 1966. Kans. Ent. Soe., Jour. 39: 317-322, 1
fig. (nest, prey transport). —Kurczewski, 1966. Kans. Ent. Soc., Jour. 39: 436-453, figs. 1,
6a, 7 (nest, male behavior, mating). —Kurezewski and Harris, 1968. N. Y. Ent. Soc., Jour.
76: 81-83 (parasites, nest). —Kurezewski and Snyder, 1968. Conservationist 23 (2): 28-31, 5
figs. (nest, prey, life cycle, parasite). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 491
(nest, prey).

SPECIES GRouP UNDATUS
ashmeadii Fox. Oreg. to Calif., east to centr. Tex., Kans. and Wyo. in deserts. Ecology: Nests
in sand. Prey: Metator sp. nymph, Cordillacris crenulata (Brun.) adult, Trachyrhachys

kiowa (Thom.) adult, Opeia sp. nymph, Phlibostroma sp. adult.
Tachysphex ashmeadii Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 509. 2.

1632 Hymenoptera in America North of Mexico

Tachysphex posterus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 510. °.
Tachysphex spinosus Fox, 1894 (1893). Acad. Nat. Sei. Phila., Proc. 45: 511. °.
Tachysphex spissatus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 515. ¢.

Larra rufipes Provancher, 1895. Nat. Canad. 22: 129. 9.

Tachysphex propinquus Viereck, 1904. Ent. News 15: 85. 2. N. syn. (W. J. Pulawski).

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Studies 8: 202-203, fig. 112 (nest, prey hunt
and transport). —Krombein, 1967. U.S. Dept. Agr., Monog. 2, Sup. 2, pp. 392-393 (prey).
—Aleock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 164 (nest, prey).

SPECIES GROUP BRULLII

alayoi Pulawski. Fla. (Dania in Broward Co.); West Indies, Cuba to Virgin Islands. Prey:
Blattidae sp. nymph.
Tachysphex alayoi Pulawski, 1974. Polskie Pismo Ent. 44: 84, figs. 121-128. 2, 6.

Biology: Pulawski, 1974. Polskie Pismo Ent. 44: 87 (prey).
belfragei (Cresson). D. C. to Fla. west to Iowa, Nebr., Tex. Prey: Conocephalus sp. nymphs.
Larrada belfragei Cresson, 1872. Amer. Ent. Soc., Trans. 4: 215. @.
Tachytes minimus Fox, 1892. Amer. Ent. Soc., Trans. 19: 248. 9.

Biology: Krombein, 1967. U. S. Dept. Agr., Monog. 2, Sup. 2, p. 392 (prey).

maurus Rohwer. Tex. to Ariz.; Mexico (Sonora).
Tachysphex maurus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 575. 2, 6.

mundus Fox. Transcontinental, north to south. Canada, south to Mexico (Chiapas).

Larra rufitarsis Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 50, pl. 4, fig. 11. d.
Preoce. N. syn. (W. J. Pulawski).

Tachysphex aequalis Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 517. ¢. N. syn. (W.
J. Pulawski).

Tachysphex exsectus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 526. 2 (¢ misdet.).
N. syn. (W. J. Pulawski).

Tachysphex mundus Fox, 1894 (1893). Acad. Nat. Sei. Phila., Proc. 45: 531. 2, ¢.

Tachysphex johnsoni Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 573. 2. N. syn. (W. J.

Pulawski).

Tachysphex opwanus Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 574. 36. N. syn. (W. J.
Pulawski). ;

Tachysphex robustior Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 164. ¢. N. syn. (W. J.
Pulawski).

Tachysphex crenuloides Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 168. 2. N. syn. (W.
J. Pulawski).

Tachysphex washingtoni Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 172. 2. N. syn. (W. J.
Pulawski).

SPECIES GROUP JULLIANI
cockerellae Rohwer. Calif., Nev., Ariz.; Mexico south to Colombia.
Tachysphex cockerellae Rohwer 1914. U.S. Natl. Mus., Proc. 47: 518. d.

coquilletti Rohwer. Kans., Okla., Colo., N. Mex., Ariz., Nev., Calif.; Mexico (Baja California,
Durango, Zacatecas). Prey: Litoneutria minor (Scudd.), L. sp., nymph.
Tachysphex coquilletti Rohwer, 1911. U.S. Natl Mus., Proc. 40: 572. 2, ¢.
Tachysphex dentatus Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 169. 2.

Biology: Kurezewski, 1966. Kans. Ent. Soc., Jour. 39: 317 (prey). —Krombein, 1967. U.S.

Dept. Agr., Monog. 2, Sup. 2, p. 393 (prey). —Aleock and Gamboa, 1975. Ariz. Acad. Sci.,
Jour. 10: 164-165 (nest, orientation flight, prey).

Genus PROSOPIGASTRA Costa

Prosopigastra Costa, 1867. Mus. Zool. Napoli Ann. 4: 88.
Type-species: Prosopigastra punctatissima Costa. Monotypice.
Homogambrus Kohl, 1889. K. K. Naturhist. Hofmus., Ann. 4: 191.
Type-species: Tachysphex globiceps Morawitz. Monotypic.
Hologambrus Morice, 1897. Ent. Soc. London, Trans., p. 309. Lapsus or emend.

Superfamily SPHECOIDEA 1633

Three Old World species make multicellular nests in soil, and two of them utilize pre-existing
burrows of other insects.. Prey consists of Hemiptera and Homoptera belonging to the families
Lygaeidae, Tropiduchidae or Pentatomidae.

nearctica Bohart. Calif., Ariz.
Prosopigastra nearctica Bohart, 1958. Ent. Soc. Wash., Proce. 60: 122, 6 figs. d, °.

SUBFAMILY MISCOPHINAE
Two inadequate revisions are listed under the heading Subfamily Larrinae.
Genus LYRODA Say

Lyrops subg. Lyroda Say, 1837. Boston Jour. Nat. Hist. 1: 372.
Type-species: Lyrops (Lyroda) subita Say. Desig. by Patton, 1881.
Morphota Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 293.
Type-species: Morphota fasciata Smith. Desig. by Pate, 1937.
Odontolarra Cameron, 1900. Ann. and Mag. Nat. Hist. (7) 5: 35.
Type-species: Odontolarra rufiventris Cameron. Monotypic.
Lyrodon Howard, 1901. Insect Book, pl. 6, fig. 5. Lapsus. Preoce.

These wasps are ground-nesting and may utilize pre-existing burrows or cavities. One North
American species preys upon crickets (Gryllidae) and several extralimital species use grouse
locusts (Tetrigidae).

subita (Say). Transcont. in south Canada and U.S. Ecology: Nests in pre-existing burrows or
cavities, makes up to 2 cells per nest and stores up to 9 prey per cell. Parasite: Metopia
argyrocephala (Meig.). Prey: Nemobius carolinus Seudd., N. fasciatus (DeG.), N. spp.;
nymphs.
Lyrops (Lyroda) subita Say, 1837. Boston Jour. Nat. Hist. 1: 372. °.
Larrada arcuata Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 293. 2.
Lyroda cockerelli Rohwer, 1909. Ent. News 20: 369. 6.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 281-282, figs. 78-84 (larva).

Biology: Patton, 1892. Ent. News 3: 90 (prey transport). —Peckham and Peckham, 1898. Wis.
Geol. Nat. Hist. Survey, Bul. 2: 169-171 (nest, prey transport, life cycle). —Peckham and
Peckham, 1905. Wasps, Social and Solitary, pp. 253-256 (nest, prey transport, life cycle).
—Evans, 1964. Amer. Ent. Soc., Trans. 90: 282 (nest, prey transport, parasite).
—Kurezewski and Kurezewski, 1971. Kans. Ent. Soc., Jour. 44: 132 (prey).

triloba (Say). Canada, D. C., Ill., Ind., Kans., La., Tex.
Lyrops (Lyroda) triloba Say, 1837. Boston Jour. Nat. Hist. 1: 372. °.
Lyrops (Lyroda) caliptera Say, 1837. Boston Jour. Nat. Hist. 1: 373. Lapsus.

Genus PLENOCULUS Fox

Plenoculus Fox, 1893. Psyche 6: 554.
Type-species: Plenoculus davisi Fox. Monotypice.
Ptygosphex Gussakovskij, 1928. Inst. Zool. Appl. Phytopath., Leningrad, Bul. 4: 18.
Misspelled Ptigosphex in generic heading.
Type-species: Ptygosphex murgabensis Gussakovskij. Orig. desig.
Pavlovskia Gussakovskij, 1935. Trav. Fil. Acad. Sci. URSS, Tadjikistan 5: 424.
Type-species: Pavlovskia tadzhika Gussakovskij. Orig. desig.

These wasps nest in sand and construct 1- or multicelled nests. In North America several taxa

prey upon Hemiptera or Homoptera but one species uses pyralidid caterpillars as prey.
Revision: Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 1-49, 90 figs. (N. Amer. spp.).
Taxonomy: Ashmead, 1899. Psyche 8: 337-338 (key to some N. Amer. spp.).

boharti Williams. Southern Calif.
Plenoculus boharti Williams, 1960. Calif. Acad. Sci., Proce. (4) 31: 28, figs. 40, 61, 83, 84. 2,

é.

1634 Hymenoptera in America North of Mexico

boregensis boregensis Williams. Calif. (Borego in San Diego Co.).
Plenoculus boregensis boregensis Williams, 1960. Calif. Acad. Sci., Proe. (4) 31: 20, figs.
26-29. 2, d.
boregensis perniger Williams. Calif. (Thousand Palms).
Plenoculus boregensis perniger Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 21. 6.
cockerellii Fox. Southern Calif. to western Tex.; Mexico (Baja California, Guerrero). Ecology:
Nests in sand. Prey: Pyralididae spp. larvae.
Plenoculus Cockerellii Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc., 45: 538. °.

Biology: Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 46-47 (nest, prey transport).
cuneatus Williams. Southern Calif. and Nev.
Plenoculus cuneatus Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 12, figs. 10, 17, 18, 24,
2 OatOr
davisi atlanticus Viereck. Coastal Conn. to Fla., Tex. Ecology: Nests in open sand, provides up
to 7 prey per cell. Prey: Phytocoris sp. nymphs and adult.
Plenoculus atlanticus Viereck, 1902. Ent. News 13: 74. 6.

Taxonomy: Krombein, 1955. Ent. Soc. Wash., Proc. 57: 146.

Biology: Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 181, 191, 192, 194-199, 202, 205 (nest,
prey transport, egg).
davisi davisi Fox. Conn. to Fla. west to Alaska, B. C., Idaho and Calif.; Mexico (Baja

California, Sinaloa, Nayarit, Durango). Ecology: Nests in sand, constructs 1-4 cells per
nest and stores 2-24 prey per cell. Prey: Arhyssus lateralis (Say) adult; Aphidae sp.
immature; Adelphocoris rapidus (Say), Amblytylus nasutus (Kirschb.), Campylomma
verbasci (Meyer), Chlamydatus associatus (Uhl.), Collaria sp. ?, Halticus bracteatus
(Say), Lopidea robiniae (Uhl.), Lygus lineolaris (Beauv.), Neolygus sp. ?, Orthocephalus
mutabilis (Fall.), Orthotylus chorionis (Say), O. marginatus (Uhl.), O. quercicola Knight,
O. sp., Pilophorus amoenus Uhl. P. sp., Plagiognathus chrysanthemi (Wolff), P. politus
Uhl., P. sp., Poecilocapsus lineatus (F.), Psallus seriatus Reut. ?, Stenotus binotatus
(F.), Trigonotylus ruficornis (Geoff.), T. sp., Mirinae spp., Phylini spp.; adults are stored
more commonly than nymphs. Predator: Philanthus pulcher D. T., P. crabroniformis
Sm.

Plenoculus davisi Fox, 1893. Psyche 6: 554. °, 6.

Plenoculus abdominalis Ashmead, 1899. Psyche 8: 339. 3.

Plenoculus apicalis Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 175. °, d.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 166-167, figs. 54-59 (larva).

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 207-208, fig. 120 (nest, prey transport).
—Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 46 (prey). —Evans, 1961. Ent. News 72:
223-228 (nest, prey transport, life cycle). —Kurezewski, 1968. Kans. Ent. Soc., Jour. 41:
179-207, 20 figs. (nest, prey transport, egg). —Evans, 1970. Mus. Compar. Zool., Bul. 140:
488 (nest, prey, predator).

davisi gracilis Williams. Calif. (Riverside Co.).
Plenoculus davisi gracilis Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 42, fig. 54. °.
davisi mojavensis Williams. Southern Calif., Ariz., N. Mex.
Plenoculus davisi mojavensis Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 39, figs. 86, 88.
OFEGr
davisi transversus Williams. Calif. (Tulare and Riverside Counties).
Plenoculus davisi transversus Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 40, figs. 8, 57,
59. 2.
deserti Williams. Southern Calif.
Plenoculus deserti Williams, 1960. Calif. Acad. Sci., Proce. (4) 31: 26, figs. 85, 87. d, 9.
gillaspyi Krombein. Tex. (Williamson Co.).
Plenoculus gillaspyi Krombein, 1938. Ent. Soc. Amer., Ann. 31: 468. °.
palmarum Williams. Southern Calif.
Plenoculus palmarum Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 22, figs. 9, 36, 37, 71,
Tom SmOnme

Superfamily SPHECOIDEA 1635

parvus Fox. N. Mex. (Las Cruces).
Plenoculus parvus Fox, 1897. Ent. News 8: 71. °.
propinquus Fox. Wash. to Calif. east to Idaho, Colo., N. Mex. Ecology: Nests in open sand.
Prey: Lygus desertus Knight, Miridae sp.; adults.
Plenoculus propinquus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 537. 9.
Plenoculus propinquus var. rufescens Cockerell, 1898. Davenport Acad. Sci., Proc. 7: 144.

Biology: Kurezewski, 1968. Kans. Ent. Soc., Jour. 41: 181, 191, 197, 198, 202 (nest, prey
transport, egg). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 488 (nest, prey).
sinuatus Williams. Southern Calif.
Plenoculus sinuatus Williams, 1960. Calif. Acad. Sei., Proce. (4) 31: 7, figs. 30, 32, 35. 2, ¢.

stygius Williams. Southern Calif. and Ariz. Prey: Miridae sp.
Plenoculus stygius Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 31, figs. 5, 62, 90. 2, d.

Biology: Williams, 1960. Calif. Acad. Sci., Proce. (4) 31: 46 (prey).
timberlakei Williams. Southern Calif. and Ariz.
Plenoculus timberlakei Williams, 1960. Calif. Acad. Sci., Proc. (4) 31: 11, figs. 2, 11, 19-22,
Ble SS AUmAD Oc audilen OeuGle

Genus SOLIERELLA Spinola

Solierella Spinola, 1851. In Gay, Hist. Fis. Pol. Chile, Zool., v. 6, p. 349.
Type-species: Solierella miscophoides Spinola. Monotypic.
Silaon Piccioli, 1869. Soc. Ent. Ital., Bol. 1: 282.
Type-species: Silaon compedita Piccioli. Monotypic.
Sylaon Piccioli, 1870. Soe. Ent. Ital., Bol. 2: pl. 1. Lapsus or emend.
Niteliopsis Saunders, 1873. Ent. Soc. London, Trans., p. 410.
Type-species: Niteliopsis pisonoides Saunders. Monotypic.
Ammosphecidium Kohl, 1878. Zool.-Bot. Gesell. Wien, Verh. 27: 701.
Type-species: Ammosphecidium Helleri Kohl. Monotypic.
Sylaon Kohl, 1885. Zool.-Bot. Gesell. Wien, Verh. 34: 290. Emend. or lapsus.
Lautara Herbst. 1920. Mus. Nace. Chile, Bol. 11: 217.
Type-species: Lautara Jaffueli Herbst. Monotypic.

Our species usually build multicelled nests in pre-existing cavities in lumber, twigs, stems,
galls, nut hulls, or in abandoned burrows in the ground. The prey of some North American spe-
cies has been recorded as Hemiptera of several families, Acrididae and Psocoptera.

Revision: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 355-417 (Calif. spp.).

Taxonomy: Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 108-110 (key to some spp.).

abdominalis Williams. Calif.
Solierella abdominalis Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 381. 2, ¢.
affinis (Rohwer). Kans., Colo., Wyo., Idaho, Calif. Ecology: Nests in soil, possibly in
pre-existing burrows, stores 4 or more prey per cell. Prey: Nabis sp. nymphs.
Niteliopsis affinis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 113. 2, ¢.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 488-489 (nest, prey).

albipes (Ashmead). Colo., Idaho, Calif.

Plenoculus albipes Ashmead, 1899. Psyche 8: 339. 6.
arcuata Williams. Calif. (San Rafael, Menlo Park).

Solierella arcuata Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 378. 2, ¢.
australis Williams. Calif. (Riverside).

Solierella australis Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 379. 2, 6.
bicolor Williams. Calif.

Solierella bicolor Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 382. @.
blaisdelli (Bridwell). Calif., Ariz., Idaho. Ecology: Nests in stems of Eriogonwm, sumac,

elderberry, raspberry, and in borings in wood. Parasite: Pseudolopyga taylori (Bod.),

1636 Hymenoptera in America North of Mexico

Hedychridium solierellae Boh. and Brum.; Lomachaeta variegata Mick. Prey: Nysius
raphanus How., N. tenellus Barber, N. ericae minutus Uhl., N. sp.; nymphs.
Silaon blaisdelli Bridwell, 1920. Hawaii. Ent. Soc., Proc. 4: 401. °.

Taxonomy: Evans, 1958. Amer. Ent. Soc., Trans. 84: 123, fig. 42 (larva). —Evans, 1964. Amer.
Ent. Soc., Trans. 90: 285 (larva).

Biology: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 394 (prey). —Parker and Bohart, 1966.

Pan-Pacific Ent. 42: 95 (nest, parasite). —Krombein, 1967. Trap-nesting wasps and bees,
pp. 177-178 (nest, prey, parasite, life cycle). —Carrillo, 1967. Pan-Pacific Ent. 43: 201-203
(larval instars). —Parker and Bohart, 1968. Pan-Pacifie Ent. 44: 3 (nest, parasites).
—Carrillo and Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 676-677 (nest, prey, parasites,
life cycle).

boharti Williams. Calif.
Solierella boharti Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 366. °.
Solierella lasseni Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 366. 2, 6.

Taxonomy: Williams, 1953. Pan-Pacific Ent. 29: 157 (synonymy).
boregensis Williams. Calif. (San Diego Co.).
Solierella boregensis Williams, 1958. Pan-Pacifie Ent. 34: 208, fig. 2. 2, ¢.
bridwelli Williams. Calif.
Solierella bridwelli Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 384. 2, d.
californica Williams. Calif. (Los Angeles).
Solierella californica Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 387. 2, d.
clypeata Williams. Calif.
Solierella clypeata Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 376. 9.
corizi Williams. Calif., Tex. Ecology: Nests in burrows of other insects in soil, makes 2-3 cells
per nest and stores 3-4 prey per cell. Prey: Corizus hyalinus (F.) adult and nymphs;
Perilabus abbreviatus (Uhl.) nymph.
Solierella corizi Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 372. 2, d.

Biology: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 392-394 (nest, prey transport).
fossor (Rohwer). Colo., N. Mex., Ariz. Prey: Oedipodinae sp. nymph.
Niteliopsis foxti Viereck, 1906. Amer. Ent. Soc., Trans. 32: 207. 2. Preoce.
Niteliopsis fossor Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 116 °, d.

Biology: Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 116 (prey).
foxii (Viereck). N. J. (North Woodbury).
Plenoculus foxii Viereck, 1902. Ent. News 13: 73. 2, d.
inermis (Cresson). N. C., Fla., Ill. to Tex. west to Idaho and Colo. Ecology: Nests in abandoned
burrows of other arthropods in sand or clay. Prey: Thyanta pallidovirens (Stal);
Harmostes reflexulus (Say), Liorhyssus hyalinatus (F.),; Chariesterus antennator (F.);
Rhynocoris ventralis (Say)?; Miridae sp.; all prey stored were nymphs.
Nysson? inerme Cresson, 1872. Amer. Ent. Soc., Trans. 4: 224. °.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 208-209, pl. 30, fig. 119 (nest, prey).

—Kurezewski, 1967. Kans. Ent. Soc., Jour. 40: 203-208, 1 fig. (nest, prey, egg).
kansensis (Williams). Kans.

Niteliopsis kansensis Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 178. °.
levis Williams. Calif.

Solierella levis Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 383. @.
lucida (Rohwer). Colo. (Boulder).

Niteliopsis lucidus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 109. ¢.
major (Rohwer). Wash., Calif.

Silaon major Rohwer, 1917. U.S. Natl. Mus., Proc. 53: 247. @.
masoni Williams. Calif. (Thousand Palms).

Solierella masoni Williams, 1959. Ent. Soe. Wash., Proc. 61: 74, 5 figs. 2, 6.
mirifica Pate. Ariz. (Pima Co.).

Solierella (Silaon) mirificus Pate, 1934. Ent. News 45: 248. ¢.

Superfamily SPHECOIDEA 1637

modesta (Rohwer). Colo. (Boulder).

Niteliopsis modestus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 116. 6.
nigrans Krombein. W. Va., Colo., Calif.

Niteliopsis niger Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 115. 2, d. Preoce.

Solierella nigrans Krombein, 1951. U. S. Dept. Agr., Monog. 2: 943. N. name.
nitens Williams. Calif. Ecology: Nests in ground.

Solierella nitens Williams, 1950. Calif. Acad. Sci., Proce. (4) 26: 376. 2.

Biology: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 395 (nest).
peckhami (Ashmead). N. Y. to Fla. west to Idaho and Calif.; adventive in Hawaii and Marshall

Islands. Ecology: Nests in cavities in twigs, stems of elderberry, sumac, raspberry, and
in cavities in almond hulls. Parasite: Psewdolopyga taylori (Bod.). Prey: Nysius
raphanus How., N. tenellus Barber, N. ericae minutus Uhl. N. sp., Pachybrachius sp.;
nymphs.

Plenoculus peckhami Ashmead, 1897. Psyche 8: 130. ¢.

Plenoculus niger Ashmead, 1899. Psyche 8: 339. 9.

Silaon rohweri Bridwell, 1920. Hawaii. Ent. Soc., Proc. 4: 398.

Solierella (Silaon) arenaria Krombein, 1939. Brooklyn Ent. Soc., Bul. 34: 139. @.

Taxonomy: Carrillo and Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 673, figs. 3-6 (egg, larva,
pupa).

Biology: Peckham and Peckham, 1905. Wasps, Social and Solitary, pp. 95-96 (nest, prey).
—Rau and Rau, 1918. Wasp Studies Afield, pp. 134-135 (nest, parasite). —Bridwell, 1920.
Hawaii Ent. Soc., Proc. 4: 399-400 (nest, prey). —Williams, 1926. Hawaii. Ent. Soc., Proc. 6:
442-444, figs. 4-7 (nest, prey, life cycle). —Rau, 1928. Acad. Sci. St. Louis, Trans. 25:
375-378, figs. 48, 49 (nest, life cycle). —Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 394
(prey). —Carrillo, 1967. Pan-Pacific Ent. 43: 201-203 (larval instars). —Carrillo and
Caltagirone, 1970. Ent. Soc. Amer., Ann. 63: 673-676, figs. 1, 2, 7 (nest, prey, life cycle,
parasites).

plenoculoides plenoculoides (Fox). N. H. to Va., west to Colo. and Tex., Ariz. Ecology: Nests
in galls on goldenrod of Eurosta solidaginis (Fitch).
Niteliopsis plenoculoides Fox, 1893. Psyche 6: 555. 2.

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 943 (nest).
plenoculoides similis (Bridwell). Calif., Oreg. Ecology: Nests in borings in stems of Sambucus,
Foeniculum, Eriogonum and Umbelliferae, stores up to 4 prey per cell. Parasite:
Senotainia trilineata (Wulp); Eurytoma stigmi Ashm.; Lomachaeta variegata Mick.
Prey: Acrididae sp. nymphs.
Silaon similis Bridwell, 1920. Hawaii. Ent. Soc., Proc. 4: 402. °.
Biology: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 391-392 (nest, prey). —Parker and
Bohart, 1966. Pan-Pacifie Ent. 42: 95 (nest, parasites).
prosopidis Williams. Calif. (San Bernardino Co.). On flowers of Prosopis.
Solierella mandibularis Williams, 1958. Pan-Pacific Ent. 34: 212, figs. 4, 4a. 2. Preoce.
Solierella prosopidis Williams, 1959. Pan-Pacifie Ent. 35: 116. N. name.
sayi (Rohwer). Colo., Calif. Ecology: Nests in sand, stores several prey per cell. Prey: Psocus
californicus Bks. adults; Lepidilla kelloggi Ribago.
Niteliopsis sayi Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 114. 2, d.

Biology: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 395 (nest, prey).
semirugosa Williams. Calif. (San Diego, Riverside and Yolo Counties).
Solierella semirugosa Williams, 1958. Pan-Pacifie Ent. 34: 210, figs. 3-3d. 2, ¢.
sonorae Williams. Calif.
Solierella sonorae Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 368. °.
striatipes (Ashmead). Calif.; Mexico (Baja California). Ecology: Makes unicellular nest in soil
which may contain 1-2 prey each bearing an egg. Parasite: Taxigramma sp.? Prey:
Melanoplus ligneolus Seudd. adult and penultimate instar nymph.
Niteliopsis striatipes Ashmead, 1899. Ent. News 10: 9. “2” = d.

1638 Hymenoptera in America North of Mexico

Biology: Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 389-391, fig. 3 (nest, prey hunt and

transport).
timberlakei Williams. Calif.

Solierella timberlakei Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 380. °.
vandykei Williams. Calif. (Tahoe).

Solierella vandykei Williams, 1950. Calif. Acad. Sci., Proc. (4) 26: 371. 2.
vierecki (Rohwer). Colo., Ariz., Calif.

Niteliopsis vierecki Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 112. 2, 6.

Niteliopsis parvus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 113. ¢.
weberi Williams. Calif. (Riverside).

Solierella weberi Williams, 1955. Pan-Pacifie Ent. 31: 85, 1 fig. d.

Genus MISCOPHUS Jurine

The shallow nests of these wasps are dug in loose sandy soil and contain one or several cells.
The prey consists of small, usually immature spiders and the number stored per cell ranges from
2 to 30.

Biology: Kurezewski, 1969. Kans. Ent. Soc., Jour. 42: 470-509, 13 figs. (comparative behavior).

Genus MISCOPHUS Subgenus MISCOPHUS Jurine

Miscophus Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 206.
Type-species: Miscophus bicolor Jurine. Monotypic.
americanus Fox. N. Y. to Fla. west to Colo., Kans. and Tex., N. W. T. Ecology: Makes a
1-celled nest in loose to well-packed sand and provides 5-11 small spiders per cell. Prey:
Theridion australe Bks., T. differens Em., T. murarium Em.
Miscophus americanus Fox, 1890 Ent. News 1: 138. 2.

Biology: Kurezewski, 1969. Kans. Ent. Soc., Jour. 42: 472-479, figs. 1, 3-5, 13 (nest, prey
transport, egg).

Genus MISCOPHUS Subgenus NITELOPTERUS Ashmead

Nitelopterus Ashmead, 1896. Jn Kohl, K. K. Naturhist. Hofmus., Ann. 11: 497.
Type-species: Nitelopterus slossonae Ashmead. Monotypic.

Miscophus subg. Hypomiscophus Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 321.
Type-species: Miscophus (Hypomiscophus) arenarum Cockerell. Monotypic.

Miscophinus Ashmead, 1898. Ent. News 9: 187.
Type-species: Miscophinus laticeps Ashmead. Orig. desig.

Biology: Powell, 1967. Kans. Ent. Soc., Jour. 40: 331-346, 1 fig. (comparative behavior of some
N. Amer. spp.).
aenescens (Bridwell). Oreg. (Mt. Jefferson).
Hypomiscophus aenescens Bridwell, 1920. Hawaii. Ent. Soc. Proc. 4: 394. ¢.
arenarum Cockerell. N. Mex. (Mesilla Park).
Miscophus (Hypomiscophus) arenarum Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2:
ozs oe
californicus (Ashmead). Calif., Ariz.
Miscophinus californicus Ashmead, 1898. Ent. News 9: 188. 6.
cyanurus (Rohwer). Colo. (Boulder).
Miscophinus cyanurus Rohwer. 1909. Amer. Ent. Soc., Trans. 35: 125. 2, d.
evansi (Krombein). Wyo., Wash. Ecology: Nests in sand, constructs up to 6 cells per nest and
stores 10-20 small spiders per cell. Parasite: Senotainia sp. in trilineata (Wulp) complex
? Prey: Dictyna sp. juveniles.
Nitelopterus evansi Krombein, 1963. Ent. News 74: 61. 2.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 283, figs. 88-92 (larva).

Biology: Evans, 1963. Ent. News 74: 234-236, figs. 1-2 (nest, prey, parasite). —Evans, 1970.
Mus. Compar. Zool., Bul. 140: 489 (nest, prey, parasite).

Superfamily SPHECOIDEA 1639

galei (Rohwer). Colo.
Miscophinus galei Rohwer, 1909. Amer. Ent. Soe., Trans. 35: 124. “2” = 6.
kansensis (Slansky). Kans. (Lawrence). Ecology: Makes a 1-celled nest in sand, usually stores
13-16 small spiders per cell, although as few as 7 or as many as 29 prey may be placed
in a single completed cell. Prey: Theridion rabuni Chamb. and Ivie, T. glaucescens
Beck., Euryopis texana Bks.; Tennesseellum formica (Em.); Grammonota sclerata Ivie
and Barr.; Tetragnatha laboriosa Hentz; Oxyopes salticus Hentz; Dictyna bicornis Em.,
D. sp.; mostly immatures are stored although adults are used occasionally.
Nitelopterus kansensis Slansky, 1969. Kans. Ent. Soc., Jour. 42: 467, 3 figs. 5, °.

Biology: Kurezewski, 1969. Kans. Ent. Soc., Jour. 42: 483-492, figs. 2, 6-8, 13 (nest, prey
transport, egg).
laticeps (Ashmead). Calif., Ariz. Ecology: Makes 1-celled nest in sand, stores 4-8 prey per
completed cell. Prey: Psilochorus rockefelleri Gertsch adults; Oecobius sp. juvenile;
Sitticus sp., Pellenes sp., both juveniles; Pardosa sp. juvenile.
Miscophinus laticeps Ashmead, 1898. Ent. News 9: 188. 2.

Biology: Cazier and Mortenson, 1965. Pan-Pacifie Ent. 41: 21-26, 1 fig. (nest, prey, egg).
—Powell, 1967. Kans. Ent. Soc., Jour. 40: 332-345 (nest, prey transport, egg, life cycle;
misdet. as californicus).

maurus (Rohwer). Colo.
Miscophinus maurus Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 126. 2, ¢.
nigrescens (Rohwer). Colo. (Rifle).
Miscophinus nigrescens Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 125. 2.
nigriceps (Rohwer). Calif. (Santa Monica).
Miscophinus nigriceps Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 587. @.
slossonae barberi (Krombein). Fla. Ecology: Makes a 1-celled nest in sand and provides up to
3 prey per cell. Prey: Meioneta formica (Em.); Pellenes sp.; Lycosa sp., Geolycosa sp.;
all prey were immatures.
Nitelopterus slossonae barberi Krombein, 1954. Amer. Ent. Soc., Trans. 80: 12, fig. 8. 2.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 283-284 (larva).

Biology: Krombein and Kurczewski, 1963. Biol. Soc. Wash., Proc. 76: 140-143 (nest, prey hunt
and transport). —Krombein, 1964. Amer. Mus. Novitates 2201: 14-15 (nest, prey, life cycle).
slossonae slossonae (Ashmead). Fla. Ecology: Makes a 1-celled nest in sand and provides up

to 11 small spiders in a completed cell. Prey: Habrocestum pulex (Hentz),
Metaphidippus sp.; Meioneta formica (Em.); Dictyna altamira Gertsch and Davis;
Arctosa sp., Lycosa spp., Pardosa sp.; Steatoda ergoniformis (Camb.); Tetragnatha
laboriosa Hentz; most prey were juveniles but occasionally adults were used.

Nitelopterus slossonae Ashmead, 1896. In Kohl, K. K. Naturhist. Hofmus., Ann. 11: 497. d.

Biology: Krombein and Evans, 1954. Ent. Soc. Wash., Proc. 56: 232 (prey transport).
—Krombein and Evans, 1955. Ent. Soc. Wash., Proc. 57: 231 (prey transport, nest).
—Krombein and Kurczewski, 1963. Biol. Soc. Wash., Proc. 76: 140-143 (nest, prey
transport). —Krombein, 1964. Amer. Mus. Novitates 2201: 13-14 (prey transport).
—Kurezewski, 1969. Kans. Ent. Soc., Jour. 42: 494-500, figs. 9-12 (nest, prey transport,
egg).

texanus (Ashmead). Tex., Ariz. Ecology: Nests in loose sand. Prey: Steatoda fulva Keys.
subadult, Latrodectus mactans F. juvenile.
Miscophinus tecanus Ashmead, 1898. Ent. News 9: 189. 9.

Biology: Cazier and Mortenson, 1965. Pan-Pacific Ent. 41: 26-28 (nest, prey transport; the
wasp was provisionally identified as texans).

timberlakei (Bridwell). Calif. (Mt. San Jacinto).
Hypomiscophus timberlakei Bridwell, 1920. Hawaii. Ent. Soc., Proc. 4: 394. °@.

1640 Hymenoptera in America North of Mexico
Genus NITELA Latreille

Genus NITELA Subgenus NITELA Latreille

Nitela Latreille, 1809. Gen. Crust. Ins., v. 4, p. 77.
Type-species: Nitela Spinolae Latreille. Monotypic.

Rhinonitela Williams, 1928. Hawaii Sugar Planters’ Assoc. Expt. Sta., Bul. Ent. Ser. 19: 97.
Type-species: Rhinonitela domestica Williams. Orig. desig.

Only the typical subgenus occurs in North America. Two of our species have been recorded as
nesting in abandoned burrows of other insects in wood or twigs. There are no prey records for
American species; several Palaearctic species have been reported to prey upon Psocoptera,
Aphididae and Psyllidae.

Taxonomy: Pate, 1937. Brooklyn Ent. Soc., Bul. 32: 5-7 (key to N. Amer. spp.).
cerasicola Pate. N. Y. (Long Isl.). Ecology: Reared from burrow in dead cherry tree.
Nitela cerasicola Pate, 1937. Brooklyn Ent. Soc., Bul. 32: 5. @.
floridana Pate. Fla.
Nitela floridana Pate, 1934. Ent. News 45: 241. 9.
leoni Krombein. Fla.
Nitela leoni Krombein, 1968. Nat. Canad. 95: 700. 2, ¢.
townesorum Krombein. Calif. (Yosemite Park).
Nitela townesorum Krombein, 1950. Pan-Pacific Ent. 26: 130. °.
virginiensis Rohwer. N. Y. to Fla., W. Va., Mich., Wis., Miss. Ecology: Nests in twigs of Rhus
labra.
Nitela virginiensis Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 100. °.

Taxonomy: Krombein, 1958. Ent. Soc. Wash., Proe. 60: 60. ¢.
Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 945 (nest).

SUBFAMILY TRYPOXYLONINAE

So far as known all members of this subfamily prey upon small spiders, storing rather large
numbers per cell. A few species nest in the ground, utilizing pre-existing cavities or burrows of
other arthropods. A number of species build free mud cells; our familiar pipe-organ wasp,
Trypargilum politwm, is the only North American representative having this habit. The majori-
ty of species nest above ground in cavities of various kinds such as abandoned beetle borings in
twigs, logs or structural timber, old mud-dauber nests and hollow stems.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 89-99, figs. 36-68 (larvae). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 157-161, figs. 64-70 (larvae).

Genus PISONOPSIS Fox

Pisonopsis Fox, 1893. Psyche 6: 553.
Type-species: Pisonopsis clypeata Fox. Monotypic.

The North American birkmanni makes a linear series of cells in borings in stems. P. clypeata
apparently usually makes a similar series of cells in pre-existing burrows in the soil, but it has
also been reported as nesting in trap stems at ground level.

Revision: Williams, 1954. Pan-Pacific Ent. 30: 235-246 (N. Amer. spp.).
birkmanni Rohwer. Tex. to south. Calif.; Mexico. Ecology: Nests in borings in Sambucus, in
stems of oats, white sage, poison hemlock, and in trap stems. Parasite: Ceratochrysis
antyga Boh.; Photopsis sp. Prey: Thomisidae spp.
Pisonopsis birkmanni Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 129. 2.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 157, figs. 64-70 (larva).

Biology: Williams, 1954. Pan-Pacific Ent. 30: 236, 238 (nest, prey, cocoon). —Parker and
Bohart, 1966. Pan-Pacific Ent. 42: 94-95 (nest, parasite). —Parker and Bohart, 1968.
Pan-Pacific Ent. 44: 3 (nest, parasite).

Superfamily SPHECOIDEA 1641

clypeata clypeata Fox. Wyo., Nev., Calif. Ecology: Nests in pre-existing burrows in soil and in
trap stems, making a linear series of cells, provides 9-17 prey per cell. Parasite:
Sarcophagidae sp. Prey: Chrysso nordica (Chamb. and Ivie), Theridion rabuni (Chamb.
and Ivie), T.-petraeum (Koch); Singa sp.; most of prey were adult females, but one
juvenile was stored.
Pisonopsis clypeata Fox, 1893. Psyche 6: 553. 2, 3.

Biology: Parker and Bohart, 1968. Pan-Pacifie Ent. 44: 3 (nest, parasite). —Evans, 1969.
Kans. Ent. Soce., Jour. 42: 118-121, figs. 1-4 (nest, prey transport).
clypeata occidentalis Williams. Calif. Ecology: Nests in ground in Diadasia burrows.
Pisonopsis clypeata occidentalis Williams, 1954. Pan-Pacific Ent. 30: 242, figs. 1-5, 8, 10,
15 e20 Noe coco coe Ge

Biology: Linsley, MacSwain and Smith, 1952. Calif. Univ. Pubs. Ent. 9: 274 (nest).
triangularis californica Williams. Calif.
Pisonopsis triangularis californica Williams, 1954. Pan-Pacific Ent. 30: 245, figs. 6, 9, 11,
18, 16-18, 24, 31. 2, ¢.
triangularis triangularis Ashmead. Colo., Wyo., Idaho, Calif.
Pisonopsis triangularis Ashmead, 1899. Ent. News 10: 9. 2.

Genus PISON Jurine
Genus PISON Subgenus PISON Jurine

Pison Jurine, 1808. Jn Spinola, Insectorum Liguriae, v. 2, p. 255.
Type-species: Pison Jurini Spinola. Monotypic.
Tachybulus Latreille, 1809. Gen. Crust. Ins., v. 4, p. 75.
Type-species: Tachybulus niger Latreille. Monotypic.
Nephridia Brulle, 1833. Soc. Ent. France, Ann. 2: 408.
Type-species: Nephridia Xanthopus Brulle. Monotypic.
Pison subg. Pisonitus Shuckard, 1838. Ent. Soc. London, Trans. 2: 79.
Type-species: Pison (Pisonitus) argentatus Shuckard. Desig. by Pate, 1937.
Pseudo-Nysson Radoszkowski, 1876. Soc. Ent. Rossica, Horae 12: 104.
Type-species: Pseudo-Nysson fasciatus Radoszkowski. Monotypic.
Taranga Kirby, 1883. Ent. Soe. London, Trans., p. 201.
Type-species: Taranga dubia Kirby. Monotypic.
Pisum Agassiz, 1847. Nomencl. Zool., fase. 12, p. 293. Emend. Preoce.
Pisum Schulz, 1906. Spolia Hym., p. 212. Emend. Preoce.

It is not at all certain that the single species of typical Pison described from North America
was correctly labeled as to locality. It has never been collected since in Georgia. In habitus the
unique holotype is very reminiscent of some of the glossy black Micronesian and Melanesian spe-
cies which suggests that perhaps it actually came from New Georgia in the Solomon Islands.

laeve Smith. Ga.
Pison laevis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 317. “2” = 6.

Genus PISON Subgenus KROMBEINIELLUM Richards

Paraceramius Radoszkowski, 1887. Soc. Ent. Rossica, Horae 21: 432. Preoce.
Type-species: Paraceramius Koreensis Radoszkowski. Monotypic.
Pison subg. Krombeiniellum Richards, 1962. A Revisional Study of the Masarid Wasps, p.
118. N. name.

Taxonomy: Menke, 1968. Canad. Ent. 100: 1100-1107, 15 figs. (review of New World spp.).
agile (Smith) Md., Va., Ill, Mich., Kans.; Japan, Korea, China, India, Sri Lanka. Ecology: Makes
delicate mud cells in cracks, small depressions, old Sceliphron nests; cells are placed side
by side, end to end, or in a clump; stores 20-31 small spiders per cell. Adventive after
World War II, probably from Japan. Parasite: Melittobia chalybii Ashm. Prey: Dictyna
bellans Chamb., D. sublata Hentz, D. sp.; both adults and juveniles.
Parapison agilis Smith, 1869. Ent. Soe. London, Trans., p. 300. °.

1642 Hymenoptera in America North of Mexico

Paraceramius Koreensis Radoszkowski, 1887. Soc. Ent. Rossica, Horae 21: 433, figs. 1-3.
9. N. syn. (K. V. Krombein).

Taxonomy: Krombein, 1958. Ent. News 69: 166-167. —Sheldon, 1968. Psyche 75: 111-114, figs.
5-10 (egg, larva, cocoon).

Biology: Krombein, 1958. Ent. News 69: 167 (nest, cocoon). —Sheldon, 1968. Psyche 75:
107-111, figs. 1-4 (nest, prey transport).

Genus TRYPOXYLON Latreille

Trypoxylon Latreille, 1796. Precis Caract. Gen. Ins., p. 121. No species.

Type-species: Trypoxylon figulus Linnaeus. First included species.
Tripoxylon Spinola, 1806. Insectorum Liguriae, v. 1, p. 65. Lapsus or emend.

Apius Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 106.

Type-species: Sphex figulus Linnaeus. Monotypic.

Apius Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 140. Preoce.

Type-species: Sphex figulus Fabricius. Desig. by Morice and Durrant, 1915.
Trypoxilon Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., pp. 2, 141. Lapsus or emend.
Trypoxylum Agassiz, 1847. Nomencl. Zool., p. 380. Emend.

Trypoxylum Schulz, 1906. Spolia Hym., p. 212. Emend. Preoce.
Trypoxylon subg. Asaconoton Arnold, 1959. South. Rhodesia Natl. Mus., Occas. Papers, no.
23, B, p. 322.
Type-species: Trypoxylon (Asaconoton) egregium Arnold. Orig. desig.

Sandhouse (1940) is the most reliable source for identification of North American species for
Richards (1934) does not include all of our species. The revisions cited below include the species
of both Trypoxylon and Trypargilum which are considered herein to be separate genera.

Most species of Trypoxylon nest in pre-existing cavities such as hollow stems or twigs, aban-
doned beetle borings in dead wood or structural timber, or, rarely, in soil. The Species Group
Fabricator is unusual in that some species build free mud cells whereas others nest in
pre-existing cavities in wood or soil. The preferred prey of the North American species are
small spiders, usually immatures, belonging to several families of snare-builders, but errant
spiders are used occasionally; as few as 4 and as many as 20 spiders may be stored in a single
cell. Trypoxylon males do not participate in some of the nesting activities as do those belonging
to Trypargilum; there is one report of a male usually being present in the nest of an extralimital
species of the Fabricator Group, but this needs confirmation. The cocoons of our North Amer-
ican species are delicate silken structures except in johnsoni which constructs a brittle cocoon
incorporating sand from the cell partition.

Revision: Fox, 1891. Amer. Ent. Soc., Trans. 18: 136-148, 1 pl. (N. Amer. spp.). —Fox, 1893.
Acad. Nat. Sci. Phila., Proc. 45: 472-474 (revised key to N. Amer. spp.). —Richards, 1934.
Roy. Ent. Soe. London, Trans. 82: 173-362, 56 text figs., 5 pls. (New World spp.).
—Sandhouse, 1940. Amer. Midland Nat. 24: 133-176, 4 pls. (N. Amer. spp.).

SPECIES GROUP FIGULUS
Members of Species Group Fabricator are included here.

aldrichi Sandhouse. Alta., Mont. and Wyo. west to B. C. and north. Calif. Ecology: Nests in
borings in trap stems. Predator: Philanthus zebratus nitens (Bks.).
Trypoxylon (Trypoxylon) aldrichi Sandhouse, 1940 Amer. Midland Nat. 24: 158, figs. 25,
538, 62, 66, 67. 2, d.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 95, figs. 56-58 (larva).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 95 (nest).

bidentatum Fox. Wash. and Idaho south to Calif. and Ariz. Ecology: Nests in borings in
Sambucus and in trap stems. Parasite: Trichrysis doriae (Grib.).
Trypoxylon bidentatum Fox, 1891. Amer. Ent. Soc., Trans. 18: 148. 2, ¢ (? in part).
Trypoxylon morrisoni Richards, 1934. Roy. Ent. Soc. London, Trans. 82: 319. °.

Biology: Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 95 (nest, parasite).

Superfamily SPHECOIDEA 1643

clarkei Krombein. Mass. to Fla., Tenn., [ll., Mo., Tex. Ecology: Nests in trap-nests, stores 4-6
prey per cell. Prey: Mangora gibberosa Hentz.
Trypoxylon (Trypoxylon) clarkei Krombein, 1962. Biol. Soc. Wash., Proc. 75: 9. 3, 9.

Biology: Krombein, 1967 Trap-nesting wasps and bees, pp. 229-230 (nest, prey, life cycle).
fastigium Fox. D. C., Ga., Miss., Tex., Okla., Mo., Ark., Kans., Nebr., Idaho, Utah, Nev., Ariz.,
Calif. Parasite: Trichrysis doriae (Grib.).
Trypoxylon carinifrons Fox, 1891. Amer. Ent. Soc., Trans. 18: 142. 2, ¢. Preoce.
Trypoxylon fastigium Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc. 45: 476. N. name.
Trypoxylon subfrigidum Rohwer, 1909. Ent. News 20: 378. 6.
Trypoxylon nigrellum Rohwer, 1909. Ent. News 20: 379. 9.
figulus figulus (Linnaeus). Que., Maine, N. H., Mass.; Europe. Ecology: In Europe it nests in
hollow stems or beetle borings and preys mostly upon Araneidae. Possibly adventive in
North America. Parasite: Perithous divinator (Rossi). Other subspp. occur in the
Palaearctic Region.
Sphex figulus Linnaeus, 1758. Syst. Nat., ed. 10, p. 570.
Sphex fuliginosa Seopoli, 1763. Ent. Carn., p. 292.
Trypoxylon figulum var. major Kohl, 1883. Schweiz. Ent. Ges., Mitt. 6: 657. 6.
Trypoxylon apicalis Fox, 1891. Amer. Ent. Soc., Trans. 18: 142, fig. 10. 2.
Trypoxylon figulwm var. minor de Beaumont, 1945. Schweiz. Ent. Ges., Mitt. 19: 478. 2,
3.

Taxonomy: Giordani Soika, 1934. Soc. Ent. France, Ann. 103: 342-343, pl. 3, fig. 2 (larva).
—Pate, 1943. Brooklyn Ent. Soc., Bul. 38: 46 (synonymy of apicale).
frigidum frigidum Smith. Hudson Bay south to N. C., west to Wash. and N. Mex. Ecology:

Nests in hollow stems and twigs, in abandoned beetle borings in dead wood and
structural timber, and in trap-nests, stores 4-16 prey per cell. Parasite: Pyemotes
ventricosus (Newp.); Megaselia sp.; Anthrax sp.; Amobia distorta (Allen); Melittobia
chalybii Ashm.; Cleonymidae sp., possibly Ptinobius magnificus (Ashm.); Trichrysis
doriae (Grib.). Prey: Achaearanea globosa (Hentz), Thymoites unimaculata
unimaculata (Em.), Theridion albidum Bks., T. differens Em., T. frondeuwm Hentz ?, T.
globoswm Hentz, T. lyra Hentz, T. murarium Em., T. sp. in Murarium Group, T.
unimaculatum Em.; Salticidae sp.; Hustala anastera (Walck.), E. sp., Araneidae spp.;
Tennesseellum formicum (Em.); Leucauge venusta (Walck.), Tetragnatha sp.;
Ceratinopsis interpres Camb., C. purpurescens Keys., Micryphantidae sp.; both adults
and immatures are stored and most are snare-building species. Other subspp. occur in
the Palaearctic Region.

Trypoxylon frigidum Smith, 1856. Cat. Hym. Brit Mus., v. 4, p. 381. 2.

Trypoxylon plesium Rohwer, 1920. U.S. Natl. Mus., Proc. 57: 229. 9.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 96, figs. 59-60 (larva).

Biology: Packard, 1867. Ent. Soc. Phila., Proc. 6: 415 (nest). —Rau, 1922. Acad. Sci. St. Louis,
Trans. 24: 22. —Blackman and Stage, 1924. N. Y. State Col. Forestry, Syracuse Univ.,
Tech. Pub. 17: 197 (nest). —Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 197 (nest). —Rau,
1928. Acad. Sci. St. Louis, Trans. 25: 439-441, figs. 65, 66 (nest, parasites). —Taylor, 1928.
Psyche 35: 225 (nest). —Pate, 1937. Brooklyn Ent. Soc., Bul. 22: 5 (nest). —Thomas, 1962.
Amer. Midland Nat. 67: 365 (nest, parasite). —Thomas, 1963. Mich. Acad. Sci., Arts,
Letters, Papers 48: 127-130 (nest, life cycle, parasite). —Parker and Bohart, 1966.
Pan-Pacific Ent. 42: 95 (nest). —Krombein, 1967. Trap-nesting wasps and bees, pp. 223-227,
fig. 11 (nest, prey, life cycle, parasites). —Medler, 1967. Amer. Midland Nat. 78: 344-358
(nest, prey, cocoon, parasites, life cycle).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 21, figs. M-P (male
genitalia).
johnsoni Fox. Ont., N. Y. to Fla. west to Mich., Mo., Okla., Tex. Ecology: Nests in hollow

stems or borings in wood, and possibly in pre-existing burrows in earth. Prey:
Tetragnatha spp.; Micrathena gracilis (Walck.), Araneus sp.?; prey consisted of
immatures and penultimate stages of both sexes.

Trypoxylon Johnsoni Fox, 1891. Amer. Ent. Soc., Trans. 18: 147. @.

Trypoxylon ornatipes Fox, 1891. Amer. Ent. Soc., Trans. 18: 148. 6.

1644 Hymenoptera in America North of Mexico

Trypoxylon (Trypoxylon) adelphiae Sandhouse, 1940. Amer. Midland Nat. 24: 151. 9°, d.
Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 97, fig. 61 (larva).

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 137-139 (nest, prey). —Rau, 1922.
Acad. Sci. St. Louis, Trans. 24: 22 (nest ?). —Krombein, 1952. Amer. Ent. Soc., Trans. 78:
93 (prey). —Krombein, 1967. Trap-nesting wasps and bees, pp. 229-231 (nest, prey, life
cycle, cocoon).

kolazyi Kohl. N. Y. to Ga., W. Va., Ill., Mo.; Austria, Mediterranean area. Ecology: Nests in
abandoned anobiid borings in structural lumber and in trap-nests. Adventive from
Europe. Parasite: Trichrysis doriae (Grib.). Prey: Tennesseellum formicum (Em.);
Micryphantidae sp.; adults and subadults.
Trypoxylon Kolazyi Kohl, 1893. Zool.-Bot. Gesell. Wien, Verhandl. 43: 29. d, 2.
Trypoxylon (Trypoxylon) backi Sandhouse, 1940. Amer. Midland Nat. 24: 164, figs. 18, 74,
75. 2, 6. N. syn. (R. E. Coville).

Biology: Sandhouse, 1940. Amer. Midland Nat. 24: 165 (nest). —Krombein, 1958. Biol. Soc.
Wash., Proc. 71: 21-22 (nest, prey). —Thomas, 1962. Amer. Midland Nat. 67: 364 (parasite).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 227-228 (nest, prey, life cycle).

pennsylvanicum pennsylvanicum Saussure. Que. and Maine to Fla. west to Colo. and Tex.
Ecology: Nests in cavity in twig. Prey: Araneidae sp. immature. Another subsp. occurs
in Japan.
Trypoxylon pennsylvanicum Saussure, 1867. Reise d. Novara, Zool. 2, Hym., p. 82. @.

Biology: Sandhouse, 1940. Amer. Midland Nat. 24: 160 (nest). —Krombein, 1961. Brooklyn
Ent. Soc., Bul. 56: 64 (prey).
regulare Viereck. Mo., Kans.
Trypoxylon regularis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 205. “2” = 6.
sculleni Sandhouse. Mont. to Ariz. west to B. C. and Calif. Ecology: Nests in cavities in twigs,
Sambucus, Rhus glabra, and Eriogonum, and in trap stems. Parasite: Anthrax irroratus
Say; Trichrysis doriae (Grib.).
Trypoxylon (Trypoxylon) sculleni Sandhouse, 1940. Amer. Midland Nat. 24: 160, figs. 15,
52, 58, 59, 65. 2, 3.

Biology: Sandhouse, 1940. Amer. Midland Nat. 24: 162 (nest). —Parker and Bohart, 1966.
Pan-Pacifie Ent. 42: 95 (nest, parasites).

SPECIES GROUP RUFIDENS

bridwelli Sandhouse. Tex. (Brownsville).
Trypoxylon (Trypoxylon) bridwelli Sandhouse, 1940. Amer. Midland Nat. 24: 168, figs. 24,
40. 6.
richardsi Sandhouse. Ont., N. Y. to Fla., west to Mich., Ind., Mo., Ala., Tex. Ecology: Nests in
cavity in twig of Chionanthus virginiana and in twig gall on oak. Prey: Small spiders.
Trypoxylon (Trypoxylon) richardsi Sandhouse, 1940. Amer. Midland Nat. 24: 167, figs. 22,
AD AOMORI

Biology: Sandhouse, 1940. Amer. Midland Nat. 24: 168 (nest). —Krombein, 1959. Biol. Soc.
Wash., Proc. 72: 101-102 (nest, prey, life cycle).
timberlakei Sandhouse. Ariz., Calif. Ecology: Nests in deserted gall of Callirhytis hamiformis
(Bass.) on Quercus wislizenii.
Trypoxylon (Trypoxylon) timberlakei Sandhouse, 1940. Amer. Midland Nat. 24: 169. °.
Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 956 (nest).
SPECIES GROUP SCUTATUM

carinatum Say. U.S. east of 100th meridian north to Mass. Ecology: Reared from burrow in
partly decayed trunk of tulip-tree, also nests in borings in wood. Prey: Theridion
lyricum Walck.
Trypoxylon carinatus Say, 1837. Boston Jour. Nat. Hist. 1: 374. ¢.

Biology: Sandhouse, 1940. Amer. Midland Nat. 24: 154 (nest). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 228-229 (nest, prey, life cycle, cocoon).

Superfamily SPHECOIDEA 1645

SPECIES GROUP MARGINATUM

punctivertex Richards. Okla., Tex. south to Brazil.
Trypoxylon (Trypoxylon) punctivertex Richards, 1934. Roy. Ent. Soc. London, Trans. 82:
333. 2.

Genus TRYPARGILUM Richards

Trypoxylon subg. Trypargilum Richards, 1934. Roy. Ent. Soc. London, Trans. 82: 191.
Type-species: Trypoxylon nitidum Smith. Orig. desig.

Sandhouse (1940) is the most reliable source for identification of North American species for
Richards (1934) does not include all of our species. The revisions cited below include the species
of both Trypargilum and Trypoxylon which are considered herein to be separate genera.

Some aspects of the ethology are unique among wasps. This genus and some species of Pison
are the only wasps in which males are known to assist in some of the nesting activities. They
remain in the nest while the female is hunting for prey and discourage attack by at least some of
the parasites that afflict wasps. Activities in which the males may participate include cleaning
out a pre-existing boring which is to serve as a nesting site, taking prey from the female and
placing it in the cell, and helping the female seal inner partitions of cells with mud which she
brings to the nest. The larvae also exhibit unusual behavior in that the cocoon which incor-
porates silk, other salivary secretions and mud from the cell partition is specifically different in
each of the North American species which have been observed.

North American species have been reported to store 3-36 small, usually immature spiders
per cell. Some species use only snare-building spiders as prey, one uses predominantly
snare-building spiders but does include some errant spiders, and others use predominantly er-
rant types but with a number of snare-builders. This suggests that there are specific differences
in the way that groups of species hunt for prey.

In North America members of the Spinosum, Nitidum and Punctulatum Groups nest in
pre-existing cavities such as hollow stems or twigs, old insect galls or mud dauber nests, aban-
doned beetle borings in dead wood or structural timber and in trap-nests. Our single member of
the Albitarse Group is a mud dauber and builds the familiar pipe-organ nest.

Revision: Fox, 1891. Amer. Ent. Soc., Trans. 18: 136-148, 1 pl. (N. Amer. spp.). —Fox, 1894
(1893). Acad Nat. Sci. Phila., Proc. 45: 472-474 (revised key to N. Amer. spp.). —Richards,
1934. Roy. Ent. Soc. London, Trans. 82: 173-362, 56 text figs., 5 pls. (New World spp.).
—Sandhouse, 1940. Amer. Midland Nat. 24: 133-176, 4 pls. (N. Amer. spp.).

Taxonomy: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 152-153 (key to red-marked Fla. taxa).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 178-185, text fig. 2 (male
behavior, cocoon differences, prey preferences, competition for nesting sites, differing
emergence dates).

SPECIES GROUP SPINOSUM

Members of this group prey upon both errant and snare-building spiders, but prefer the
former by a substantial margin.

bicalearatum (Richards), n. comb. South. Ariz.; Mexico.
Trypoxylon (Trypargilum) bicalcaratum Richards, 1934. Roy. Ent. Soe. London, Trans. 82:
235. 6.
californicum Saussure. Tex., N. Mex., Ariz., Utah, Nev., Calif., Oreg., Wash. Ecology: Nests in
borings in wood, stores 8-19 prey per cell. Parasite: Chrysididae sp. Prey: Agassa sp.,
Habronattus sp., Metaphidippus sp., Phidippus sp., Synemosyna sp., Salticidae spp.;
Ebo sp., Miswmenops sp., Philodromus sp., Thomisidae spp.; Oxyopes tridens Brady, O.
sp.; Clubionidae sp.; Dictyna sp.; Araneidae spp.; stores mostly errant spiders, only a
few snare-builders.
Trypoxylon californicum Saussure, 1867. Reise d. Novara, Zool. 2, Hym., p. 78. 9.
Trypoxylon arizonense Fox, 1891. Amer. Ent. Soc., Trans. 18: 145. 2. N. syn. (R. E.
Coville).

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 92, figs. 47-48 (larva).
Biology: Matthews and Matthews, 1968. Psyche 75: 285-293, 2 figs. (nest, prey, cocoon).

1646 Hymenoptera in America North of Mexico

clavatum clavatum (Say), n. status (R. E. Coville). Ont., U. S. east of Rockies except New

England, Ariz. Ecology: Nests in borings in wood, old mud dauber and Polistes nests,
and old mining bee burrows; stores 5-22 spiders per cell. Parasite: Pyemotes ventricosus
(Newp.); Anthrax aterrima (Big.); Megaselia sp. ?; Amobia distorta (Allen),
Miltogramminae spp.; Melittobia chalybii Ashm.; Sphaeropthalma p. pensylvanica
(Lep.), S. p. scaeva (Bl); Trichrysis carinata (Say). Prey: Dictyna sublata (Hentz);
Clubiona sp., Clubionidae sp.; Anyphaena pectorosa Koch, A. sp., Anyphaenella
saltabunda (Hentz), Anyphaenidae spp.; Misuwmenops asperatus (Hentz), M. oblongus
(Keys.), M. spp., Miswmenoides aleatorius (Hentz), Synema parvula (Hentz), Xysticus
triguttatus Keys., X. spp., Philodromus rufus Walck., P. marxii Keys., P. pernix
Blackw. P. satullus Keys., P. washita Bks., P. spp., Thanatus formicinus (Oliv.), T.
striatus Koch, Misumeninae spp.; Salticus scenicus (L.), Evarcha hoyi Peckh., Onondaga
lineata (Koch), Habronattus sp., Phidippus audax (Hentz), P. clarus Keys., P. spp.,
Paraphidippus marginatus (Walck.), P. spp., Zygoballus bettini Peckh., Z. nervosus
(Peckh.), Z. sexpunctatus (Hentz), Thiodina iniquies (Walck.), T. puerpera (Hentz),
Hentzia mitrata (Hentz), Maevia vittata (Hentz), Icius elegans (Hentz), I. hartii Em.,
Metaphidippus insignis (Bks.), M. protervus (Walck.), M. galathea (Walck.), Salticidae
spp.; Dapanus mira (Walck.); Pardosa sp., Lycosidae spp.; Oxyopes salticus Hentz,
Oxyopidae spp.; Argiope aurantia Luc., A. trifasciata (Forsk.), Mangora gibberosa
(Hentz), M. maculata (Keys.), Eustala anastera (Walck.), E. sp., Neoscona minima
Camb., N. domiciliorwm (Hentz), N. sp., Araneus juniperi (Em.), A. spp., Araniella
displicata (Hentz), Araneidae spp.; Theridiidae spp.; Tetragnatha sp.; prefers errant to
snare-building spiders by ratio of 4: 1. Predator: Cymatodera sp.?, Lecontella cancellata
(LeC.); Trogoderma ornatum Say.

Trypoxylon clavatus Say, 1837. Boston Jour. Nat. Hist. 1: 374.

Trypoxylon annulare Dahlbom, 1844. Hym. Europaea, v. 1, pp. 282, 509. °.

Trypoxylon rufozonalis Fox, 1891. Amer. Ent. Soc., Trans. 18: 145. 2, d. N. syn. (R. E.
Coville).

Trypoxylon quintilis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 206. d.

Trypoxylon cockerellae Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 130. °.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 91, figs. 36-43 (larva). —Evans, 1959.
Amer. Ent. Soc., Trans. 85 160 (larva).

Biology: Ashmead, 1894. Psyche 1: 45 (nest). —Rau and Rau, 1916. Jour. Anim. Behavior 6:
34, figs. 16, 19, 24, 25 (nest). —Rau and Rau, 1918. Wasp Studies Afield, pp. 135-137, fig. 32
(nest). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 22 (nest). —Blackman and Stage, 1924
N. Y. State Col. Forestry, Syracuse Univ., Tech. Pub. 17: 196 (nest). —Rau, 1926. Acad.
Sci. St. Louis, Trans. 25: 198 (nest). —Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 406 (prey,
life cycle). —Rau, 1931. Ent. News 42: 200 (homing experiments). —Muma and Jeffers,
1945. Ent. Soc. Amer., Ann. 38: 246, 252, 255, pl. 2, figs. 5, 6 (nest, prey). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 203-210, text fig. 2b, figs. 11, 133 (nest, prey, life cycle,
egg, cocoon, parasites, predators).

clavatum johannis (Richards), n. status (R. E. Coville). Ga., Fla. Ecology: Nests in borings in
wood, stores 8-17 spiders per cell. Parasite: Amobia floridensis (Tns.), Miltogrammini sp.
Prey: Lyssomanes viridans (Hentz), L. viridis (Walck.); Misumenops bellulus (Bks.), M.
celer (Hentz), M. sp., Tmarus sp., Tibellus sp., Thomisidae sp.; Pellenes sp.,
Paraphidippus marginatus (Walck.), Phidippus audax (Hentz), P. variegatus Luc., P.
clarus Keys., P. sp., Icius sp., Hentzia ambigua (Walck.), H. palmarum Hentz, H. sp.,
Maevia hobbsi Barnes, M. michelsoni Barnes, Thiodina sylvana Hentz, T. pseustes
Chamb. and Ivie; Mimetus notius Chamb.; Pardosa sp.; Oxyopes salticus Hentz, O. sp.,
Peucetia abboti (Walck.); Gea heptagon (Hentz), Drexelia directa (Hentz), Eustala
anastera (Walck.), Mangora placida (Hentz), Wagneriana tauricornis Chamb., Neoscona
arabesca (Walck.), N. minima (Keys.), N. sp., Argiope aurantia Luc., A. trifasciata
(Forsk.), Araneidae sp.; Lewcauge venusta (Walck.); prefers errant to snare-building
spiders by a ratio of 3:1.

Trypoxylon (Trypargilum) johannis Richards, 1934. Roy. Ent. Soc. London, Trans. 82: 238.

é.

’

Superfamily SPHECOIDEA 1647

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 92, figs. 44-46 (larva). —E vans, 1959.
Amer. Ent. Soc., Trans. 85: 160 (larva).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 210-214, text fig. 2c, fig. 52 (nest,
prey, life cycle, egg, cocoon, parasites).
saussurei (Rohwer), n. comb. South. Ariz. south to Guanacaste Prov., Costa Rica. Ecology:
Nests in old Sceliphron cells.
Trypoxylon Mexicanum Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 78, pl. 4, fig.
45. 9, 6. Preoce.
Trypoxylon saussurei Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 478. N. name.

Biology: Rau, 1943. Ent. Soc. Amer., Ann. 36: 649 (nest).

spinosum (Cameron). Tex. to Panama. Ecology: Nests in holes in masonry, crevices along tree

trunk and in bamboo stems; stores 11-20 spiders per cell: Parasite: Sarcophagidae sp.
Prey: Aysha decepta Bks., A. gracilis (Hentz), Anyphaena sp.; Gea ergaster (Walck.),
Neoscona sp.; Mimetus notius Chamb.; Peucetia viridans (Hentz); Hentzia palmarum
(Hentz), Metaphidippus sp., Paraphidippus marginatus (Walck.); Misumenops oblongus
(Keys.); Uloborus glomosus Walck.; both adults and immatures are stored.

Trypoxylon spinosum Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 46. 2, d.

Trypoxylon cinereo-hirtum Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 44. 2.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 93, fig. 49 (larva).

Biology: Lin, 1969. Wasmann Jour. Biol. 27: 125-128 (nest, prey hunt, life cycle, egg, cocoon).
texense (Saussure). Ga., Fla., Ala. to Ill. west to N. Mex., Colo., S. Dak. Ecology: Nests in

abandoned mud-dauber cells, old burrows of other arthropods in sand banks, and
crevices in wooden or stone wall; stores 9-15 spiders per cell. Parasite: Anthrax
limatulus artemisia Marst. Prey: Oxyopes salticus Hentz, Peucetia viridis (Walck.);
Neoscona arabesca (Walck.), Gea ergaster (Walck.), Argiope trifasciata (Forsk.), A. sp.,
Eustala cepina (Walck), Metepeira sp., Araneus cornutus Clerck; Tetragnatha
pallescens Camb., T. versicolor Walck.; Misumenops delphinus (Walck.), M. oblongus
(Keys.), Tibellus duttoni (Hentz), Xysticus sp., Philodromus sp., Misumena sp.;
Zygoballus nervosus Peckh., Habronattus brunneus (Peckh.), Hentzia ambigua (Walck.),
Marpissa pikei (Peckh.), Metaphidippus galathea (Walck.), Paraphidippus marginatus
(Walck.), Phidippus audax Hentz, P. rimator (Walck.); Pardosa distincta (Blackw.);
Mimetus sp.; Theridion intritum Bish. and Cros., T. murarium Em.; Rucinia sp.;
Dendryphantes sp.; stores both immatures and adults, and about equal numbers of
errant and snare-building species.

Typoxylon texense Saussure, 1867. Reise d. Novara, Zool. 2, Hym., p. 77. °.

Trypoxylon sulcus La Munyon, 1877. Nebr. Assoc. Adv. Sci., Proc., March 8.

Trypoxylon aureolum Rohwer, 1909. Ent. News 20: 381. 9, 6.

Trypoxylon relativum Rohwer, 1909. Ent. News 20: 382. °.

Taxonomy: Evans, 1959. Amer. Ent. Soe., Trans. 85: 159 (larva).

Biology: Hartman, 1905. Tex. Acad. Sci., Trans. 7: 71-78, pl. 4, fig. 23 (nest, prey).
—Hungerford and Williams, 1912. Ent. News 23: 248, fig. 5 (nest, prey). —Rau, 1928.
Acad. Sci. St. Louis, Trans. 25: 441 (nest). —Rau, 1940. Ent. Soc. Amer., Ann. 33: 592
(nest). —Evans, 1959. Amer. Ent. Soc., Trans. 85: 159 (nest). —Kurezewski, 1963. Fla. Ent.
46: 243-245 (nest, prey, egg). —Lin, 1969. Wasmann Jour. Biol. 27: 128-129 (nest, prey).

xantianum (Saussure), n. comb. South. Calif.; Mexico (Baja California).
Trypoxylon xantianum Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 78. d, °.

SPECIES GROUP PUNCTULATUM

lactitarse (Saussure). Ont. and Mass. to Fla., west to Wis., Tex. and Ariz., south to Argentina.
Ecology: Nests in borings of other insects in dead wood and structural timber, in trap
nests, in old mud dauber nests, and in old burrows of mining bees. Parasite: Amobia
aurifrons (Tns.), A. distorta (Allen), Senotainia sp. in trilineata (Wulp) complex;
Anthrax a. argyropygus Wied., A. aterrimus (Big.); Megaselia sp.; Tyrophagus sp.;
Melittobia chalybii Ashm.; Trichrysis carinata (Say); Sphaeropthalma pensylvanica

1648 | Hymenoptera in America North of Mexico

scaeva (Bl.). Prey: Clubiona obesa Hentz, C. pallens Hentz; Aysha gracilis (Hentz),
Anyphaena celer (Hentz), A. fraterna (Bks.), A. pectorosa Koch; Philodromus washita
Bks., P. infuscatus Keys., P. praelustris Keys., P. rufus (Walck.), P. pernix Blackw., P.
spp.; Salticidae sp.; Pisaurina mira (Walck.), P. sp.; Mimetus puritanus Chamb.; Wixia
ectypa (Walck.), Eustala anastera (Walck.), E. emertoni Bks., Acacesia hamata (Hentz),
Mangora gibberosa (Hentz), M. maculata (Keys.), Neoscona arabesca (Walck.), N.
domiciliorum (Hentz), N. minima Camb., N. spp., Araneus juniperi (Em.), A.
patagiatus Clerck, A. marmoreus Clerck, A. spp., Araniella displicata Hentz,
Neosconella pegnia (Walck.), Argiope trifasciata (Forsk.), Conepeira glyphica Archer,
Araneidae spp.; Theridion differens (Em.), T. murarium (Em.); Poecilochroa capulata
(Walck.); snare-building spiders are preferred to errant by ratio of 13: 1. Predator:
Trogoderma ornatum Say; Lecontella cancellata (LeC.).

Trypoxylon lactitarse Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 81. ¢.

Trypoxylon striatum Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 283. 9.

Trypoxylon cinereum Cameron, 1889. Biol. Cent.-Amer., Hym., v. 2, p. 40. 2, d.

Trypoxylon albopilosum Fox, 1891. Amer. Ent. Soc., Trans. 18: 139. ?, d.

Trypoxylon albopilosum planoense Rohwer, 1909. Ent. News 20: 380. °.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 94, fig. 55 (larva). —Evans, 1959. Amer.
Ent. Soc., Trans. 85: 161 (larva). —Menke, 1974. Ent. Soc. Wash., Proc. 76: 418 (identity of
lactitarse Sauss.).

Biology: Peckham and Peckham, 1895. Psyche 7: 303 (nest, prey, male behavior in nest).
—Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 77-87 (nest, prey,
male behavior in nest). —Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 199 (nest). —Rau,
1928. Acad. Sci. St. Louis, Trans. 25: 423 (nest, prey). —Krombein, 1956. Ent. Soc. Wash.,
Proc. 58: 155-156 (nest, prey). —Balduf, 1961. Brooklyn Ent. Soc., Bul. 56: 82-83 (nest).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 214-222, text fig. 2e, figs. 53-56, 132,
134, 135 (nest, prey, life cycle, egg, cocoon, parasites, predator). —Medler, 1967. Amer.
Midland Nat. 78: 344-358 (nest, prey, cocoon, parasites, life cycle).

SPECIES GRouP NITIDUM
Members of this group prey entirely upon snare-building spiders.

collinum collinum (Smith). Ga., Fla. Ecology: Nests in cavities in twigs and in trap nests,
stores 13-25 spiders per cell. Parasite: Anthrax a. argyropygus Wied.; Phoridae sp.;
Trichrysis carinata (Say). Prey: Theridion flavonotatum Beck., T. glaucescens Beck.;
Eustala anastera (Walck.), E. triflex (Walck.), Acacesia folifera (Marx), A. hamata
(Hentz), Neoscona minima Camb., N. spp., Araneus juniperi (Em.), Conaranea
floridensis Bks., C. sp., Neosconella pegnia (Walck.), Araneidae spp. Predator:
Crematogaster sp.

Trypoxylon collinum Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 381. @.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 160 (larva).

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 18 (nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 185-187 (nest, prey, life cycle, cocoon, parasites,
predator). —Krombein, 1970. Smithsn. Contrib. Zool. 46: 22-26, figs 62-78 (nesting
behavior, prey, life cycle).

collinum rubrocinctum (Packard). Que. to Ga., west to Minn., Kans., Nev. Ecology: Nests in
cavities in twigs, trap nests, abandoned beetle borings, hollow straws, and in crevices in
mortar; stores 5-27 spiders per cell. Parasite: Anthrax a. argyropygus Wied., A.
aterrimus (Big.); Megaselia sp.; Miltogramminae sp.; Sphaeropthalma pensylvanica
scaeva (Bl.); Trichrysis carinata (Say), T. doriae (Grib.), Chrysis pellucidula Aar., C.
sp.; Messatoporus compressicornis Cush. Prey: Conopistha sp., Theridula opulenta
(Walck.), Theridion lyricum Walck., T. murariwm Em., T. spirale Em., T. flavonotatum
Beck., T. lyra Hentz, T. alabamense Gertsch and Archer, T. albidum (Bks.), T. differens
(Em.), T. spp., Argyrodes sp., Theridiidae spp.; Eustala anastera (Walck.), E. emertoni
Bks., E. spp., Cyclosa conica (Pall.), Neoscona arabesca (Walck.), N. spp., Araneus
attestor Petrunk., A. juniperi (Em.), A. trifolium (Hentz), A. spp., Argiope aurantia

Superfamily SPHECOIDEA 1649

Lue., A. trifasciata (Forsk.), Mangora gibberosa (Hentz), Araneidae spp.; Linyphia
clathrata Sund., L. sp.; Leucauge venusta (Walck.).
Trypoxylon rubro-cinctum Packard, 1867. Ent. Soc. Phila., Proc. 6: 416. °.

Taxonomy: Evans, 1957. Amer. Ent. soc., Trans. 83: 94, fig. 63 (larva). —Evans, 1959. Amer.
Ent. Soe., Trans. 85: 161 (larva).

Biology: Peckham and Peckham, 1895. Psyche 7: 303 (nest, prey, life cycle). —Peckham and
Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 77-84 (nest, prey, life cycle).
—Peckham and Peckham, 1905. Wasps, social and solitary, pp. 178-193, 2 figs. (nest, prey,
male behavior in nest). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 22 (nest, parasite).
—Krombein, 1954. Brooklyn Ent. Soc., Bul. 49: 5 (nest, prey). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 187-193, text fig. 2a, figs. 126, 135-139 (nest, prey, life
cycle, cocoon, parasites). —Medler, 1967. Amer. Midland Nat. 78: 344-358 (nest, prey,
cocoon, parasites, life cycle).

orizabense (Richards), n. comb. South. Ariz.; Mexico (Orizaba).
Trypoxylon (Trypargilum) orizabense Richards, 1934. Roy. Ent. Soe. London, Trans. 82:
Zhan Fi, 801:
tridentatum archboldi (Krombein). Fla. Ecology: Nests in borings in wood, stores 9-36 spiders
per cell. Parasite: Anthrax a. argyropygus Wied., Lepidophora lepidocera (Wied.) ?;
Pyemotes ventricosus (Newp.). Prey: Mimetus sp.; Gea heptagon (Hentz), Drexelia
directa (Hentz), Eustala anastera (Walck.) Larinia directa (Hentz), Singa sp., Neoscona
arabesca (Walck.), N. sp., Conaranea floridensis Bks., C. sp., Araneidae spp.; Theridula
quadripunctata Keys., Theridion flavonotatum Beck., Chrysso clementinae (Petrunk.),
Theridiidae spp.
Trypoxylon (Trypargilum) tridentatum archboldi Krombein, 1959. Ent. Soc. Wash., Proc.
61: 150. 2, 6.

Biology: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 151-152 (nest, prey, cocoon). —Krombein,
1967. Trap-nesting wasps and bees, pp. 200-202 (nest, prey, life cycle, cocoon, parasites).
tridentatum tridentatum (Packard). Conn. to Fla., west to B. C. and Calif. Ecology: Nests in
Sambucus and white sage stems, borings in wood and old mud dauber nests. Parasite:
Trichrysis mucronata (Br.), T. deversor Boh., Ceratochrysis antyga Boh., Chrysis
pellucidula Aar.; Sphaeropthalma uro (Bl.), S. amphion (Fox), S. abdominalis (B1.);
Tetrastichus sp.; Amobia floridensis (Tns.), A. spp. ?; Bombyliidae sp.; Pyemotes
ventricosus (Newp.). Prey: Latrodectus mactans (F.), Theridion murarium Em., T.
dilutum Levi, T. sp.; Mimetus hesperus Chamb.; Eustala rosae Chamb. and Ivie, E. sp.,
Metepeira arizonica Chamb. and Ivie, M. sp., Neoscona vertebrata (McCook), N. sp.,
Araneus sp., Acanthepeira stellata (Walck.), Cyclosa conica (Pall.), Araneidae spp.
Trypoxylon tridentatwm Packard, 1867. Ent. Soc. Phila., Proc. 6: 417. °.
Trypoxylon projectum Fox, 1891. Amer. Ent. Soc., Trans. 18: 141. 2, ¢.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 93, figs. 50-52 (larva). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 161 (larva).

Biology: Rau and Rau, 1918. Wasp Studies Afield, p. 134 (nest, life cycle). —Blackman and
Stage, 1924. N. Y. State Col. Forestry, Syracuse Univ., Tech. Pub. 17: 196-197 (nest).
—Rau, 1934. Canad. Ent. 66: 259 (nest, parasite). —Hicks, 1934. Colo. Univ., Studies 21:
267 (nest, parasite). —Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 95 (nest, parasites).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 193-200, text fig. 2d, figs. 115-118
(nest, prey, life cycle, egg, cocoon, parasites). —Paetzel, 1973. Pan-Pacific Ent. 49: 26-30, 3
figs. (male behavior in nest, mating; misdet. as rwbrocinctum). —Paetzel, 1973. Outdoor
World 6 (2): 42-45, 4 figs. (nest, prey, life cycle, male behavior in nest; misdet. as
rubrocinctum).

SPECIES GROUP ALBITARSE
politum (Say). Mass. to Fla. west to Kans. and Tex. Ecology: Builds linear series of mud cells
in parallel rows resembling pipes of an organ, stores 3-18 prey per cell. Parasite: Vidia

sp.; Anthrax limatulus fur (O. S.), A. aterrimius (Big.); Amobia aurifrons (Tns.), A.
distorta (Allen), Helicobia rapax Wlkr., Senotainia sp.; Sphaeropthalma p. pensylvanica

1650 Hymenoptera in America North of Mexico

(Lep.), S. p. scaeva (Bl.), Dasymutilla v. vesta (Cr.); Melittobia chalybii Ashm.;
Trichrysis tridens (Lep.). Prey: Neoscona minima Keys., N. benjamina Walck., N. spp.,
Eustala anastera (Walck.), Wixia ectypa (Walck.), Aranea spp.; Theridiidae spp.; preys
upon adults and immatures. Because of its nest this wasp is popularly known as the
pipe-organ wasp.

Trypoxylon politus Say, 1837. Boston Jour. Nat. Hist. 1: 373.

Trypoxylon neglectum Kohl, 1884. Zool.-Bot. Gesell. Wien, Verh. 33: 340. 6.

Trypoxylon basale Rohwer, 1912. U.S. Natl. Mus., Proe. 41: 475. 2, ¢.

Trypoxylon politiforme Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 476. °, d.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 95, fig. 62 (larva). —Evans, 1959. Amer.
Ent. Soc., Trans. 85: 161 (larva).

Biology: Rau, 1913. Ent. News 24: 401 (larval feeding). —Rau and Rau, 1916. Jour. Anim.
Behavior 6: 31-35, fig. 5 (nest, prey, male behavior in nest). —Rau, 1928. Acad. Sci. St.
Louis, Trans. 25: 428-439, figs. 57-62 (nest, prey, life cycle, male behavior in nest, cocoon,
parasites). —Osborne, 1929. Conn. Agr. Expt. Sta., Bul. 305: 751-753, 1 pl., 1 fig. —Dow,
1930. Brooklyn Ent. Soe., Bul. 25: 98-101 (nest). —Fattig, 1936. Canad. Ent. 68: 44 (nest,
prey). —Hartman, 1944. Ent. News 55: 7 (prey). —Rau, 1944. Ent News 55: 9 (prey hunt).
—Muma and Jeffers, 1944. Ent. News 55: 50 (nest). —Muma and Jeffers, 1945. Ent. Soc.
Amer., Ann. 38: 246, 252, 254-255, figs. 3, 4 (nest, prey). —Lin, 1969. Wasmann Jour. Biol.
27: 129-130 (prey). —Johnson, 1974. Ent. Soc. Wash., Proc. 76: 448-449 (parasite). —Cross,
Stith and Bauman, 1975. Ent. Soc. Amer., Ann. 68: 901-916, 10 figs. (mating, nest
construction and provisioning, prey hunting, cocoon, life history, parasites).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 21, figs. L, Q, R (male
genitalia).
SUBFAMILY BOTHYNOSTETHINAE
Genus BOTHYNOSTETHUS Kohl
Bothynostethus Kohl, 1884. Zool.-Bot. Gesell. Wien, Verhandl. 33: 344.

Type-species: Bothynostethus Sausswrei Kohl. Monotypic.

distinctus Fox. N. Y. to Fla. west to Idaho and Ariz. Ecology: Nests in sand, occasionally
utilizing a rodent burrow, makes up to 8 cells per nest, stores 4-9 chrysomelid beetles
per cell. Parasite: Senotainia sp. ? Prey: Monowia sp., Pyrrhalta decora (Say), P.
perplexa Fall., P. spiraeae Fall, P. cavicollis (LeC.), Ophraella notata (F.).
Bothynostethus distinctus Fox, 1891. Ent News 2: 31. @, 6.

Taxonomy: Kurezewski and Evans, 1972. Psyche 79: 97-99, figs. 8-15 (larva).
Biology: Cazier and Mortenson, 1965. Pan-Pacifie Ent. 41: 31-32 (nest, prey). —Kurcezewski

and Evans, 1972 Psyche 79: 88-97, figs. 1-7 (nest, prey transport, cocoon, parasites).
Family CRABRONIDAE

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 99-113, figs 69-115 (larvae). —Evans,
1959. Amer. Ent. Soc., Trans. 85: 162-165, figs. 43-53, 61 (larvae). —Evans, 1964. Amer.
Ent. Soc., Trans. 90: 289-296, figs. 104-114 (larvae).

SUBFAMILY OX YBELINAE

So far as known all species nest in the ground making uni- or multicellular nests and prefer-
‘ing rather friable soil.
Taxonomy: Pate, 1940. Amer. Ent. Soc., Trans. 66: 3-5 (group characters and key to world
genera).
Genus BELOMICRUS Costa

Belomicrus Costa, 1871 (1866). R. Univ. Napoli Mus. Zool., Ann. 6: 80.
Type-species: Belomicrus italicus Costa. Monotypic.

Superfamily SPHECOIDEA 1651

Oxybeloides Radoszkowski, 1877. In Fedtschenko, Reise in Turkestan, Imp. Obschch.
Moskva Izv. 26 (1): 68.
Type-species: Oxybeloides fasciatus Radoszkowski. Monotypic.
Oxybelomorpha Brauns, 1896. In Kohl, K. K. Naturhist. Hofmus, Ann. 11: 475.
Type-species: Oxybelomorpha Kohlii Brauns. Monotypic.
Belomicrus subg. Nototis Arnold, 1927. Transvaal Mus., Ann. 12: 64.
Type-species: Belomicrus (Nototis) bicornutus Arnold. Monotypic.
Belomicrus subg. Pseudoxybelus Gussakovskij, 1933. Akad. Nauk S. 8S. S. R., Zool. Inst.,
Trudy 1: 286.
Type-species: Belomicrus (Pseudoxybelus) persa Gussakovskij. Monotypice.

Seanty information is available on the biology of five species, two of which have been reported
to construct multicellular nests with up to 4 cells per nest. Three North American species prey
upon nymphal or adult mirid bugs; one North American and one European species prey upon
adult melyrid beetles.

Revision: Pate, 1940. Amer. Ent. Soc., Trans. 66: 11-96, 209-257, 55 figs. (N. Amer. spp.)
apache Pate. N. Mex., Ariz.
Belomicrus apache Pate, 1940. Amer. Ent. Soc., Trans. 66: 15. 2.
bridwelli Pate. Va., Tenn., Fla.
Belomicrus bridwelli Pate, 1940. Amer. Ent. Soc., Trans. 66: 93. d, 2.
cahuilla Pate. Calif.
Belomicrus‘cahuilla Pate, 1940. Amer. Ent. Soc., Trans. 66: 39. d.
cladothricis (Cockerell). Tex. to Calif., Nev., Idaho; Mexico (Baja California).
Oxybelus cladothricis Cockerell, 1895. Canad. Ent. 27: 309. d, 2.
Belomicrus cladothricis prosopidis Pate, 1940. Amer. Ent. Soc., Trans. 66: 72. d, °.
Belomicrus minidoka Pate, 1940. Amer. Ent. Soc., Trans. 66: 79. d.
coloratus Baker. Idaho, Nev., Calif. Prey: Miridae sp. nymphs.
Belomicrus colorata Baker, 1909. Pomona Col., Jour. Ent. 1: 29. 9.

Biology: Bohart and Menke, 1976. Sphecid wasps of world, p. 363 (prey).
cookii Baker. Calif.
Belomicrus Cookii Baker, 1909. Pomona Col., Jour. Ent. 1: 29. d, °.
cucamonga Pate. Wash., Calif.
_Belomicrus cucamonga Pate, 1940. Amer. Ent. Soc., Trans. 66: 76. 2.
eriogoni Pate. Calif., Oreg., Nev., Utah.
Belomicrus cladothricis eriogoni Pate, 1940. Amer. Ent. Soc., Trans. 66: 70. d, 2.
forbesii (Robertson). Colo. and Wyo. north to Man. and west to B. C. and Calif. Ecology: Nests
in compact, sandy-clay soil sometimes mixed with gravel, makes 4-8 cells per nest,
stores 5-13 prey per cell. Parasite: Senotainia trilineata (Wulp). Prey: Orectoderes
obliquus Uhl. nymphs and a few adults, Miridae sp. nymphs. Predator: Philanthus
pulcher D. T.
Oxybelus Forbesii Robertson, 1889. Amer. Ent. Soc., Trans. 16: 85. 6.
Oxybelus (Oxybeloides) columbianus Kohl, 1892. K. K. Naturhist. Hofmus., Ann. 7: 208. d,
9

Belomicrus larimerensis Rohwer, 1908. Ent. News 19: 417. °.
Taxonomy: Evans, 1969. Kans. Ent. Soc., Jour. 42: 123-124, figs. 6-9 (larva).

Biology: Evans, 1969. Kans. Ent. Soc., Jour. 42: 122-123, fig. 5 (nest, prey transport).
franciscus Pate. Calif. Ecology: Nests in sand. Prey: Trichochrous antennatus Mots. adults.
Belomicrus franciscus Pate, 1931. Ent. News 42: 77. 6, °.
Belomicrus quemaya Pate, 1940. Amer. Ent. Soc., Trans. 66: 47. ¢.
Biology: Williams, 1936. Pan-Pacific Ent. 12: 3-6 (nest, prey).

istam Pate. Calif.
Belomicrus istam Pate, 1940. Amer. Ent. Soc., Trans. 66: 81. d, 2.

jurumpa Pate. Calif.
Belomicrus jurumpa Pate, 1940. Amer. Ent. Soc., Trans. 66: 53. d, °.

1652 Hymenoptera in America North of Mexico

maricopa Pate. Ariz., N. Mex.

Belomicrus maricopa Pate, 1947. Ent. Soc. Wash., Proc. 49: 54. °.
mescalero Pate. Calif., N. Mex.

Belomicrus mescalero Pate, 1940. Amer. Ent. Soc., Trans. 66: 87. 9.
mono Pate. Calif.

Belomicrus serrano mono Pate, 1940. Amer. Ent. Soce., Trans. 66: 46. °.
pachappa Pate. Calif.

Belomicrus pachappa Pate, 1940. Amer. Ent. Soc., Trans. 66: 73. 2.
penuti Pate. Calif., Nev., Oreg. Prey: Miridae sp. nymphs.

Belomicrus forbesii penuti Pate, 1940. Amer. Ent. Soc., Trans. 66: 27. 6, °.

Biology: Bohart and Menke, 1976. Sphecid wasps of world, p. 363 (prey).

potawatomi Pate. Iowa.

Belomicrus potawatomi Pate, 1947. Ent. Soc. Wash., Proc. 49: 57. 2.
querecho Pate. N. Mex.

Belomicrus querecho Pate, 1940. Amer. Ent. Soc., Trans. 66: 36. d.
sechi Pate. South. Calif.

Belomicrus sechi Pate, 1940. Amer. Ent. Soc., Trans. 66: 60. 6.
serrano Pate. Calif.

Belomicrus serrano serrano Pate, 1940. Amer. Ent. Soc., Trans. 66: 42. d, 9.
timberlakei Pate. Calif.

Belomicrus timberlakei Pate, 1940. Amer. Ent. Soe., Trans. 66: 91. ¢.
tuktum Pate. Calif.

Belomicrus tuktum Pate. 1940. Amer. Ent. Soc., Trans. 66: 84. 6.
vanyume Pate. Calif.

Belomicrus vanyume Pate, 1940. Amer. Ent. Soc., Trans. 66: 17. °.
vierecki Pate. Calif., N. Mex.; Mexico (Baja California, Puebla).

Belomicrus vierecki Pate, 1940. Amer. Ent. Soc., Trans. 66: 56. d, °.

Genus ENCHEMICRUM Pate

Enchemicrum Pate, 1929. Ent. News 40: 219.
Type-species: Enchemicrum australe Pate. Orig. desig.

The genus contains only one species.

Revision: Pate, 1940. Amer. Ent. Soc., Trans. 66: 257-264.
australe Pate. Ga. to Ariz. north to Okla. and Ill. Ecology: Makes multicellular nest in firmly
packed sand, stores 12-16 flies per cell. Prey: Paralimna texana Cr., Zeros flavipes Cr.;
Medetera californiensis Whlr. Predator: Iridomyrmex pruinosus analis (Andre),

Dorymyrmex pyramicus (Rog.).
Enchemicrum australe Pate, 1929. Ent. News 40: 220. 6, °.

Biology: Bohart and Holland, 1966. Pan-Pacific Ent. 42: 161 (nest, prey, predators).

Genus OXYBELUS Latreille

Oxybelus Latreille, 1796. Precis Caract. Gen. Ins., p. 129. No species.
Type-species: Crabro uniglumis of Fabricius. First included species.
Notoglossa Dahlbom, 1845. Hym. Europaea, v. 1, p. 514.
Type-species: Notoglossa sagittata Dahlbom. Monotypice.
Alepidaspis Costa, 1882. Accad. delle Sci. Fis. e Mat. Napoli, Atti 9 (11): 35.
Type-species: Alepidaspis diphyllus Costa. Monotypic.
Oxybelus subg. Anoxybelus Kohl, 1923. Konowia 2: 274.
Type-species: Oxybelus (Anoxybelus) Maidlii Kohl. Monotypic.
Gonioxybelus Pate, 1937. Amer. Ent. Soc., Mem. 9: 28.
Type-species: Oxybelus nigripes Olivier. Orig. desig.
Orthoxybelus Pate, 1937. Amer. Ent. Soc., Mem. 9: 45.
Type-species: Vespa uniglumis Linnaeus. Orig. desig.
Latroxybelus Noskiewiez and Chudoba, 1950. Polskie Pismo Ent. 19: 300.

Superfamily SPHECOIDEA 1653

Type-species: Oxybelus latro Olivier. Monotypic.

Bohart and Schlinger (1957) recognized six species groups in the Nearctic fauna. However,
Peckham, Kurezewski and Peckham (1973) reported that some ethological information is not in
concordance with this classification. It seems preferable not to recognize species groups at this
time pending further systematic studies based on both morphology and ethology.

The majority of species nest in easily friable soil such as sand, and construct uni- or multicellu-
lar nests. Adult Diptera are preyed upon, some species using flies as large as themselves, others
much smaller kinds. The prey is transported in flight either impaled on the sting or carried by
the legs; some species which carry the prey with their legs may land and impale the prey on the
sting before entering the nest. A number of species store mosty male flies, and a few are known
to prey upon males only. The cell may be stored with as few as 2 flies or as many as 38, depend-
ing upon the species of wasp.

Taxonomy: Robertson, 1889. Amer. Ent. Soc., Trans. 16: 77-85. —Mickel, 1918 (1917). Nebr.
Univ. Studies 17: 44-51 (Nebr. spp.). —Bohart and Schlinger, 1956. Pan-Pacifie Ent. 32:
157-165 (annotated synonymical list of N. Amer. spp.). —Bohart and Schlinger, 1957. Calif.
Ins. Survey, Bul. 4: 103-142, 23 maps, 93 figs. (deser. of Calif. spp. and key to N. Amer.
spp.).

Biology: Bohart, Lin and Holland, 1966. Ent. Soc. Amer., Ann. 59: 820 (prey records of N.
Amer. spp.). —Peckham, Kurezewski and Peckham, 1973. Ent. Soc. Amer., Ann. 66: 658-660
(ethology of N. Amer. spp.).

abdominalis Baker. U. S. west of 100th meridian, north to Idaho, Nebr,; north. Mexico.
Oxybelus abdominalis Baker, 1896. Ent. News 7: 158. 6.
Notolossa calligaster Viereck, 1906. Amer. Ent. Soc., Trans. 32: 214. 2.

argenteopilosus Cameron. West. N. Amer. from Colo. and Oreg. south to Mexico
(Tehuantepec).
Oxybelus argenteopilosus Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 158, pl. 9, figs.
23, 23a. 9.

argypheus Bohart and Schlinger. Calif., Ariz., south. Colo.
Oxybelus argypheum Bohart and Schlinger, 1956. Biol. Soc. Wash., Proc. 69: 38. °.

bipunctatus bipunctatus Olivier. Maine to Va.; Europe, Japan. Ecology: Nests in sand, makes

1-4 cells per nest, stores 2-12 prey per cell. Apparently adventive from Europe in 1935.
Parasite: Phrosinella fulvicornis (Coq.), Senotainia trilineata (Wulp), Sarcophagidae
spp. Prey: Allognosta fuscitarsis (Say), Caloparyphus tetraspilus (Lw.), Euparyphus
stigmaticalis Lw., Microchrysa polita (L.), Nemotelus nigrinus Fall., Oxycera picta
Wulp, O. variegata Oliv., Sargus decorus Say; Chrysopilus quadratus (Say),
Symphoromyia sp.; Psilocephala Haemorrhoidalis (Macq.); Diaphorus pseudopacus
Robinson; Callomyia venusta Snow, Platypeza anthrax Lw.; Pipunculus sp;
Chrysogaster inflatifrons Shann., C. nigripes Lw., Heringia salax (Lw.), Neocnemodon
elongatus (Curr.), Platycheirus erraticus Curr.; Lonchaea nudifemorata Mall., L. spp.;
Pholeomyia indecora (Lw.); Hylemya florilega (Zett.), H. fugax (Meig.), H. platura
(Meig.), H. cilicrura (Rond.), H. sp.; Azelia sp., Fannia canicularis (L.), F. coracina
(Lw.), F. depressa (Stein), F. manicata (Meig.), F. nidicola Mall., F. sociella (Zett.),
Gymnodia cilifera (Mall.), Hebecnema vespertina (Fall.), Hydrotaea armipes (Fall.), H.
basdeni Coll., H. occulta (Meig.), Spilogona sp.; Melanomya sp.; Metopia argyrocephala
(Meig.); Alophorella aeneoventris (Will.), A. fumosa (Coq.), A. pulverea (Coq.),
Hyalomya purpurascens (Tns.), Hyalomyodes triangulifer (Lw.); prey consists of male
Brachycera and Cyclorrhapha, rarely females of latter. Another subsp. occurs in the
Mediterranean area.

Oxybelus bipunctatus Olivier, 1811. Encycl. Meth. Ins., v. 8, p. 597. °.

Oxybelus nigroaeneus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 113. d.

Oxybelus laevigatus Schilling, 1848. Schles. Gesell. f. Vaterland. Kult. Arb. im Jahre 1847,
p. 105.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 111, figs. 107-115 (larva).

Biology: Pate, 1946 (1945). Brooklyn Ent. Soc., Bul. 40: 165 (prey). —Krombein, 1948. Ent.
Soc. Wash., Proc. 50: 67 (nest, prey). —Krombein, 1956. Brooklyn Ent. Soc., Bul. 51: 43

1654 Hymenoptera in America North of Mexico

(prey). —Kurezewski and Harris, 1968. N. Y. Ent. Soe., Jour. 76: 81-83 (nest, parasites).
—Peckham, Kurezewski and Peckham, 1973. Ent. Soc. Amer., Ann. 66: 647-651, figs. 1, 3-7
(nest, prey transport, egg, mating, parasites).
californicus Bohart and Schlinger. West. U.S. and north. Mexico.
Oxybelus californicum Bohart and Schlinger, 1956. Pan-Pacifie Ent. 32: 147. d, ?.
canalis Bohart and Schlinger. Southwest. U. S., north to Utah; north. Mexico.
Oaybelus canalis Bohart and Schlinger, 1956. Pan-Pacific Ent. 32: 149. d, °.
cochise Pate. Calif., Ariz., N. Mex., Tex.; Mexico (Chihuahua, Baja California).
Oxybelus cochise Pate, 1943. Brooklyn Ent. Soc., Bul. 38: 93. d.
cocopa Pate. Calif.
Oxybelus cocopa Pate, 1943. Pan-Pacifiec Ent. 19: 121. d.
cornutus Robertson. U.S. west of 100 degrees W., Montana south to Mexico (Guadalajara).
Oxybelus cornutus Robertson, 1889. Amer. Ent. Soc., Trans. 16: 80. d.
Oxybelus quadricolor Cockerell and Baker, 1896. Psyche 7 (sup.): 21. °.
Oxybelus polygoni Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 116. 2.

crandalli Bohart and Schlinger. Ariz., Ark.
Oxybelus crandalli Bohart and Schlinger, 1956. Pan-Pacifie Ent. 32: 154. °.
cressonii Robertson. Centr. U. S., Utah, and Tex., east to Va., north to Mich. Ecology: Makes
unicellular nest in sand, stores 10-15 prey per cell. Prey: Neopachygaster maculicornis
(Hine); Thaumatomyia bistriata (Wlkr.); Coelotanypus scapularis (Lw.).
Oxybelus Cressonii Robertson, 1889. Amer. Ent. Soc., Trans. 16: 88. 2.

Biology: Krombein, 1948. Ent. Soc. Wash., Proc. 50: 67 (prey). —Kurezewski, 1972. Kans. Ent.
Soce., Jour. 45: 398-400, fig. la, b (nest, prey transport).
decorosus (Mickel). U. S. east of 100 degrees W., Vt. and Minn., south to Fla.
Notoglossa decorosa Mickel, 1916. Amer. Ent. Soc., Trans. 42: 430. °.

emarginatus Say. U.S. and south. Canada, Mexico south nearly to Mexico City. Ecology:

Usually makes unicellular nest in sand, occasionally a 2-celled nest, stores 4-38 prey per
cell. Parasite: Diptera spp., Miltogrammini sp. ? Prey: Anarete buscki (Felt), Lestremia
sp.; Chironomus sp.; Chaoborus punctipennis Say; Culicoides stellifer (Coq.); Simulium
sp. in jenningsi Mall. group; Microchrysa polita (L.); Syneches simplex Wlkr.;
Chrysotus obliquus Lw., C. pomeroyi Parent, C. spp., Condylostylus caudatus (Wied.),
C. flavipes (Ald.), C. nigrofemoratus (Wlkr.), C. patibulatus (Say), Diaphorus gibbosus
Van D., D. similis Van D., Gymnopternus exilis Lw., G. spp., Medetera veles Lw., M.
vittata Van D., M. sp., Neurigona lateralis (Say); Pipunculus sp.; Mesograpta
marginata (Say), Toromerus geminatus (Say); Euxesta notata (Wied.); Amphicnephes
pullus (Wied.), Rivellia variabilis Lw., R. winifredae Namba, R. sp.; Euaresta bella
(Lw.); Sepsis punctum (F.); Camptoprosopella borealis Shew., Lyciella annulata (Mel.),
L. sp., Minettia lupulina (F.); Leptocera ferruginata (Sten.), L. Pararoralis Duda, L. sp.;
Desmometopa sordida (Fall.), D. tarsalis Lw., Leptometopa latipes (Meig.), Madiza
glabra Fall., P. indecora (Lw.); Notiphila sp.; Drosophila busckii Coq., D. robusta Sturt.,
D. sp., Scaptomyza pallida (Zett.); Conioscinella minor (Adams), Diplotoxa versicolor
(Lw.), Ectecephala sulcata Sabr., Gaurax sp., Hippelates particeps (Beck.), H. spp.,
Oscinella frit (L.), O. luteiceps Sabr., O. soror (Macq.), O. umbrosa (Lw.), O. sp.,
Thawmatomyia glabra (Meig.); Agromyza sp., Cerodontha sp., Melanagromyza sp.,
Ophiomyia nasuta (Mel.), O. sp.; Mumetopia terminalis (Lw.); Hylemya florilega
(Zett.), H. platura (Meig.), H. sp.; Coenosia sp., Atherigona orientalis Schin., Fannia sp.,
Schoenomyza dorsalis (Lw.); Medina barbata (Coq.); usual prey are Brachycera and
Cyclorrhapha, rarely Nematocera. Predator: Ants.

Oxybelus emarginatus Say, 1837. Boston Jour. Nat. Hist. 1: 375. ¢.

Oxybelus dilutus Baker, 1896. Ent. News 7: 159. °.

Oxybelus trifidus Cockerell and Baker, 1896. Psyche 7 (sup): 23. d.

Notoglossa americana Robertson, 1901. Amer. Ent. soc., Trans. 27: 204. d, 2.

Notoglossa pacifica Rohwer, 1909. Amer. Ent. Soe., Trans. 35: 119. d.

Notoglossa minor Mickel, 1916. Amer. Ent. Soc., Trans. 42: 428. d.

Superfamily SPHECOIDEA 1655

Biology: Hartman, 1905. Tex. Univ. Bul. 65: 29-31 (prey). —Pate, 1930. Brooklyn Ent. Soe.,
Bul. 25: 40 (prey). —Krombein and Kurezewski, 1963. Biol. Soc. Wash., Proc. 76: 150-151
(nest, prey carriage, parasite ?). —Krombein, 1964 (1963). Brooklyn Ent. Soc., Bul. 58: 120
(prey). —Krombein, 1964. Amer. Mus. Novitates 2201: 24-26 (nest, prey transport, mating,
male burrows). —Snoddy, 1968. Ent. Soe. Amer., Ann. 61: 1029-1030 (prey capture on
cattle). —Peckham, Kurezewski and Peckham, 1973. Ent. Soc. Amer., Ann. 66: 655-656,
figs. 12, 15 (nest, prey transport, mating, parasites).

exclamans Viereck. West. U.S., north to 8S. Dak., east to Fla., south to Mexico (Chihuahua).
Ecology: Makes unicellular nest in sand stored with 5 prey. Prey: Senotainia sp.
Oxybelus exclamans Viereck, 1906. Amer. Ent. Soc., Trans. 32: 215. 9.
Oxybelus townsendi Rohwer and Cockerell, 1908. Ent. News 19: 180. d.
Oxybelus argentarius Mickel, 1916. Amer. Ent. Soc., Trans. 42: 431. 2.
Oxybelus pectorosus Mickel, 1918 (1917). Nebr. Univ. Studies 17: 321. ¢.

Biology: Kurezewski, 1972. Kans. Ent. Soc., Jour. 45: 400-402, fig. 1e (nest, prey transport).
fossor Rohwer and Cockerell. U. S. west of 100 degrees W., north to Nebr.; Mexico (Baja
California).
Oxybelus fossor Rohwer and Cockerell, 1908. Ent. News 19: 179. 9.
Oxybelus wmbrosus Mickel, 1916. Amer. Ent Soc., Trans. 42: 432. 6.
Oxybelus puente Pate, 1943. Pan-Pacific Ent. 19: 125. dg, 2.
frontalis Robertson. U. 8. east of Rocky Mts., Tex. north to Mich. and Pa.; Mexico (Veracruz).
Oxybelus frontalis Robertson, 1889. Amer. Ent. Soc., Trans. 16: 83. 6, 2.
inornatus (Robertson). Northeast. U. S., Mich. to Mass., south to N. C. Prey: Pholeomyia
indecora (Lw.). Predator: Efferia albibarbis (Macq.).
Notoglossa inornata Robertson, 1901. Amer. Ent. Soc., Trans. 27: 203. 6, 2.

Biology: Krombein, 1956. Brooklyn Ent. Soc., Bul. 51: 43-44 (prey).
krombeini Bohart and Schlinger. Calif.
Oxybelus krombeini Bohart and Schlinger, 1956. Pan-Pacific Ent. 32: 149. d, 2.
laetus fulvipes Robertson. Southeast. Coastal States north to Va.
Oxybelus fulvipes Robertson, 1889. Amer. Ent. Soc., Trans. 16: 82. d.
Oxybelus floridanus Robertson, 1901. Amer. Ent. Soc., Trans. 27: 203. 6.
laetus laetus Say. U.S. east of 100 degrees W., Tex. to N. C., north to Mich. and Mass.
Oxybelus laetus Say, 1837. Boston Jour. Nat. Hist. 1: 375. 6.
linsleyi Bohart and Schlinger. Calif. Prey: Therevidae sp.
Oxybelus linsleyi Bohart and Schlinger, 1956. Pan-Pacific Ent. 32: 151. d, 9.

Biology: Bohart and Menke, 1976. Sphecid wasps of world, p. 366 (prey).

macswaini Bohart and Schlinger. Calif., Ariz.

Oxybelus macswaini Bohart and Schlinger, 1956. Pan-Pacific Ent. 32: 153. d, @.
major Mickel. East. U. S., Tex. to N. C., north to Nebr., Mich. and Va.

Oxybelus major Mickel, 1916. Amer. Ent. Soc., Trans. 42: 434. ¢.
niger Robertson. Southeast. Canada and northeast. U. S., Minn. and IIl. to N. Y. Prey:

Lonchaea polita Say.
Oxybelus niger Robertson, 1889. Amer. Ent. Soc., Trans. 16: 82. d.

Biology: Krombein, 1958. U. S. Dept. Agr., Monog. 2, Sup. 1: 202 (prey).
packardii Robertson. U.S., Calif. to Fla., north to Ill., Ohio, and N. J.
Oxybelus Packardii Robertson, 1889. Amer. Ent. Soc., Trans. 16: 80. d, 2.
Oxybelus Packardii var. tecanus Robertson, 1889. Amer. Ent. Soc., Trans. 16: 81. d, 9.
Oxybelus heterolepis Cockerell and Baker, 1896. Psyche 7 (sup.): 22. d.
Oxybelus heterolepis var. defectus Cockerell and Baker, 1896. Psyche 7 (sup.): 23
Oxybelus unicus Mickel, 1918 (1917). Nebr. Univ. Studies 17: 323. °.
Oxybelus carolinus Banks, 1921. Ent. Soc. Amer., Ann. 14: 18. 9.
paenemarginatus (Viereck). Kans.
Notoglossa paenemarginatus Viereck, 1906 Amer. Ent. Soc., Trans. 32: 214. °.
paracochise Bohart and Schlinger. Ariz., Tex.; north. Mexico.
Oxybelus paracochise Bohart and Schlinger, 1956. Biol. Soc. Wash., Proc. 69: 37. d, 2.

1656 Hymenoptera in America North of Mexico

parvus Cresson. U. S. west of 100° W., Nebr., and Utah to south. Calif. and north. Mexico.
Oxybelus parvus Cresson, 1865. Ent. Soe. Phila., Proc. 4: 476. SO Zeno
Oxybelus intermedius Baker, 1896. Ent. News 7: 160. 2.
Oxybelus coloradensis Baker, 1896. Ent. News 7: 160. d.
Notoglossa incisura Mickel, 1916. Amer. Ent. Soc., Trans. 42: 4380. 6.
pitanta Pate. Calif., Nev., Ariz., N. Mex.
Oxybelus pitanta Pate, 1943. Pan-Pacifie Ent. 19: 128. 3.
rancocas Pate. N. J.
Oxybelus rancocas Pate. 1943. Brooklyn Ent. Soc., Bul. 38: 91. 6.
rejectus Baker. Colo.
Oxybelus rejectus Baker, 1896. Ent. News 7: 59. 6d.
robertsonii Baker. Northwest. U.S., Nebr. to Oreg., south to Nev. and Calif.
Oxybelus robertsonii Baker, 1896. Ent. News 7: 156. ¢.
Oxybelus varicoloratus Baker, 1896. Ent. News 7: 157. °.
Oxybelus hirsutus Baker, 1896. Ent. News 7: 157. 9.
Oxybelus apicatus Smith, 1908. Nebr. Univ. Studies 8: 409. 3.
Oxybelus glenensis Smith, 1908. Nebr. Univ. Studies 8: 410. 2.
sericeus Robertson. Mass. to Fla. west to S. Dak., Utah, Oreg. and Calif.; Mexico (Baja
California, Nayarit). Ecology: Makes 1-4 cells in moist sand, stores 10-20 flies per cell.
Parasite: Senotainia litoralis Allen. Prey: Ephydra riparia Fall.; Chaetopsis aenea
(Wied.), C. fulvifrons (Macq.).
Oxybelus sericeus Robertson, 1889. Amer. Ent. Soc., Trans. 16: 81. @.
Oxybelus delicatus Mickel, 1918 (1917). Nebr. Univ. Studies 17: 322. ¢.
Oxybelus sericeus crocatus Krombein, 1955. Brooklyn Ent. Soc., Bul. 50: 73. d, 2.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 112, figs. 105-106 (larva).

Biology: Pate, 1930. Brooklyn Ent. Soc., Bul. 25: 41 (prey). —Krombein, 1955, Brooklyn Ent.
Soe., Bul. 50: 74 (nest, mating, prey). —Bohart and Marsh, 1960. Pan-Pacific Ent. 36:
115-118 (nest, prey carriage, mating, life cycle, parasite).

similis Cresson. U. S. west of Rocky Mts.
Oxybelus similis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 476. 9.
Notoglossa striatifrons Mickel, 1916. Amer. Ent. Soc., Trans. 42: 429. 6.
sparideus Cockerell. Southwest. U.S., Ariz. to Tex., Okla., south to Mexico (Veracruz).
Ecology: Nests in damp sand, makes 2-3 cells per nest, stores 3-6 prey per cell. Parasite:
Sarcophagidae spp. from maggots emerging from female prey. Prey: Ceracia dentata
Cogq., Chaetonodexodes sp., Pseudoperichaeta erecta (Coq.), Spathimeigenia hylotomae
(Coq.), Winthemia rufopicta (Big.), Lespesia flavifrons Ben.; Helicobia rapax (Wlkr.),
Oxysarcodexia ventricosa (Wulp), Ravinia derelicta (Wlkr.), Senotainia sp., Sarcophaga
davidsoni Coq.; Psilocephala haemorrhoidalis (Macq.).
Oxybelus sparideus Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 292. d.

Biology: Bohart, Lin and Holland, 1966. Ent. Soc. Amer., Ann. 59: 818-820 (nest, prey
transport, egg, life cycle, cocoon, parasitism by prey).
subcornutus Cockerell. U. S. from Ariz. to East Coast, N. Y. and Mich. south to Mexico
(Chihuahua). Ecology: Nests in sand, makes 1-6 cells per nest, stores 2-8 prey per cell.
Parasite: Macronychia aurata (Coq.), Senotainia trilineata (Wulp). Prey: Allograpta
obliqua (Say), Carposcalis obscura (Say), Melangyna sp., Neocnemodon coxalis (Curr.);
all prey were males.
Oxybelus subcornutus Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 293. °.
Oxybelus punctatus Baker, 1896. Ent. News 7: 60. 6.
Oxybelus striatus Baker, 1896. Ent. News 7: 60. 6.
Oxybelus cockerellii Baker, 1896. Ent. News 7: 61. ¢.
Oxybelus denverensis Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 118. 2.

Biology: Pate, 1945. Brooklyn Ent. Soc., Bul. 40: 165 (prey). —Peckham, Kurezewski and
Peckham, 1973. Ent. Soc. Amer., Ann. 66: 656-658, figs. 14, 16, 17 (nest, prey transport,
parasites).

Superfamily SPHECOIDEA 1657

subulatus Robertson. Northeast. U. S., Colo. to Mass. north to Canada. Ecology: Nests in sand,

makes 1-8 cells per nest, stores 3-11 prey per cell; male wasp guards nest entrance.
Parasite: Phrosinella fulvicornis (Coq.), P. sp., Senotainia trilineata (Wulp), Metopia
argyrocephala (Meig.), Macronychia sp. Prey: Psilocephala haemorrhoidalis (Macq.), P.
frontalis Cole, Thereva candidata Lw., Furcifera rufiventris (Lw.); preys only upon
males.

Oxybelus mucronatus Packard, 1867. Ent. Soc. Phila., Proc. 6: 486. d, 2. Preoce.

Oxybelus subulatus Robertson, 1889. Amer. Ent. Soc., Trans. 16: 79. d, 9.

Oxybelus Packardi Dalla Torre, 1890. Wien. Ent. Ztg. 9: 203. Preoce.

Oxybelus acutus Baker, 1896. Ent. News 7: 61. 9.

Oxybelus albosignatus Smith, 1908. Nebr. Univ. Studies 8: 407. 2.

Oxybelus mottensis Mickel, 1918 (1917). Nebr. Univ. Studies 17: 323. 2.

Biology: Peckham, Kurezewski and Peckham, 1973. Ent. Soc. Amer., Ann. 66: 652-655, figs. 8,
10, 11, 18 (male behavior, nest, prey transport, parasites).
taenigaster (Viereck). Cent. U.S., incl. Ariz., N. Mex., Nebr., south to Mexico (Oaxaca).
Notoglossa taenigaster Viereck, 1906. Amer. Ent. Soc., Trans. 32: 215. 9.
Oxybelus fastigatus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 433. d, @.
Notoglossa albomaculata Mickel, 1918 (1917). Nebr. Univ. Studies 17: 320. d.

timberlakei Bohart and Schlinger. Calif.
Oxybelus timberlakei Bohart and Schlinger, 1956. Pan-Pacifie Ent. 32: 150. d, °.
uniglumis (Linnaeus). Alaska, N. W. T., South. Canada and U.S. south in Mexico to Puebla;

Palaearctic, Europe to Mongolia. Ecology: Nests in sand, makes 1-5 cells per nest,
transports prey on sting, stores 2-13 prey per cell. Parasite: Senotainia rubriventris
(Macq.)?, Metopia argyrocephala (Meig.), Phrosinella fulvicornis (Coq.); these records
are all from U.S. Prey: Allognosta fuscitarsis (Say); Chrysopilus modestus Lw.,
Symphoromyia atripes Big., S. montana Ald.; Anthrax albofasciatus Macq.;
Condylostylus patibulatus (Say); Metasyrphus venablesi (Curr.); Rivellia sp. in
melliginis group; Anthomyia procellaris Rond., Emmesomyia socia (Fall.),
Eustalomyia vittipes (Zett.), Hydrophoria conica (Wied.), H. implicata Huck., Hylemya
depressa Stein, H. fugax (Meig.), H. lasciva Zett., H. platura Meig., H. cilicrura (Rond.),
Pegomyia affinis Stein; Sapromyza monticola Mel.; Fannia coracina (Lw.), F.
enatohensis Seago, F. pellucida (Stein), F. scalaris (F.), Helina duplicata (Meig.),
Musca domestica L., Myospila meditabunda (F.), Ophyra aenescens (Wied.), O.
leucostoma (Wied.), Phaonia fusca (Stein), Spilogona sp.; Bufolucilia silvarum (Meig.),
Lucilia sp., Phaenicia sericata (Meig.), Pollenia rudis (F.); Blaesoxipha hunteri
(Hough), B. reversa (Ald.), Boettcheria cimbicis (Tns.), Helicobia rapax (Wlkr.), Metopia
argyrocephala (Meig.), Oxysarcodexia cingarus (Ald.), Phrosinella fulvicornis (Coq.),
Ravinia acerba (Wlkr.), R. lherminieri (Desv.), Senotainia trilineata (Wulp); Admontia
nasoni (Coq.), Alophora sp., Eulasiona comstocki Tns., Lixophaga sp.; Medina barbata
(Coq.), Voria aurifrons (Tns.); prey in U. S. consist principally of male Brachycera and
Cyclorrhapha, rarely females of latter group. Predator: Philanthus flavifrons Cr., P.
pulcher D. T.

Vespa uniglumis Linnaeus, 1758. Syst. Nat., Ed. 10, v. 1, p. 573. @.

Vespa uniglummis (!) Christ, 1791. Naturgesch. Class. Nomencl. Ins., p. 246.

Nomada punctata Fabricius, 1793. Ent. Syst., v. 2, p. 346.

Crabro tridens Fabricius, 1798. Sup. Ent. Syst., p. 270. °.

Vespa decim-maculata Donovan, 1806. Nat. Hist. Brit. Ins., v. 11, pl. 376, fig. 1. This is a
questionable synonym.

Oxybelus pygmaeus Olivier, 1811. Eneyel. Meth., Ins., v. 8, p. 597.

Oxybelus quadrinotatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app., p. 338.

Oxybelus impatiens Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 390. °.

Oxybelus interruptus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 475. °.

Oxybelus fallax Gerstaecker, 1867. Ztschr. Gesam. Naturw. Halle 30: 91. °.

Oxybelus Brodiei Provancher, 1883. Nat. Canad. 14: 36. °.

Oxybelus quadrinotatus var. montanus Robertson, 1889. Amer. Ent. Soc., Trans. 16: 78. d,
OF

Oxybelus hispanicus Giner, 1943. Inst. Espan. Ent., Trab., Fam. Sphecidae, p. 260. d, 2.

1658 Hymenoptera in America North of Mexico

Biology: Peckham and Peckham, 1898. Wis. Geol. and Nat. Hist. Survey, Bul. 2: 73-76 (nest,
prey transport). —Parker, 1915. Ent. Soc. Wash., Proc. 17: 74-75 (nest, prey transport,
parasites). —Pate, 1930. Brooklyn Ent. Soc., Bul. 25: 41 (prey). —Williams, 1936.
Pan-Pacific Ent. 12: 1-3, fig. 1 (nest, prey transport, predator). —Krombein, 1936. Ent.
News 47: 95 (prey transport). —Strandtmann, 1945. Ent. Soc. Amer., Ann. 38: 313 (nest,
prey transport, parasite). —Krombein, 1948. Ent. Soc. Wash., Proc. 50: 67 (prey).
—Krombein, 1956. Brooklyn Ent. Soc., Bul. 51: 44 (prey transport). —Evans, 1970. Mus.
Compar. Zool., Bul. 140: 493 (nest, prey transport, parasites, predator). —Peckham,
Kurezewski and Peckham, 1973. Ent. Soc. Amer., Ann. 66: 651-652, figs. 2, 9 (nest, prey
transport, parasites).

ventralis Fox. Pacific Coast, Wash., south to Mexico (Baja California).
Oxybelus ventralis Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 107. @.
Oxybelus manni Rohwer, 1909. Amer. Ent. Soc., Trans. 35: 117. 9.

xerophilus Bohart and Schlinger. Calif., Ariz.
Oxybelus xerophilum Bohart and Schlinger, 1956. Pan-Pacific Ent. 32: 154. d.

SUBFAMILY CRABRONINAE

This large subfamily includes species which nest in the ground, in pre-existing cavities in
wood, twigs, stems, and in the soft pith of various plants or shrubs.

Revision: Fox, 1895. Amer. Ent. Soc., Trans. 22: 129-226 (N. Amer. spp.).

Taxonomy: Pate, 1944. Amer. Midland Nat. 31: 329-384 (gen. of world). —Leclercgq, 1954.
Monog. Hym. Crabron., 371 pp., 40 figs., 84 maps (phylogeny, zoogeography, key to world
gen., catalog of world spp.).

Genus ANACRABRO Packard

Anacrabro Packard, 1866. Ent. Soc. Phila., Proc. 6: 67.
Type-species: Anacrabro ocellatus Packard. Monotypic.
ocellatus boerhaviae Cockerell. N. Mex., Ariz.; Mexico (Chihuahua, Sonora, Sinaloa, Durango,
Colima 2).
Anacrabro boerhaviae Cockerell, 1895. Canad. Ent. 27: 308. ¢.
ocellatus ocellatus Packard. S. Canada and U. S. east of Rockies, except Fla. Ecology: Nests
in sand, makes 1-9 or more cells per nest, stores 4-7 prey per cell. Parasite:
Miltogrammini sp.?, Diptera sp. Prey: Lygus oblineatus (Say), L. lineolaris (Beauv.),
Plagiognathus politus Uhl.; only adults are stored. Predator: Acanthomyops claviger

(Rog.), Monomorium sp.
Anacrabro ocellatus Packard, 1866. Ent. Soc. Phila., Proc. 6: 68. 2.
Thyreopus rugoso-punctatus Provancher, 1883. Faune Ent. Canada, Hym., p. 664. 9. A

secondary homonym in Crabro.
Crabro rugulosopunctatus Dalla Torre, 1897. Cat. Hym., v. 8, p. 624. N. name.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 102, figs. 75-84 (larva).

Biology: Barth, 1909 (1908). Wis. Nat. Hist. Soc., Bul. 6: 147-153, 10 figs. (nest, prey, parasite,
predator). —Kurcezewski and Peckham, 1970. Ent. Soc. Amer., Ann. 63: 1419-1424, 7 figs.
(nest, prey transport, egg, parasite, predator).

ocellatus robertsoni Rohwer. Fla.
Anacrabro robertsoni Rohwer, 1920. Ent. Soc. Wash., Proce. 22: 58. 2.

Taxonomy: Krombein, 1948. Brooklyn Ent. Soc., Bul. 43: 20-21 (status of N. Amer. spp.).
—Lecleregq, 1973. Acta Zool. Lilloana 30: 38-41 (key to New World spp., and revision of

Neotropical spp.).
Genus ENCOPOGNATHUS Ashmead

The name Encopognathus was first employed by Kohl, 1896 (K. K. Naturhist. Hofmus., Ann.
11: 485, 486). However, it may not be validated from that date, for Kohl proposed it as a name
for a species group (“Artengruppe”) of the subgenus Lindenius. The earliest use of the name in

Superfamily SPHECOIDEA 1659

a generic or subgeneric sense was by Ashmead, 1899, who characterized it as a genus.

The species nest in soil, and African forms have been reported to provision their nests with
ants. One species in North America and two species in Sri Lanka have been observed recently
preying upon Miridae, both nymphs and adults.

Revision: Pate, 1943. Lloydia 6: 53-76, 15 figs. (key to subg. and revision of N. Amer. spp.).

Genus ENCOPOGNATHUS Subgenus ENCOPOGNATHUS Ashmead

Encopognathus Ashmead, 1899. Canad. Ent. 31: 166.
Type-species: Crabro (Encopognathus) Braueri Kohli. Orig. desig.

The typical subgenus does not occur in North America.

Genus ENCOPOGNATHUS Subgenus RHECTOGNATHUS Pate

Encopognathus subg. Rhectognathus Pate, 1936. Ent. News 47: 147.
Type-species: Encopognathus (Rhectognathus) pectinatus Pate. Orig. desig.
pectinatus Pate. Calif.
Encopognathus (Rhectognathus) pectinatus Pate, 1936. Ent. News 47: 148. 9, d.
rufiventris Timberlake. Calif. (San Bernardino Co.).
Encopognathus (Rhectognathus) rufiventris Timberlake, 1940. Ent. News 51: 167. °.

Genus ENCOPOGNATHUS Subgenus TSAISUMA Pate

Encopognathus subg. Tsaiswma Pate, 1943. Lloydia 6: 57.
Type-species: Lindenius wenonah Banks. Orig. desig.

wenonah (Banks). Calif., Oreg.
Lindenius wenonah Banks, 1921. Ent. Soc. Amer., Ann. 14: 16. ¢.

Genus ENTOMOGNATHUS Dahlbom

These wasps nest in the ground and prey upon adult chrysomelid beetles.
Biology: Miller and Kurezewski, 1972. Psyche 79: 61-69, 72-75 (nesting behavior).

Genus ENTOMOGNATHUS Subgenus ENTOMOGNATHUS Dahlbom

Entomognathus Dahlbom, 1844. Hym. Europaea, v. 1, p. 295.
Type-species: Crabro brevis Vander Linden. Monotypic.

The typical subgenus does not occur in North America.

Genus ENTOMOGNATHUS Subgenus TONCAHUA Pate

Entomognathus subg. Toncahua Pate, 1944. Amer. Midland Nat. 31: 341.
Type-species: Entomognathus texanus Cresson. Orig. desig.

Encopognathus subg. Florkinus Leclercq, 1956. Inst. Roy. Sci. Nat. Belg., 32 (20): 2.
Type-species: Encopognathus (Florkinus) evolutionis Leclereq. Orig. desig.

Taxonomy: Krombein, 1963. Biol. Soc. Wash., Proc. 76: 247-254 (spp. of eastern U. S.).

arenivagus Krombein. Fla. (Arcadia), N. C. (?). Ecology: Nests in sand.
Entomognathus (Toncahua) arenivaga Krombein, 1963. Biol. Soc. Wash., Proc. 76: 249. 3,
Oe

lenapeorum Viereck. N. J. to Va., Kans. Ecology: Nests in open wooded areas.
Entomognathus lenapeorum Viereck, 1904. Amer. Ent. Soc., Trans. 30: 239. 9.
memorialis Banks. Conn. to Va., Kans. Ecology: Nests in sandy-loam cliffs, occasionally in
pre-existing burrows, makes 9-20 cells per nest which may be arranged singly, in
fan-shaped clusters, or in tandem; stores 3-9 prey per cell. Prey: Altica ulmi Woods, A.
marevagans Horn; adults.
Entomognathus memorialis Banks, 1921. Ent. Soc. Amer., Ann. 14: 16. 9.

Taxonomy: Miller and Kurczewski, 1972. Psyche 79: 69-72, figs. 5-12 (larva).

1660 Hymenoptera in America North of Mexico

Biology: Miller and Kurezewski, 1972. Psyche 79: 64-69, figs. 1-4 (mating, nest, prey
transport, egg, cocoon).
texanus Cresson. Pa., Kans. to Tex., N. Mex.; Mexico (Tamaulipas). Prey: Diabrotica tricincta
Say adult.
Entomognathus tecanus Cresson, 1887. Amer. Ent. Soc., Trans., Sup. Vol., p. 286. °, d.
Anothyreus panurgoides Viereck, 1904. Amer. Ent. Soc., Trans. 30: 239. d (2 misdet.).

Biology: Cazier and Mortenson, 1965. Pan-Pacific Ent. 41: 33 (prey).

Genus LINDENIUS Lepeletier and Brulle

Lindenius Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 791.
Type-species: Crabro albilabris Fabricius. Desig. by Westwood, 1839.

Lindenius subg. Chalcolamprus Wesmael, 1852. Acad. Roy. Sci. Belg., Bul. 19: 590.
Type-species: Crabro albilabris of Vander Linden. Monotypic.

Crabro subg. Trachelosimus Morawitz, 1866. Acad. Imp. Sci. St. Petersburg, Bul. 9: 249.
Type-species: Crabro armatus Vander Linden. Monotypic.

These small wasps nest in firmly packed sand or fine gravel, frequently in large aggregations,
and construct a vertical main burrow which usually leads to a short horizontal passage. The prey
consists of small adult Hymenoptera and Diptera, and both nymphal and adult Hemiptera;
specific differences as to the kinds of prey stored are found among the six species studied.

Taxonomy: Pate, 1947. Notulae Nat. 185: 4-7 (key to subg. and syn. of N. Amer. spp.).

Biology: Miller and Kurezewski, 1974 (1973). Ins. Sociaux 20: 365-378, 1 fig. (male-female
interactions in aggregations of several spp.). —Miller and Kurezewski, 1975. N. Y. Ent.
Soc., Jour. 83: 82-120, 12 figs. (comparative behavior of several Nearctic and Palaearctic
spp.)-

armaticeps (Fox). South. Canada, U.S. northeast from Colo. and Tex. Ecology: Nests in sand

cliffs or in flat hard-packed sand, makes 3-11 cells per nest, stores 3-15 prey per cell.
Parasite: Phrosinella fulvicornis (Coq.), Senotainia sp. ? Prey: Diplotoxa versicolor
(Lw.), Meromyza sp. near pratorum Meig., Parectecephala eucera (Lw.), P.
sanguinolenta (Lw.), Thaumatomyia glabra (Meig.), T. sp., Chlorops sp.; all prey
records are of Chloropidae.

Crabro armaticeps Fox, 1895. Amer. Ent. Soc., Trans. 22: 185. ¢.

Crabro flaviclypeus Fox, 1895. Amer. Ent. Soc., Trans. 22: 186. 2.

Crabro (Lindenius) zellus Rohwer, 1909. Ent. News 20: 151. °.

Biology: Kurezewski, 1972. Kans. Ent. Soc., Jour. 45: 399 (prey). —Miller and Kurezewski,
1974 (1973). Ins. Sociaux 20: 368, 372-378 (male-female intraspecific interaction, nest,
parasites). —Miller and Kurezewski, 1975. N. Y. Ent. Soc., Jour. 83: 90-96, figs. 1-5 (nest,
prey capture and transport, egg, cocoon, life cycle, parasite).

buccadentis Mickel. U.S. east from Wyo., north from Ark. Ecology: Nests in bare, level
hard-packed roadbeds of coarse sand, makes up to 4 cells per completed nest, stores
22-77 prey per cell. Prey: Drapetis sp. near divergens Lw.; Agathis spp., Apanteles
paralechiae Mues., A. sp., Bracon sp., Diaeretus sp., Orgilus sp., Pauesia sp.,
Phanerotoma sp., Rhaconotus cressoni Mues. and Walkl.; Acrolytina sp., Mesochorus
sp., Toxophoroides scitulus (Cr.); Achrysocharella silvia Gir., Chrysocharis sp.,
Closterocerus tricinctus (Ashm.), Euderus sp., Hulophus anomocerus (Cwfd.),
Euplectrus sp., Hyssopus sp., Tetrastichus whitmani (Gir.), T. spp.; Perilampus
fulvicornis Ashm., P. robertsoni Cwfd.; Ormyrus brunneipes Prov.; Capellia sp.,
Erythromalus sp., Gastrancistrus aphidis (Gir.), Pachyneuron siphonophorae (Ashm.),
Pteromalinae sp.; Bruchophagus sp., Eudecatoma sp.; Spilochalcis sp.; Charips spp.,
Cynipinae sp.; Apenesia parapolita Evans, Goniozus sp.; Monomorium minimum
(Buckl.), Tapinoma sessile (Say); preferred prey were species of Ichneumonoidea and
Chalcidoidea, and only one dipteran was stored.

Lindenius buccadentis Mickel, 1916. Amer. Ent. Soc. Trans. 42: 427. 2, 6.

Biology: Miller and Kurezewski, 1974 (1978). Ins. Sociaux 20: 368, 372-378 (male-female
intraspecific interaction). —Miller and Kurezewski, 1975. N. Y. Ent. Soc., Jour. 83: 96-101,
fig. 6 (nest, prey transport, egg, cocoon).

Superfamily SPHECOIDEA 1661

californicus Court and Bohart. Calif.

Lindenius californicus Court and Bohart, 1958. Pan-Pacifie Ent. 34: 161, figs. 1, 8, 11. d,

oe
columbianus (Kohl). South. Canada, U. S. Ecology: Nests in dense aggregations in firmly

packed clay-sand, occasionally in pre-existing burrows and in sand cliffs, may make up
to 10 cells per nest, stores 8-76 prey per cell. Parasite: Phrosinella fulvicornis (Coq.),
Senotainia sp. ?; Phalacrotophora halictorum Mel. and Brues ?; Myrmosa wnicolor Say?
Prey: Dasyhelea grisea (Coq.), D. spp., Forcipomyia brevipennis (Macq.), Jenkinshelea
magnipennis (Joh.); Chironomus spp., Cricotopus sp., Orthocladius spp., Paratendipes
subaequalis (Mall.), Pentaneura sp., Procladius spp., Psectrocladius sp., Tanytarsus sp.;
Bradysia sp.; Svammerdamella obtusa Cook, S. sagittata Cook, Scatopse fuscipes Meig.;
Anarete johnsoni (Felt), A. pritchardi Kim, A. spp., Asteromyia carbonifera (O. S.),
Clinodiplosis sp., Dasineura sp., Mayetiola sp., Neolasioptera spp., Ozirhincus millefolii
(Wachtl), Porricondyla sp., Procystiphora sp.; Drapetis septentrionalis Mel., D. sp.,
Platypalpus trivialis Lw., P. sp., Rhamphomyia sp.; Chamaemyia juncorum (Fall.),
Leucopis sp.; Madiza parva (Adams), Leptometopa halteralis (Coq.), L. latipes (Meig.),
Paramyia nitens (Lw.); Philygria debilis Lw., Hydrellia sp.; Conioscinella
melancholica (Beck.), C. minor (Adams), C. triorbiculata (Sabr.), Diplotoxa versicolor
(Lw.), Hippelates bishoppi Sabr., H. sp. near bishoppi Sabr., Meromyza sp., Olcella
cinerea (Lw.), O. parva (Adams), O. quadrivittata Sabr., O. trigramma (Lw.), Oscinella
carbonaria (Lw.), O. frit (L.), O. luteiceps Sabr., O. soror (Macq.), O. umbrosa (Lw.), O.
sp., Siphonella nigripalpis (Mall.); Agromyza sp., Cerodontha dorsalis (Lw.), C. sp.,
Liriomyza sp., Ophiomyia sp., Phytoliriomyza arctica (Lundb.), Pseudonapomyza
lacteipennis (Mall.); Diptera spp.; Oriws insidiosus (Say), O. tristicolor (White);
Chlamydatus associatus (Uhl.), Miridae spp.; Lygaeidae sp.; Aphis sp., Capitophorus
elaeagni (Del Guer.), Rhopalosiphum maidis (Fitch), Schizaphis sp.; Hemiptera spp.;
Agathis spp., Apanteles limentidis Riley, A. xylinus (Say), A. spp., Aphidius obscuripes
Ashm., A. spp., Bracon sp., Chelonus sp., Dacnusa sp., Diaeretiella spp., Elasmosoma
sp., Huphoriana uniformis Gah., Lysaphidus sp., Lysiphlebus spp., Microplitis sp.,
Orgilus gelechiae (Ashm.), O. sp., Praon spp., Trioxys spp.; Adelognathus flavopictus
Davis; Polynema sp.; Aprostocetus sp., Chrysocharis sp., Diaulinopsis callichroma
Cwfd., Euderus swhopacus (Gah.), Euderus sp., Hemiptarsenus americanus (Gir.),
Hyssopus novus Gir., Necremnus sp., Notanisomorpha ainsliei Cwfd., Pnigalio sp.,
Sympiesis bimaculatipennis (Gir.), Tetrastichus bruchophagi Gah., T. chlamytis Ashm.,
T. fumipennis (Gir.), T. incertus (Ratz.), T. semilongifasciatus (Gir.), T. tesserus Burks,
T. spp., Entedontini sp.; Copidosoma sp., Anagyrini sp., Bothriothoracini sp.; Ewpelmella
vesicularis (Retz.), Eupelmus allynii (French), E. sp.; Pseudometagea schwarzii
(Ashm.), Eucharitidae sp.; Hiridontomerus isosomatis (Riley), Pseudotorymus lazulellus
(Ashm.); Asaphes lucens (Prov.), Ecrizotes sp., Erixestus winnemanna Cwfd.,
Habrocytus sp., Halticoptera patellanna (Dalm.), H. sp., Heteroschema sp., Homoporus
chalcidiphagus (Walsh and Riley), H. febriculosus (Gir.), Mesopolobus sp., Pachyneuron
allograptae Ashm., P. siphonophorae (Ashm.), P. sp., Parecrizotes marylandensis Gir.,
Pteromalus puparum (L.), P. vanessae Harr., Systasis sp., Tridymus sp., Pirenini sp.,
Pteromalini sp., Tridymini sp.; Bruchophagus sp., Hudecatoma sp., Eurytoma sp.,
Harmolita sp., Systole sp., Spilochalcis albifrons (Walsh); Chalcidoidea sp.; Lygocerus
sp.; Lasius sp.; Spilomena pusilla (Say); prey preferences vary at different localities
depending upon ecological factors; Chironomidae are preferred prey in nesting sites
near water; in other areas Pteromalidae, Scatopsidae, Chloropidae, Eulophidae,
Ceratopogonidae, Milichiidae, Cecidomyiidae, Eucharitidae and Anthocoridae are
important as prey. Predator: Philanthus pulcher D. T., P. pacificus Cr.

Crabro (Lindenius) columbianus Kohl, 1892. K. K. Naturhist. Hofmus., Ann. 7: 203. °.

Crabro errans Fox, 1895. Amer. Ent. Soc., Trans. 22: 184. 9, 6.

Crabro pinguis Fox, 1895. Amer. Ent. Soc., Trans. 22: 186 °.

Ammoplanus salicis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 402. 9.

Biology: Evans, 1970. Mus. Compar. Zool. 140: 491 (nest, prey, predators). — Miller and
Kurezewski, 1974 (1973). Ins. Sociaux 20: 368-378, 1 fig. (male-female intraspecific
interactions, nest, prey, parasites ?). —Miller and Kurczewski, 1975. N. Y. Ent. Soc., Jour.

1662 Hymenoptera in America North of Mexico

83: 101-115, figs. 7-15 (nest, prey capture and transport, egg, cocoon, life cycle). —Bohart
and Menke, 1976. Sphecid wasps of world, p. 383 (nest, prey capture and transport).
inyoensis Court and Bohart. Calif., Nev. '
Lindenius inyoensis Court and Bohart, 1958. Pan-Pacific Ent. 34: 164, figs. 3, 5, 14. 2, d.
latifrons (Fox). Tex.
Crabro latifrons Fox, 1895. Amer. Ent. Soc., Trans. 22: 185. 9.
montezuma (Cameron). South. Calif. to west. Tex.; Mexico (Guanajuato, Guerrero).
Crabro montezwma Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 151, pl. 9, fig. 13, 13a.
RIOr
Lindenius (Trachelosimus) dugesianus Leclercq, 1950. Inst. Roy. Sci. Nat. Belg., Bul. 26,
nNosOnpr las.
neomexicanus Court and Bohart. N. Mex., Colo.
Lindenius neomexicanus Court and Bohart, 1958. Pan-Pacific Ent. 34: 166, figs. 2, 4, 6, 13.
Om Cr
tecuya Pate. Calif. Ecology: Nests in mixed sand-gravel at edge of river. Prey: Diptera spp., 6
families; Chalcidoidea spp., 3 families; Anthocoridae sp.
Lindenius (Trachelosimus) tecuya Pate, 1947. Notulae Nat. 185: 5. 2, d.

Biology: Bohart and Menke, 1976. Sphecid wasps of world, pp. 383-384 (nest, prey).
tylotis Court and Bohart. Calif. Ecology: Nests in silty soil at creek edge, makes 4-24 cells per
nest, and stores an average of 8 prey per cell. Parasite: Myrmosa bradleyi Rob. Prey:
Diptera spp.; Hemiptera spp.; Hymenoptera spp.
Lindenius tylotis Court and Bohart, 1958. Pan-Pacifie Ent. 34: 164, figs. 9, 10, 12. 6, @.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 162-1638, figs. 51-53 (larva).
Biology: Bohart and Menke, 1976. Sphecid wasps of world, p. 383 (nest, prey, parasite).

Genus RHOPALUM Stephens

The North American species, and most of the extralimital taxa, nest in the pith of twigs, in
hollow stems or reeds, in rotten wood, and in abandoned beetle borings in wood. One extralimital
species is known to nest in the ground. Prey of the species occurring in North America includes
small Diptera and Psocoptera. Some extralimital taxa prey upon Aphididae, Psyllidae, winged
Formicidae and Microlepidoptera.

Taxonomy: Bohart, 1974. Ga. Ent. Soc., Jour. 9: 252-260, 33 figs. (review of N. Amer. spp.).

Genus RHOPALUM Subgenus RHOPALUM Stephens

Euplilis Risso, 1826. Hist. Nat. Europ. Merid., v. 5, p. 227.
Type-species: Crabro rufiventris Panzer. Desig. by Pate, 1935.
Rhopalum Stephens, 1829. Nomencl. Brit. Ins., p. 34.
Type-species: Crabro rufiventris Panzer. Desig. by Curtis, 1837.
Physoscelus Lepeletier and Brulle, 1835. Soc. Ent. France, Ann. 3: 804.
Type-species: Crabro rufiventris Panzer. Desig. by Westwood, 1839.
Physoscelis Westwood, 1837. Introd. Mod. Classif. Ins. 2, Gen. Synop., p. 80. Lapsus or
emend.

Taxonomy: Menke, Bohart and Richards, 1974. Bul. Zool. Nomencl. 30: 219-220 (petition to
suppress Euplilis Risso, 1826, and place Rhopalum Stephens, 1829, on Official List of
Generic Names). —Day et al., 1975. Bul. Zool. Nomencel. 32: 96-99 (comments on petition
by Menke et al.). —Krombein, 1976. Bul. Zool. Nomenel. 32: 205-207 (comment on petition
by Menke et al.).

atlanticum Bohart. Ga. (Athens, Atlanta), N. Y. (Long Island). Ecology: Nests in
abandoned borings of Ceratina sp. in dead Erianthus sp. (plume grass), stores 31-41
adult or nymphal Psocoptera per cell. Parasite: Eurytoma inornata Bugbee. Prey:
Caecilius pinicola Bks.; Peripsocus madidus Hagen, P. quadrifasciatus Harr.;
Lachesilla forcepeta Chapm., L. sp.; Psocidus pollutus Walsh, Trichadenotecnum
circuluroides Bed.
Rhopalum atlanticum Bohart, 1974. Ga. Ent. Soe., Jour. 9: 256, figs. 26-33. 3, 2.

Superfamily SPHECOIDEA 1663

clavipes clavipes (Linnaeus). Maine, N. H., R. L., Md., Idaho, Wash., Oreg., Calif.; cent. and

south Europe. Ecology: In Europe nests in straws, stores 16-26 psocid prey per cell.
Prey: In Europe Graphopsocus cruciatus (L.), Peripsocus phaeopterus (Ste.), Ellipsocus
westwoodi Mclack., Mesopsocus immunis (Ste.), Loenisia variegata (Latr.), Hyperates
questfalicus Kolb.; mostly adult females, a few nymphs, one adult male. Another subsp.
occurs in Japan.

Sphex clavipes Linnaeus, 1758. Syst. Nat., ed. 10, p. 569.

Crabro rufiventris Panzer, 1799. Faunae Ins. German., heft 72, fig. 12.

Taxonomy: Marechal, 1929. Soc. Ent. France, Ann. 98: 115-116 (larva).

Biology: Freeman, 1938. Roy. Ent. Soc. London, Proc., Ser. A, 13: 1-2 (nest, prey, cocoon, life
cycle in England).

Genus RHOPALUM Subgenus CORYNOPUS Lepeletier and Brulle

Corynopus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 802.
Type-species: Crabro tibialis Panzer. Monotypic.

Dryphus Herrich-Schaeffer, 1840. Nomencl. Ent., heft. 2, p. 123.
Type-species: Crabro tibialis Fabricius. Monotypic.

Alliognathus Ashmead, 1899. Canad. Ent. 31: 219.
Type-species: Crabro occidentalis Fox. Orig. desig.

Taxonomy: Pate, 1947. Notulae Nat. 185: 7-10 (synonymic notes on N. Amer. spp.).
coarctatum (Scopoli). Canada and U. S. in Transit. and U. Austr. Zones east of Rocky Mts.;

Europe, Siberia. Ecology: Nests in soft pith of dead hibiscus stems and in borings in
wood, makes up to 12 cells per nest, stores as many as 20 prey per cell. Parasite:
Tetrabaeus americanus (Brues); Eurytoma inornata Bugb.; Melittobia chalybui Ashm.;
Megaselia sp.? Prey: Palpomyia subasper (Coq.); Chironomus fulvus Joh., C. viridulus
(L.), C. nervosus Staeg., Chironomidae sp.?

Sphex coarctata Scopoli, 1763. Ent. Carn., p. 293, pl. 42, fig. 778. d.

Crabro crassipes Fabricius, 1798. Sup. Ent. System., p. 270.

Crabro tibialis Fabricius, 1798. Sup. Ent. System., p. 271. Preoce.

Rhopalum modestum Rohwer, 1908. Ent. News 19: 257. d.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 292-293, figs. 111-112 (larva).

Biology: Krombein, 1964. Biol. Soc. Wash., Proc. 77: 90-92 (nest, prey, life cycle, cocoon,
parasites). —Krombein, 1967. Trap-nesting wasps and bees, pp. 259-260 (nest, prey, life
cycle, parasites).

occidentale (Fox). Maine, Conn., N. Y., N. C., Mich., Colo., Wyo., Nev., Calif., Oreg., B. C.
Crabro occidentalis Fox, 1895. Amer. Ent. Soc., Trans. 22: 200. °.
Rhopalum (Alliognathus) carolina Banks, 1921. Ent. Soc. Amer., Ann. 14: 17. 2.
pacificum Bohart. Nev., Calif., Oreg. Ecology: Nests in Sambucus stems. Parasite:
Diomorus zabriskiei Cr.
Rhopalum pacificum Bohart, 1974. Ga. Ent. Soe., Jour. 9: 258, figs. 6-14. d, °.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest, parasite; misdet. as arapaho
Pate).
pedicellatum Packard. Maine, Mass., Conn., N. Y., Md., Wis., Minn., Colo. Ecology: Nests in

pre-existing cavities in old stump and in raspberry stalks, stores 25-33 prey per cell.
Prey: Chironomus sp.

Rhopalum pedicellatum Packard, 1867. Ent. Soc. Phila., Proc. 6: 380. 2, d.

Rhopalum rubrocinctum Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul.
2: 43. 2.

Euplilis (Corynopus) arapaho Pate, 1947. Notulae Nat. 185: 8. 2, d.

Taxonomy: Evans, 1957. Amer. Ent. Soe., Trans. 83: 104-105, figs. 99-104 (larva).

Biology: Packard, 1869. Guide Study Ins., p. 158 (nest). —Peckham and Peckham, 1895. Wis.
Geol. Nat. Hist. Survey, Bul. 2: 42-43 (nest, prey).

1664 Hymenoptera in America North of Mexico

rufigaster Packard. U.S. e. of 100th meridian. Ecology: Nests in pith of green and dead

Hibiscus stems and in old anobiid borings in wood, makes 3-27 cells per nest, stores
29-50 prey per cell. Parasite: Tetrabaeus americanus (Brues); Eurytoma inornata
Bugb.; Diomorus zabriskiei Cr.; Ptychoneura aristalis (Coq.). Prey: Chironomus fulvus
Joh., C. modestus Say, C. neomodestus Mall., C. nervosus Staeg., C. nigrovittatus Mall.,
C. xenolabis (K.), Cricotopus spp., Calopsectra sp., Procladius culiciformis (L.),
Tanytarsus sp., Orthocladiinae sp., Chironomidae spp.; Polymeda cana (Wlkr.); Bezzia
setulosa (Lw.).

Rhopalum rufigaster Packard, 1867. Ent. Soc. Phila., Proc. 6: 382. 2, d.

Rhopalum lucidum Rohwer, 1909. Ent. News 20: 324. 9.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 291-292, fig. 110 (larva).

Biology: Krombein, 1958. Biol. Soc. Wash., Proc. 71: 26 (nest, prey transport). —Krombein,
1964. Biol. Soc. Wash., Proc. 77: 92-98, figs. 12-14 (nest, prey, life cycle, cocoon, parasites).

Genus MONIAECERA Ashmead

Moniaecera Ashmead, 1899. Canad. Ent. 31: 220.
Type-species: Crabro abdominalis Fox. Orig. desig.

The described species are all recorded from America north of Mexico, but there are some un-
described taxa in Mexico. These wasps nest in the soil and prey upon a variety of insect groups
including leafhoppers, psyllids, mirids and chironomids.

Revision: Pate, 1948. Amer. Ent. Soc., Trans. 74: 41-60, 6 figs., 1 map.
abdominalis (Fox). Ga., Tex., Kans., Ariz. Ecology: Nests in sand. Prey: Tylozygus bifidus

(Say).
Crabro abdominalis Fox, 1895. Amer. Ent. Soc., Trans. 22: 198. 2.

Biology: Hartman, 1905. Tex. Acad. Sci., Trans. 7: 57-58 (nest, prey).
asperata (Fox). Tex., N. Mex., Ariz., Calif. Ecology: Nests in powdery clay-sand or hard

packed soil, 2-3 females sharing same burrow entrance but presumably making
individual cells stored with about 20 prey each. Prey: Procladius sp. near bellus (Lw.);
Heteropsyllus tecana Cwfd., Aphalaroida spp., Paratrioza sp.; Circulifer tenellus
(Bak.), Empoasca abrupta DeLong, Erythroneura sp., Typhlocybinae sp.; Miridae sp.; all
prey were adults.
Crabro asperatus Fox, 1895. Amer. Ent. Soc., Trans. 22: 199. ¢.
Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 293-294, figs. 74-77 (larva).
Biology: Evans, 1964. Ins. Sociaux 11: 71-78, 3 figs. (communal nesting, prey transport).
—Cazier and Mortenson, 1964. Pan-Pacific Ent. 40: 111-114, 1 fig. (nest, prey).
evansi Pate. Ariz. (Tucson).
Moniaecera evansi Pate, 1947. Ent. News 57: 289. 6.

foxiana Pate. Calif.
Moniaecera (Moniaecera) foxiana Pate, 1948. Amer. Ent. Soc., Trans. 74: 49. 2, 6.

pinal Pate. Ariz., Calif.
Moniaecera pinal Pate, 1947. Notulae Nat. 185: 10. d.

Genus HUAVEA Pate
Moniaecera subg. Huavea Pate, 1948. Amer. Ent. Soc., Trans. 74: 58.
Type-species: Moniaecera (Huavea) chontale Pate. Orig. desig.
The morphological features of the female suggest that members of this genus nest in the
ground.

pima Court and Bohart. Ariz. (Santa Cruz and Pima Co’s.).
Huavea pima Court and Bohart, 1966. Pan-Pacifie Ent. 42: 331, figs. 1, 2. 6, 2.

Superfamily SPHECOIDEA 1665

Genus CROSSOCERUS Lepeletier and Brulle

This large genus contains a number of subgenera and numerous species. Some of the sub-
genera are ground-nesters, others nest in pre-existing burrows or cavities in wood, or construct
their own burrows in soft pith of herbs and stems or in rotten wood. Most species prey upon
small Diptera, but some prey upon Homoptera, Trichoptera, Microlepidoptera, Hemiptera, and,
rarely, Mecoptera and Ephemeroptera. So far as known, our North American species do not
store this entire range of prey.

Taxonomy: Pate, 1944 (1943). Lloydia 6: 267-271 (key to subg.). —Lecleregq, 1968. Soc. Roy.
Sci. Liege, Bul. 27: 99-100 (key to xylicolous subg.).

Genus CROSSOCERUS Subgenus CROSSOCERUS Lepeletier and Brulle

Crossocerus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 763.
Type-species: Crabro scutatus Fabricius. Desig. by Westwood, 1839.
Crabro subg. Microcrabro Saussure, 1892. Jn Grandidier, Hist. Nat. Madagascar 20: 574. N.
syn. (R. C. Miller).
Type-species: Crabro (Microcrabro) micromegas Saussure. Monotypic.
Stenocrabro Ashmead, 1899. Canad. Ent. 31: 216.
Type-species: Crabro planipes Fox. Orig. desig.
Synorhopalum Ashmead, 1899. Canad. Ent. 31: 218.
Type-species: Crabro decorus Fox. Orig. desig.
Ischnolynthus Holmberg, 1903. Mus. Nac. Buenos Aires, An. 9: 472.
Type-species: Ischnolynthus foveolatus Holmberg. Monotypic.
Crossocerus subg. Yuchiha Pate, 1944 (1943). Lloydia 6: 272. N. syn. (R. C. Miller).
Type-species: Crossocerus (Yuchiha) xanthochilos Pate. Orig. desig.

Relatively little is known of the biology of members of this subgenus. Some species are known
to nest in the ground; this is substantiated by some of the morphological features of the females.
However, a few species are known to nest in pre-existing burrows in wood above the ground. It
is possible that many species utilize pre-existing burrows, modifying them as required as nesting
sites. If this is so, perhaps the few species which have been reported as nesting in wood may be
found eventually to nest in the ground also.

chromatipus Pate. Nev., Calif., Oreg., Wash.
Crabro pictipes Fox, 1895. Amer. Ent. Soc., Trans. 22: 187. ¢. Preocce.
Crossocerus (Crossocerus) chromatipus Pate, 1944 (1943). Lloydia 6: 280. N. name.
decorus (Fox). Nebr., Colo., Utah, west. Tex. to south. Calif., south in Mexico to Chiapas.
Ecology: Nests in sand, clay, and softer spots of rocky root-shrouded stream banks;
pre-existing burrows of other insects are used and new side branches are constructed,
resulting in twisted burrows 8.5-15.0 em long, weakly branched, with tight clusters or
linear series of cells grouped around end of burrow; up to 9 cells per nest with 8-29
prey per cell. Prey: Chloropidae, Pipunculidae, Stratiomyidae, Ceratopogonidae,
Empididae, Sciaridae, Dolichopodidae, Tachinidae, Simuliidae, Tephritidae, Agromyzidae,
Chironomidae.
Crabro decorus Fox, 1895. Amer. Ent. Soc., Trans. 22: 200. 2.

Biology: Cockerell, 1898. Davenport Acad. Nat. Sci., Proc. 7: 148 (nest). —Smith, 1908. Nebr.
Univ. Studies 8: 395 (nest).

elongatulus elongatulus Vander Linden. Transit. and U. Austr. Zones, N.S., Que., Ont., Maine
to Va. west to Utah and Wash; Palaearctie south of Arctic Circle including North
Africa; adventive in Argentina. Ecology: In Europe it is recorded as nesting in soil or
wood, and sometimes in pre-existing burrows of wood-boring insects; nests have been
found in such odd habitats as soil clinging to roots of fallen trees, old mortar of
buildings, crevices in stone walls, and in cracks of rock outcrops; communal nesting has
been reported once. Parasite: In Europe a species of Macronychia has been recorded.
Prey: Chloropidae, Lauxaniidae, Scatopsidae, Cecidomyiidae, Empididae, Phoridae,
Dolichopodidae, Syrphidae, Agromyzidae, Milichiidae, Tachinidae, Ephydridae,

1666 Hymenoptera in America North of Mexico

Drosophilidae, Diastatidae, Calliphoridae, Stratiomyidae, Muscidae, Sciaridae, and one
record of Aphidoidea. Another subspecies occurs in Sicily.

Crossocerus elongatulus Vander Linden, 1829. Nouv. Mem. Acad. Roy. Sci., Belles-Lettres
Bruxelles 5: 64. 6, °.

Crossocerus varipes Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 773. 2, d.

Crossocerus pallidipalpis Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 779. °, d.

Crossocerus morio Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 781. 2, d.

Crossocerus affinis Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 781. 6.

Crossocerus luteipalpis Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 785. ¢.

Crossocerus annulatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 787. °.

Crabro proximus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 156. ¢. This is a
questionable synonym.

Crabro hyalinus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 161. °.

Crabro transversalis Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 162. d.

Crabro obliquus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 167. 2, d.

Crabro propinquus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 168. d.

Crossocerus elongatus Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 1938. Lapsus or
emend.

Crabro scutellaris Smith, 1851. List Brit. Anim. Brit. Mus. 6: 121. 2. Preoce.

Crabro sulcus Fox, 1895. Amer. Ent. Soe., Trans. 22: 187. 9.

Ischnolynthus foveolatus Holmberg, 1903. Mus. Nat. Buenos Aires, An. 9: 472. 6.

Stenocrabro plesius Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 472. d.

Crossocerus elongatus berlandi Richards, 1928. Soe. Ent. France, Bul., p. 223. d.

Biology: Leclercq, 1954. Monog. Syst., Phylogen., Zoogeogr. Crabron., Hym. p. 307 (nest,
prey, refs. to biol. in Europe).
eriogoni (Rohwer). Alta., Colo., N. Mex., Ariz.; Mexico (Chihuahua, Coahuila, Zacatecas,
Durango, Mexico). Ecology: Nests in softer spots of rocky root-shrouded stream banks;
apparently nests in pre-existing burrows of other insects and makes a sometimes
branched burrow 8-16 cm long; makes up to 8 cells per nest in a small cluster at end of
burrow and stores 18-22 prey per cell. Prey: Pipunculidae, Chloropidae, Tephritidae,
Empididae, Chamaemyiidae.
Crabro (Crossoceros (!)) eriogoni Rohwer, 1908. Ent. News 19: 256. 9.
lentus (Fox). Que. west to B. C., Yukon Terr., N. W. T., Alaska south to Fla. and N. Mex.,
apparently not occurring west of Rocky Mts. in U. S. Ecology: Nests in vertical banks
of sand and in stony, root-shrouded banks of firm clayey sand; pre-existing burrows are
used and new side branches are constructed with clusters of cells located around, off to
the side, or at the end of the main burrow; burrows are 7.0-10.5 cm long, linear or
branched; up to 12 cells per nest with 5-30 prey per cell, newly captured prey stored at
end of burrow, not in cell. Prey: Simuliidae, Chloropidae, Empididae, Ceratopogonidae,
Chironomidae, Agromyzidae.
Crabro scutellatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 341. 2. Preoce.
N. syn. (R. C. Miller).
Crabro lentus Fox, 1895. Amer. Ent. Soc., Trans. 22: 190. 9.
Crabro scutellifer Dalla Torre, 1897. Cat. Hym., v. 8, p. 625. N. name. N. syn. (R. C. Miller).

Biology: Peckham and Peckham, 1905. Wasps, Social and Solitary, p. 101 (nest, prey).

maculiclypeus (Fox). Newfoundland west to B. C., Yukon Terr., N. W. T., Alaska, south to Pa.
and Kans. in the east, and to N. Mex. and Calif. in the west. Ecology: Nests in sparsely
vegetated sand, burrows 4.0-11.5 em long, extremely variable in shape, but main burrow
usually enters at acute angle to surface; entrances usually concealed under small sand
ledges or leaves, at bases of plants, or in depressions; cells arranged in clusters at the
end of or off to the side of the main burrow with up to 9 cells per nest, each cell with
5-20 prey; pre-existing burrows are often used but new burrows are begun in small
crevices or depressions in sand; newly captured prey sometimes stored in open cell at
end of burrow, sometimes at end of burrow in a section only slightly or not at all
widened. Prey: Helobia sp., Molophilus sp.; Palpomyia sp.; Cricotopus sp., Hydrobaenus
sp., Chironomidae sp.; Bradysia sp.; Drapetis sp., Hilara femorata Lw., Platypalpus

Superfamily SPHECOIDEA 1667

holosericus Mel., P. cellarius Mel., P. xanthopodus Mel., Bicellaria pectinata Mel.,
Rhamphomyia sp. Chrysotus sp., Peloropeodes sp., Thrypticus willistoni (Whlr.);
Conioscinella melancholica Beck., C. sp., Meromyza sp., Oscinella sp., Chlorops sp.;
Agromyza sp., Liriomyza sp., Melanagromyza sp., Ophiomyia labiatarum Her.,
Phytobia sp., Phytomyza sp.; Psila angustata Cr., P. rosae F.; Rhagoletis fausta (O. S.);
Plunomia elegans Curr.; Coenosiinae sp.; preferred prey are Brachycera and
acalyptrate Cyclorrhapha. Predator: Philanthus pacificus Cr.

Crabro maculiclypeus Fox, 1895. Amer. Ent. Soc., Trans. 22: 189. 2, d.

Thyreopus (Crossocerus) daeckei Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 51. °.

Biology: Kurezewski, Burdick and Gaumer, 1969. N. Y. Ent. Soc., Jour. 77: 92-104, 8 figs.
(nest, prey transport). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 491-492 (nest, prey).
minimus (Packard). N. B. to Ont., N. W. T., Maine to N. C. west to S. Dak. and Tex.
Blepharipus minimus Packard, 1867. Ent. Soc. Phila., Proe. 6: 3877. 2, d.
Crabro propinquus Fox, 1895. Amer. Ent. Soc., Trans. 22: 189. 2. Preoce.
Crossocerus (Crossocerus) pelas Pate, 1944 (1943). Lloydia 6: 280. N. name.
planifemur Krombein. N. Y. to N. C., W. Va. Ecology: Nests in abandoned beetle borings in
logs.
Crossocerus (Crossocerus) planifemur Krombein, 1952. Ent. Soc. Wash., Proc. 54: 181. °,
(3 misdet.).
Crossocerus (Crossocerus) spangleri Krombein, 1962. Biol. Soc. Wash., Proc. 75: 16. d, (¢
misdet.). N. syn. (R. C. Miller).

Biology: Krombein, 1952. Ent. Soc. Wash., Proc. 54: 182 (nest).
planipes (Fox). Newfoundland to B. C., Yukon, N. W. T., Alaska, Maine to Ga. west to Wash.

and Calif. Ecology: Usually nests in vertical sand or clay banks, but has been found
nesting in level or gently sloping bare soil; nests are 10-40 em long, straight or curved,
rarely branched; cells arranged in irregular linear series or tight clusters, up to 7 cells
per nest, each new group of cells started at a point farther along main burrow than
most recent cell of preceding group; females investigate and eventually renovate
pre-existing burrows and holes; 6-20 prey stored per cell; newly captured prey stored in
cell at end of burrow or at unwidened end of burrow. Prey: Procladius bellus (Lw.);
Chelifera sp., Chersodromia sp., Drapetis sp., Hilara testacea Lw., Leptopeza borealis
Zett., Oedalia ohioensis Mel., Platypalpus holosericus Mel.; Rhamphomyia pusio Lw.,
Trichina nura Mel.; Chrysotus sp.; Psila sp.; Homoneura disjuncta (Johns.), Minettia
sp.; Meromyza sp., Parectecephala sanguinolenta (Lw.).

Crabro incavus Fox, 1895. Amer. Ent. Soc., Trans. 22: 188. °.

Crabro planipes Fox, 1895. Amer. Ent. Soe., Trans. 22: 193. 6.

Crabro (Crossoceros (!)) cockerelli Rohwer, 1908. Ent. News 19: 255. “2?” = 6, 6.

Taxonomy: Bohart and Menke, 1976. Sphecid wasps of world, p. 402 (synonymy).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 21 (nest). —Krombein, 1964. Brooklyn
Ent. Soc., Bul. 58: 119-120 (nest, prey).
similis (Fox). Ont., Vt. and N. Y. to Fla., Mich., Wis., Ohio, Ill., Tenn.
Crabro similis Fox, 1895. Amer. Ent. Soc., Trans. 22: 191. °.
Stenocrabro flavitrochantericus Viereck, 1904. Amer. Ent. Soe., Trans. 30: 242. °.
wesmaeli Vander Linden. N. W. T.; widespread in Europe, Russia, Manchuria and Japan.
Ecology: In Europe nests in sand, loess and clay cliffs, cells seattered 2-5 cm below
surface, burrows linear or simply branched, up to 9 cells per nest, stores 8-39 prey per
cell; newly captured prey stored in open cell at end of burrow. Prey: In Europe
Ceratopogonidae, Chironomidae, Cecidomyiidae, Sciaridae, Agromyzidae, Ephydridae,
Tipulidae, Chloropidae, Empididae, Chamaemyiidae, Simuliidae; Psocoptera;
Cicadellidae; Anthocoridae.
Crabro Wesmaeli Vander Linden, 1829. Nouv. Mem. Acad. Roy. Sci., Belles-Lettres
Bruxelles 5: 65. 2, d.
Ceratocolus Ziegleri Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 748. °.
Ceratocolus maurus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 748. °.

1668 Hymenoptera in America North of Mexico

Biology: Leclereq, 1954. Monog. Syst., Phylogen., Zoogeogr. Hym. Crabron., pp. 307-308 (nest,
prey, refs. to biol. in Europe). —Tsuneki, 1960. Fac. Liberal Arts, Fukui Univ., Mem., Ser.
2, No. 10, pp. 47-48 (nest, prey).
xanthochilos Pate. Md. to Ga. west to Tex., Mo., Kans., Ill. Ecology: Nests in clay bank. Prey:
Condylostylus sp.
Crossocerus (Yuchiha) xanthochilos Pate, 1944 (1943). Lloydia 6: 274. 9.

Taxonomy: Krombein, 1952. Amer. Ent. Soc., Trans. 78: 96. 6.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 21 (nest; misdet. as scutellatus).

xanthognathus (Rohwer). Ariz.; Mexico (Chihuahua).
Thyreopus (Crossocerus) xanthognathus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 566. 2.

Genus CROSSOCERUS Subgenus ABLEPHARIPUS Perkins

Ablepharipus Perkins, 1913. Ent. Soe. London, Trans., p. 390.
Type-species: Crabro podagricus Vander Linden. Monotypic.

The European podagricus usually nests in abandoned burrows of wood-boring Coleoptera, and
provisions the cells with small nematocerous and acalyptrate Diptera. A female of the rare
North American unicus was collected on the trunk of a dead hemlock, where, presumably, it
nested in an abandoned beetle boring.

unicus (Patton). Conn., N. Y., Pa., Mich., Ind., Minn., Mont., Man.
Blepharipus unicus Patton, 1879. Canad. Ent. 11: 214. °.
Stenocrabro nelli Viereck, 1904. Amer. Ent. Soc., Trans. 30: 241. °.

Genus CROSSOCERUS Subgenus EPICROSSOCERUS Ashmead

Epicrossocerus Ashmead, 1899. Canad. Ent. 31: 215.
Type-species: Crabro insolens Fox. Orig. desig.
insolens (Fox). N. Y., Mich., Mo., Colo., Utah, Ariz., Calif., Oreg. Ecology: Nests in Sambucus

stems. Parasite: Diomorus zabriskiei Cr. Prey: Pachyneuron sp.; Tetrastichus
pattersonae Full., T. sp.

Crabro insolens Fox., 1895. Amer. Ent. Soe., Trans. 22: 192. 2.

Crabro (Epicrossocerus) wniversitatis Rohwer, 1909. Ent. News 20: 152. 2. N. syn. (R. C.
Miller).

Epicrossocerus raui Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 99. 2. N. syn. (R. C. Miller).

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest, prey, parasite).
Genus CROSSOCERUS Subgenus HOPLOCRABRO Thomson

Crabro subg. Hoplocrabro Thomson, 1874. Hym. Seand., v. 3, p. 277.
Type-species: Crabro quadrimaculatus Fabricius. Monotypic.

The European quadrimaculatus nests in sandy soil and provisions with nematocerous and
brachycerous Diptera and Trichoptera.

Taxonomy: Pate, 1942. Canad. Ent. 74: 177-185, 5 figs.
angelicus (Kincaid). Nebr., Alta., Mont. and Wyo. west to B. C. and north. Calif. Prey: Muscoid
Diptera.
Crabro angelicus Kincaid, 1900. Ent. News 11: 358. “2” = 6.
Crabro (Hoplocrabro) vierecki Smith, 1908. Nebr. Univ. Studies 8: 401. 2.
Crabro (Hoplocrabro) boulderensis Rohwer, 1909. Ent. News 20: 323. 6.
Crabro (Hoplocrabro) spinibuccus Viereck, 1909. Ent. Soc. Wash., Proc. 11: 44. 9.

Biology: Krombein, 1967. U. S. Dept. Agr., Monog. 2, Sup. 2: 420 (prey).

Genus CROSSOCERUS Subgenus BLEPHARIPUS Lepeletier and Brulle

Blepharipus Lepeletier and Brulle, 1834. Soe. Ent. France, Ann. 3: 728.
Type-species: Blepharipus nigrita Lepeletier and Brulle. Desig. by Ashmead, 1899.
Crabro subg. Coelocrabro Thomson, 1874. Hym. Scand., v. 3, p. 262.

Superfamily SPHECOIDEA 1669

Type-species: Crabro pubescens Shuckard. Desig. by Richards, 1935.
Dolichocrabro Ashmead, 1899. Canad. Ent. 31: 216.
Type-species: Dolichocrabro Wickhamti Ashmead. Orig. desig.
Acanthocrabro Perkins, 1913. Ent. Soe. London, Trans., p. 391. N.. syn. (R. C. Miller).
Type-species: Crabro vagabundus Panzer. Monotypic.
Crossocerus subg. Nothocrabro Pate, 1944 (1943). Lloydia 6: 314. N. syn. (R. C. Miller).
Type-species: Crabro nitidiventris Fox. Orig. desig.
Crossocerus subg. Stictoptila Pate, 1944 (1943). Lloydia 6: 315. N. syn. (R. C. Miller).
Type-species: Crabro confertus Fox. Orig. desig.
Crossocerus subg. Neoblepharipus Leclercq, 1968. Soc. Roy. Sci. Liege, Bul. 27: 98. N. syn.
(R. C. Miller).
Type-species: Crossocerus (Neoblepharipus) potosus Leclercq. Orig. desig.
Crossocerus subg. Fentis Tsuneki, 1971. Etizenia 51: 13. N. syn. (R. C. Miller).
Type-species: Crossocerus (Fentis) quinquedentatus Tsuneki. Orig. desig.
Crossocerus subg. Bnunius Tsuneki, 1971. Etizenia 51: 15. N. syn. (R. C. Miller).
Type-species: Crossocerus (Bnunius) domicola Tsuneki. Orig. desig.

These species nest in the soft pith of plant stems or twigs, or in decaying wood. Most species
make a series of linear cells separated by partitions of particles of the substrate.

Revision: Pate, 1944 (1943). Lloydia 6: 290-312 (N. Amer. spp.).
annulipes annulipes (Lepeletier and Brulle). N. S. to B. C. south to Ga., Utah, Nev. and Calif;

widely distributed in Palaearctic Region except Japan. Ecology: In North America nests
in rotting stumps and logs, and in old anobiid burrows in structural timber, stores 4-46
prey per cell. Prey: Alebra albostriella (Fall.), Riboutiana ? sp., Empoa ? sp., Empoasca
alboneura Gill., E. bifurcata DeL., E. bipunctata (Osh.), E. birdii Godg., E. erigeron
DeL., E. fabae Harr., E. gelbata DeL. and Dav., E. livingstoni Gill., E. maligna Walsh,
E. obtusa Walsh, E. patula DeL., E. pergandei Gill. E. solana DeL., E. spp.,
Edwardsiana rosae (L.), Typhlocyba gillettei Van D., T. pomaria McAtee, T. australis
Frogg., T. modesta Gibson, T. melite Mcatee, T. sp. or Ossiannilssonola sp.,
Erythroneura comes (Say), E. confirmata McAtee, E. dowelli Beamer, E. lawsoniana
Baker, E. tricincta Fitch, E. vulnerata Fitch, E. ziczac Walsh, E. lawsoni Rob., E. vitis
(Harr.), E. hartii (Gill.), E. magnacalx Beamer, E. elegans McAtee, E. rubra (Gill.), E.
albescens Beamer; adults are the preferred prey but some nymphs are stored; many
other leafhoppers are recorded as prey in Europe; Miridae sp.; Chironomidae sp.
Another subsp. occurs in Japan.

Blepharipus annulipes Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 729. °.

Crossocerus gonager Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 785. °.

Crabro nigritus Gimmerthal, 1836. Soc. Imp. Nat. Moscow, Bul. 9: 435.

Crabro ambiguus Dahlbom, 1842. Dispos. Method. Spec. Scand. Ins. Hym., p. 14. ?.

Crabro (Crossocerus) capito Zeller, 1845. In Dahlbom, Hym Europaea, v. 1, p. 524.

Blepharipus parkeri Banks, 1921. Ent. Soc. Amer., Ann. 14: 17. 2.

Crabro (Blepharipus) davidsoni Sandhouse, 1938. Ent. Soc. Amer., Ann. 31: 1. 2, d.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 106, figs. 69-74 (larva).

Biology: Davidson and Landis, 1938. Ent. Soc. Amer., Ann. 31: 5-8, 3 figs. (nest, prey, life
cycle). —Krombein, 1958. Biol. Soc. Wash., Proc. 71: 26 (nest, prey, life cycle). —Michener,
1971. Kans. Ent. Soe., Jour. 44: 405-407 (nest, prey).

barbipes (Dahlbom). Que. to B. C., Alaska, Yukon Terr. south to N. C., Tenn., and S. Dak., in

mts. to N. Mex. and Calif.; north. Palaearctic, Lapland and Holland to Japan. Ecology:
Nests in hole in wooden shingle and in twigs of Salix and Sambucus, and in dead
standing tree. Prey: Empoasca sp., possibly obtusa Walsh or patula DeL.; in Japan
preys upon small Diptera, storing 3-6 prey per cell.

Crabro barbipes Dahlbom, 1845. Hym. Europaea, v. 1, p. 521. d.

Crabro ater Cresson, 1865. Ent. Soc. Phila., Proc. 4: 477. 2. Preoce. N. syn. (R. C. Miller).

Crabro hirtipes Morawitz, 1866. Acad. Imp. Sci. St. Petersburg, Bul. 9: 258. d.

Dolichocrabro Wickhamii Ashmead, 1899. Canad. Ent. 31: 215. ¢. N. syn. (R. C. Miller).

Dolichocrabro wickhami Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 133. ¢. Preoce. N. syn.
(R. C. Miller).

1670 Hymenoptera in America North of Mexico

Crossocerus (Blepharipus) pammelas Pate, 1944 (1943). Lloydia 6: 299. N. name. N. syn.
(R. C. Miller).

Biology: Steyskal, 1944. Brooklyn Ent. Soc., Bul. 39: 170 (nest, prey). —Tsuneki, 1960. Fac.
Lib. Arts Fukui Univ., Mem. Ser. 2, Nat. Sci. (10) 1: 42 (nest, prey).
cinctipes (Provancher). Newfoundland, N. B., N. S. west to Alaska, Yukon Terr., N. W. T.,
south to Md., Mich., Minn. and in the mts. to N. Mex. and Calif.
Plepharipus(!) cinctipes Provancher, 1882. Nat. Canad. 13: 133. ¢.
Crabro niger Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 419. 2. Preoce.
Crabro nigror Fox, 1895. Amer. Ent. Soc., Trans. 22: 196. 2.
Crabro nigrior Fox, 1896. Amer. Ent. Soc., Trans. 23: 80. Emend.
Crabro servus Dalla Torre, 1897. Cat. Hym., v. 8, p. 626. N. name.
Stenocrabro cinctitarsis Ashmead, 1901. Psyche 9: 185. d.
Blepharipus columbiae Bradley, 1906. Canad. Ent. 38: 380. 2.
Thyreopus (Blepharipus) utensis Mickel, 1916. Amer. Ent. Soc., Trans. 42: 421. °.
Thyreopus (subgenus?) stygius Mickel, 1916. Amer. Ent. Soc., Trans. 42: 422. 9.
fergusoni Pate. Utah, Nev., Calif., Oreg. Ecology: Nests in Sambucus stems. Parasite:
Diomorus zabriskiei Cr.; Amobia floridensis (Tns.), Macronychia sp. Prey:
Stratiomyidae ? sp.; Empididae; Muscoid Diptera.
Crossocerus (Blepharipus) fergusoni Pate, 1944 (1943). Lloydia 6: 307. d.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 107, figs. 97-98 (larva). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 165, fig. 61 (larva).

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1022 (prey). —Parker and Bohart, 1966.
Pan-Pacifie Ent. 42: 94 (nest, parasites).
harringtonii (Fox). Que. west to B. C., south to Ga., Tenn., Kans. and Mont. Ecology: Reared
from branches of white oak.
Crabro Harringtonii Fox, 1895. Amer. Ent. Soc., Trans. 22: 195. 9.

Taxonomy: Krombein, 1952. Ent. Soc. Wash., Proc. 54: 184. ¢.
impressifrons (Smith). Ont., Mass. south to Fla., west to Nebr. and Ark. Ecology: Nests in

rotten log and in dense frass beneath tight bark of dead American elm logs, and in
hickory; one nest 10 cm long, branching and with 6 cells, some in linear series. Prey:
Condylostylus sp., Dolichopodidae sp.; Tephritidae; Empididae; Syrphidae;
Chironomidae; Trichoptera sp.

Crabro tibialis Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 340. 2. Preoce.

Crabro pusillus Harris, 1835. In Hitchcock, Rpt. Geol., Mineral. Bot. Zool. Mass., p. 68.
Nom. nud.

Crabro impressifrons Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 417. N. name.

Blepharipus scutellatus Packard, 1867. Ent. Soe. Phila., Proc. 6: 375. 3. Preoce.

Blepharipus Harristi Packard, 1867. Ent. Soe. Phila., Proc. 6: 376. d.

Crabro (Blepharipus) tridentatus Rohwer, 1909. Ent. News 20: 150. 2, 3. Preoce.

maculipennis (Smith). Que. to B. C. south to Va., W. Va., Ill., Minn. and in mts. to N. Mex. and

Calif. Ecology: Nests in cavities in log and old apple tree. Prey: Nephrotoma tenuis
(Lw.), Pales ferrugineus (F.), Tipulidae sp.

Blepharipus maculatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 730. @.
Preoce.

Crabro pictus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 417. N. name. Preoce.

Crabro maculipennis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 417. N. name.

Crabro confertus Fox, 1895. Amer. Ent. Soc., Trans. 22: 181. 2, 6.

Crabro ventralis Fox, 1895. Amer. Ent. Soe., Trans. 22: 183. ?.

Crabro canonicola Viereck, 1908. Amer. Ent. Soc., Trans. 33: 402. 2.

Crabro albertus Carter, 1925. Canad. Ent. 57: 135. 3.

Biology: Erikson, 1940. Brooklyn Ent. Soe., Bul. 35: 172 (nest, prey). —Krombein, 1951. U. S.
Dept. Agr., Monog. 2: 1023 (prey).
maculitarsis (Cameron). Ariz., Tex.; Mexico (Guerrero, Morelos, Chiapas). Ecology: Reared
from Quercus.
Crabro maculitarsis Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 154. 9.

Superfamily SPHECOIDEA 1671

melanius (Rohwer). Alta., Colo., N. Mex., Ariz.; Mexico (Chihuahua, Durango, Morelos).
Thyreopus (Blepharipus) melanius Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 565. 2, ¢.
nigricornis (Provancher). Newfoundland to B. C., Yukon and Alaska, south to N. C., Tenn.,
Nebr., in mts. south to N. Mex. and Calif. Ecology: Nests in Sambucus stems and in
twig of living box elder tree; one nest 20.5 em long, linear, contained 11 cells each with
6-14 prey per cell. Prey: Primarily Dolichopodidae and Empididae especially
Platypalpus sp., but also Ceratopogonidae, Mycetophilidae, Chironomidae, Tipulidae,
Phoridae, Chloropidae, other acalyptrate Cyclorrhapha, Muscidae and Anthomyiidae.
Blepharipus nigricornis Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 294.
3.

Biology: Pate, 1944 (1943). Lloydia 6: 306-307 (nest, prey).
nitidiventris (Fox). Que. and Ont. to Ga. west to Minn. and Tex. Prey: Nephrotoma virescens
Lw., Tipulidae sp.
Crabro nitidiventris Fox, 1892. Ent. News 3: 9. 9.
Taxonomy: Krombein, 1953. Wasmann Jour. Biol. 10: 338. 2.

Biology: Krombein, 1967. U. S. Dept. Agr., Monog. 2, Sup. 2, p. 420 (prey).
stictochilos Pate. Mass. to Ga., Mich., Ill., Ark. Ecology: Nests in pith of green Hibiscus stem.
Prey: Diptera.
Crossocerus (Blepharipus) stictochilos Pate, 1944 (1943). Lloydia 6: 304. 2, ¢.
Biology: Krombein, 1964. Biol. Soc. Wash., Proc. 77: 90 (nest, prey, cocoon, life cycle).
stricklandi Pate. Alta., B. C., Wyo., Colo., Utah.
Crossocerus (Blepharipus) stricklandi Pate, 1944 (1943). Lloydia 6: 301. ¢.
tarsalis (Fox). Que., Ont., Maine south to Ga., west to Tenn., Mo., Mich., Colo., Idaho.
Crabro tarsalis Fox, 1895. Amer. Ent. Soc., Trans. 22: 193. 6.

Genus TRACHELIODES Morawitz

Crabro subg. Brachymerus Dahlbom, 1845, Hym. Europaea, v. 1, p. 519. Preoce.

Type-species: Crabro (Brachymerus) Megerlei Dahlbom. Monotypic.
Tracheliodes Morawitz, 1866. Acad. Imp. Sci. St. Petersburg, Bul. 9: 249.

Type-species: Brachymerus megerlei Dahlbom. Desig. by Ashmead, 1899.
Fertonius Perez, 1892. In Ferton, Soc. Linn. Bordeaux, Actes 44: 341.

Type-species: Crossocerus luteicollis Lepeletier and Brulle. Desig. by Pate, 1937.

Our species nest in plant stems or borings in wood, but one European species is known to nest
in soil, another in abandoned beetle borings. Our species prey upon worker ants of the genus
Liometopum; European species use workers of both Liometopum and Tapinoma.

Revision: Pate, 1942. Lloydia 5: 222-224, 7 figs. (N. Amer. spp.).
amu Pate. N. Mex., Ariz. Ecology: Nests in borings in wood, constructs a brood chamber in
which 1-3 wasps develop, stores as many as 42 prey per egg. Prey: Liometopum

occidentale luctuosum Whlr. workers.
Tracheliodes amu Pate, 1942. Lloydia 5: 235. ¢.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 290-291, figs. 104-109 (larva).
Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 257-259, figs. 68, 69 (nest, prey,
life cycle, cocoon).
foveolineatus (Viereck). Colo., Oreg., Calif. Ecology: Nests in Sambucus stems. Prey:
Liometopum occidentale luctuwosum Whlr. workers.
Crabro (Cuphopterus) foveolineatus Viereck, 1909. Ent. Soc. Wash., Proc. 11: 44. °.
Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 95 (nest, prey).
hicksi Sandhouse. Colo., Ariz. Prey: Liometopum sp. workers.
Tracheliodes hicksi Sandhouse, 1936. Ent. News 47: 2. °.

Biology: Hicks, 1936. Ent. News 47: 4-7 (prey capture).

1672 Hymenoptera in America North of Mexico

Genus CRABRO Fabricius

Crabro Fabricius, 1775. Systema Ent., p. 373. Preoce., but placed on Official List of
Generic Names of Internat]. Comn. Zool. Nomencl., 1936., Op. 144.
Type-species: Vespa cribraria Linnaeus. Desig. by Internat]. Comn. Zool. Nomencl.,
1943.
Carabro Say, 1823. West. Quart. Rptr. 2: 78. Lapsus.
Crabro subg. Thyreopus Lepeletier and Brulle, 1835. Soc. Ent. France, Ann. 3: 519.
Type-species: Sphex cribrarius (Linnaeus). Desig. by Westwood, 1839.
Crabro subg. Thyreocnemus Costa, 1871. Mus. Zool. Napoli, Ann. 6: 64.
Type-species: Crabro pugillator Costa. Monotypic.
Anothyreus Dahlbom, 1845. Hym. Europaea, v. 1, p. 526.
Type-species: Anothyreus lapponicus of Dahlbom. Desig. by Ashmead, 1899.
Paranothyreus Ashmead, 1899. Canad. Ent. 31: 213.
Type-species: Crabro hilaris Smith. Orig. desig.
Synothyreopus Ashmead, 1899. Canad. Ent. 31: 213.
Type-species: Thyreopus tumidus Packard. Orig. desig.
Pemphilis Pate, 1944. Amer. Midland Nat. 31: 340.
Type-species: Vespa cribraria Linnaeus. Orig. desig.
Dyscolocrabro Pate, 1944. Amer. Midland Nat. 31: 349.
Type-species: Crabro chalybeus Kohl. Orig. desig.
Agnosicrabro Pate, 1944. Amer. Midland Nat. 31: 349.
Type-species: Crabro occultus Fabricius. Orig. desig.
Hemithyreopus Pate, 1944. Amer. Midland Nat. 31: 349.
Type-species: Crabro (Ceratocolus) Loewi Dahlbom. Orig. desig.
Parathyreopus Pate, 1944. Amer. Midland Nat. 31: 349.
Type-species: Crabro filiformis Radoszkowski. Orig. desig.
Pemphilis subg. Noruwmbega Pate, 1947. Notulae Nat. 185: 12.
Type-species: Thyreopus argus Packard. Orig. desig.

Subgenera have frequently been recognized in this large genus, but Bohart and Menke believe
that on morphological grounds it is preferable to recognize only species groups. This concept is
substantiated by a consideration of the ethology, for clearcut differences have not been demon-
strated for the several species groups. These wasps nest in the soil, usually constructing mul-
ticellular nests, some nesting gregariously, others are solitary. The prey consists of Diptera and,
frequently, each species has preferences for flies belonging to one or another suborder or sec-
tion.

Biology: Kurezewski and Acciavatti, 1968. N. Y. Ent. Soc., Jour. 76: 196-212 (nesting behavior
of N. Amer. spp.).

SPECIES GROUP HILARIS
This is the equivalent of Paranothyreus Ashm.

aequalis Fox. U.S. east of 100th meridian, north to N. J. and N. Dak., south to Fla.
Crabro aequalis Fox, 1895. Amer. Ent. Soc., Trans. 22: 164. 9.
Paranothyreus rugicollis Viereck, 1904. Amer. Ent. Soc., Trans. 30: 241. d.
Thyreopus (subgenus?) knoxensis Mickel, 1916. Amer. Ent. Soc., Trans. 42: 424. 6.

cingulatus (Packard). N. J. to Ala. west to Wis., Nebr., Tex.; Mexico. Ecology: Nests in
vertical sand banks, makes 2-8 cells per nest, stores 11-20 flies per cell. Prey:
Paralimna punctipennis (Wied.).
Thyreopus cingulatus Packard, 1867. Ent. Soc. Phila., Proc. 6: 366. ¢, (2 misdet.).
Crabro clarconis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 213. °.

Biology: Rau and Rau, 1918. Wasp Studies Afield, pp. 96-101, figs. 24-26 (nest, prey
transport). —Rau, 1938. Ent. Soc. Amer., Ann. 31: 548.
cognatus Fox. Alta., U. S. except Pacific States, N. H. to Ga. west to Mont., Utah and Tex.
Crabro cognatus Fox, 1895. Amer. Ent. Soc., Trans. 22: 178. 2, 6.
hilaris Smith. N. J. to Fla., Ill., Nebr., Tex.
Crabro hilaris Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 416. 2.

Superfamily SPHECOIDEA 1673

rufibasis (Banks). Fla. Ecology: Nests in vertical face of sand pit. Prey: Furcifera pictipennis
(Wied.), Psilocephala festina (Coq.), P. notata (Wied.); Bombyliidae sp.; Chrysotus
discolor Lw., Condylostylus caudatus (Wied.), C. chrysoprasi (Wlkr.), C. graenicheri
(Van D.), Plagioneurus wnivittatus Lw.; Allograpta obliqua (Say), Baccha costata Say,
B. sp. near loewi Sedman, Mesograpta spp., Mixogaster sp., Toxomerus sp.; Euxesta
basalis (Wlkr.), E. eluta Lw., E. notata (Wied.); Xanthaciura insecta (Lw.);
Atrichomelina pubera (Lw.); Homoneura sp., Poecilominettia valida (Wlkr.);
Pholeomyia decorior Steysk., P. dispar (Beck.); Chlorops abdominalis Coq.; Pegomya
gopheri Johns.; Atherigona orientalis Schin., Coenosia spp., Coenosopsia prima Mall.,
Fannia sp., Gymnodia arcuata (Stein), G. cilifera (Mall.), G. debilis (Will.), Limnophora
narona (Wlkr.), Ophyra aenescens (Wied.), Orthellia caesarion (Meig.); Cochliomyia
macellaria (F.), Phaenicia cluvia (Wlkr.); Johnsonia elegans Coq., Ravinia derelicta
(Wlkr.); Acronarista cornuta Reinh., Actia sp., Catharosia nebulosa (Coq.),
Chaetonodexodes vanderwulpi (Tns.), Chaetophlepsis townsendi (Sm.), Clausicella
floridensis (Tns.), Elfia mellissopodis (Coq.), Exoristoides sp., Gaediopsis flavipes Coq.,
Leskiella brevirostris James, Lespesia aletiae (Riley), Livophaga mediocris Ald., Lidella
sp. near thomsoni Hert., Medina sp., Miamimyjia cincta Tns., Paradidyma singularis
(Tns.), Phasiopsis floridana Tns., Pseudochaeta sp. near finalis Reinh.,
Pseudomyothyria ancilla (Wlkr.), Prophryno parviteres (Ald. and Webb.), Trichopoda
plumipes (F.).

Thyreopus rufibasis Banks, 1921. Ent. Soc. Amer., Ann. 14: 17. @.
snowii Fox. N. Y., Md., Va., Minn., Kans.
Crabro snowi Fox, 1896. Amer. Ent. Soce., Trans. 23: 79. 2, ¢.

SPECIES GROUP TUMIDUS
This is Synothyreopus Ashm., in part.

lacteipennis Rohwer. Tex., Colo., Ariz.; Mexico (Nuevo Leon, Durango).
Crabro (Thyreopus) lacteipennis Rohwer, 1909. Ent. News 20: 150. °.

peltista Kohl. Tex., Ariz.; Mexico to Nicaragua.
Crabro (Thyreopus) peltista Kohl, 1888. Zool. Jahrb., Ztschr. f. System. 3: 586. 9, d.
Crabro incertus Fox, 1895. Amer. Ent. Soc., Trans. 22: 174. 2? (¢ misdet.).

tenuiglossa Packard. Ont., Mich., Ill., Minn., N. Dak., S. Dak., Alta.
Crabro tenuiglossa Packard, 1866. Ent. Soc. Phila., Proc. 6: 98. 9.
Thyreopus discifer Packard, 1867. Ent. Soc. Phila., Proc. 6: 363. d.

tumidus (Packard). N. Y. to S. C., Ill., Nebr.
Thyreopus tumidus Packard, 1867. Ent. Soc. Phila., Proc. 6: 364. °, 6.

SPECIES GROUP ADVENA
This is Synothyreopus Ashm., in part.

advena Smith. South. Canada and U.S. east of 100th meridian. Ecology: Nests in hard clay

loam or sand, makes 2 to more than 16 cells per nest, stores 1-10 prey per cell. Prey:
Chrysopilus proximus (Wlkr.), Symphoromyia pluralis (Curr.); Chrysops univittata
Macq.; Syrphus rectus O. S.; Euxesta notata (Wied.), Callopistromyia annulipes
(Macq.), Otitidae sp.; Pegomya finitima Stein, P. lipsia (Wlkr.), Hydrophoria conica
(Wied.), Anthomyiidae sp.; Muscina assimilis (Fall.), Musca autumnalis DeG., M.
domestica L., Coenosia tigrina (F.), Fannia scalaris (F.); Pollenia rudis (F.), Phaenicia
sericata (Meig.), Calliphoridae sp.; Sarcophaga scoparia Pand., S. sp., Senotainia sp.;
Belvosia unifasciata Desv., Aplomyiopsis sp., Lespesia sp., Blondeliini sp.; preferred
prey are calyptrate Cyclorrhapha but Orthorrhapha are also used.

Crabro pegasus Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p. 68.
Nom. nud.

Crabro advena Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 421. 2 (6 misdet.).

Crabro succinctus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 479. 9.

Thyreopus signifer Packard, 1867. Ent. Soc. Phila., Proc. 6: 361. 2, ¢.

Thyreopus pegasus Packard, 1867. Ent. Soc. Phila., Proc. 6: 362. 2, d.

Thyreopus elegans Provancher, 1883. Faune Ent. Canada, Hym., p. 665. 2, d.

Crabro discretus Fox, 1895. Amer. Ent. Soc., Trans. 22: 165. °.

1674 Hymenoptera in America North of Mexico

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 104, figs. 95-96 (larva). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 164, figs. 47, 48 (larva).

Biology: Patton, 1897. Canad. Ent. 29: 248 (nest, prey). —Krombein, 1951. U.S. Dept. Agr.,
Monog. 2: 1017 (prey). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 53 (prey). —Evans,
1960. N. Y. Ent. Soc., Jour. 68: 123-127, fig. 1 (nest, prey transport, egg, larval life).
—Kurezewski and Acciavatti, 1968. N. Y. Ent. Soc., Jour. 76: 200-209, figs. 1-11 (nest, prey
transport, egg). —Kurezewski, Burdick and Gaumer, 1969. N. Y. Ent. Soc., Jour. 77:
152-170, figs. 1-8 (nest, prey transport, egg).

bruneri (Mickel). Nebr., Oreg.
Thyreopus (Synothyreopus) bruneri Mickel, 1916. Amer. Ent. Soc., Trans. 42: 422. 2, 3.

florissantensis Rohwer. Colo. and Wyo. to Oreg. and Wash. Predator: Philanthus pulcher D.

aly.
Crabro (Synothyreopus) florissantensis Rohwer, 1909. Ent. News 20: 149. ¢.
henrici Krombein. N. J., Va.
Thyreopus (Synothyreopus) vierecki Rohwer, 1910. Ent. Soc. Wash., Proc. 12: 50. 6.
Preoce.
Crabro (Synothyreopus) henrici Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1016. N.
name.
hispidus Fox. Oreg., Wash., B. C.
Crabro hispidus Fox, 1895. Amer. Ent. Soc., Trans. 22: 192. °.
vernalis (Packard). Transcont. in N. Amer. in Transit. Zone including Alaska.
Thyreopus vernalis Packard, 1867. Ent. Soc. Phila., Proce. 6: 369. 2.
Crabro (Thyreopus) brachycarpae Rohwer, 1908. Ent. News 19: 252. d.
Crabro (Paranothyreus) gillettei Rohwer, 1908. Ent. News 19: 418. °.
virgatus Fox. Calif., Nev., Oreg., Idaho, Alta.
Crabro virgatus Fox, 1895. Amer. Ent. Soc., Trans. 22: 174. 6.
Crabro veles Carter, 1925. Canad. Ent. 57: 134. d.

SPECIES GROUP THYREOPHORUS
This is Synothyreopus Ashm., in part.

thyreophorus Kohl. Nev., Calif., Oreg.
Crabro (Thyreopus) thyreophorus Kohl, 1888. Zool. Jahrb., Ztschr. f. System. 3: 585, pl. 14.
Onn

SPECIES GROUP CRIBRARIUS
This is Crabro F., sens. str., and Norumbega Pate.

argusinus Bohart. Transcont. in south. Canada and U. S. Ecology: Nests in sand, makes 1-4 or

more cells per nest, stores 10-19 flies per cell. Parasite: Senotainia sp. near trilineata
(Wulp), Phrosinella fumosa Allen ? Prey: Dolichopus coercens Wlkr., D. gladius Van D.,
D. sp., Rhaphium vanduzeei Curr., Argyra albicans Lw., A. calceata Lw., Hercostomus
barbatulus Lw., H. crassicauda Lw., H. frequens Lw., H. ornatus Van D., Liancalus
genualis Lw., Plastoneurus vagans Lw.; Parydra bituberculata Lw., P. borealis Cr., P.
breviceps Lw.; Lispe albitarsis Stein, L. nasoni Stein; Dolichopodidae are the preferred
prey, but some Ephydridae and a few Muscidae are also used.

Crabro argus Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p. 68. Nom.
nud.

Thyreopus argus Packard, 1867. Ent. Soc. Phila., Proc. 6: 359. ¢. Preoce.

Crabro argusinus Bohart, 1976. In Bohart and Menke, Sphecid wasps of world, p. 407. N.
name.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 164, figs. 49, 50 (larva).
Biology: Hartman, 1905. Tex. Acad. Sci., Trans. 7: 44 (prey, nest). —Dow, 1930. Psyche 37:
181 (prey). —Evans, 1960. N. Y. Ent. Soc., Jour. 68: 129-132, fig. 1 (mest, prey, parasites).
conspicuus Cresson. Colo., Nev., Calif., Wash., Alta.

Crabro conspicuus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 480. °.
Crabro medius Fox, 1895. Amer. Ent. Soc., Trans. 22: 167. cd.

Superfamily SPHECOIDEA 1675

cribrellifer (Packard). N. H. to Fla., Ont., Mich., Ind. Prey: Ommatius tibialis Say, Tolmerus
novaescotiae (Macq.).
Thyreopus cribrellifer Packard, 1867. Ent. Soe. Phila., Proc. 6: 358. d.
Thyreopus sinuatus Provancher, 1883. Faune Ent. Canada, Hym., p. 664. 9. Preoce.
Crabro Provancheri Fox, 1895. Amer. Ent. Soc., Trans. 22: 168. N. name.

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1017 (prey). —Kurezewski and
Acciavatti, 1968. N. Y. Ent. Soc., Jour. 76: 198 (prey).

juniatae Krombein. Pa., Va., W. Va. Prey: Hylemya cilicrura (Rond.).
Crabro (Crabro) juniatae Krombein, 1938. Ent. Soc. Amer., Ann. 31: 469. 2.

Biology: Krombein, 1952. Ent. Soc. Wash., Proc. 54: 181 (prey).
largior Fox. U. S., transcont. in Transit. Zone.
Crabro largior Fox, 1895. Amer. Ent. Soc., Trans. 22: 161. 2, d.
latipes Smith. Transcont. in Alaska, Canada, U. S. in Transit. and Canad. Zones. Ecology:

Nests in sand, makes 8 or more cells per nest, stores 6-10 flies per cell. Prey:
Platycheirus peltatus (Meig.); Symphoromyia pluralis Curr.; Dolichopus adultus Van
D., D. nigricornis Meig., D. nodipennis Van D., D. remus Van D., D. socius Lw.,
Gymnopternus spectabilis Lw., Rhaphium armatum Curr., R. crassipes (Meig.); Musca
domestica L., M. autumnalis DeG., Fannia maniceta (Meig.), Muscina assimilis (Fall.),
Phaonia bysia (Wlkr.); Lucilia illustris (Meig.); Aplomyiopsis sp., Icteriophyto tibialis
(Curr.), Oswaldia assimilis (Tns.); Hydrophoria conica (Wied.), Hylemya setigera (Joh.),
H. stratifrons Huck., Pegomya affinis Stein, P. lipsia (Wlkr.); preferred prey are
calyptrate Cyclorrhapha, but Orthorrhapha are also used. Predator: Philanthus zebratus
nitens (Bks.).

Crabro gryphus Harris, 1835. In Hitchcock Rpt. Geol. Mineral. Bot. Zool. Mass., p. 68.
Nom. nud.

Crabro latipes Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 396. d.

Crabro vicinus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 479. 9.

Thyreopus coloradensis Packard, 1867. Ent. Soc. Phila., Proc. 6: 356. d.

Thyreopus elongatus Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 293. d.

Crabro canadensis Dalla Torre, 1897. Cat. Hym., v. 8, p. 585. N. name.

Crabro (Anothyreus) viciniformis Viereck, 1907. Amer. Ent. Soc., Trans. 33: 381. °.

Crabro pratus Carter, 1925. Canad. Ent. 57: 133. 9.

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1017 (prey). —Krombein, 1955.
Brooklyn Ent. Soc., Bul. 50: 17 (prey). —Kurezewski and Acciavatti, 1968. N. Y. Ent. Soc.,
Jour. 76: 199 (prey). —Kurezewski, Burdick and Gaumer, 1969. N. Y. Ent. Soc., Jour. 77:
152-170, figs. 1, 3-8 (nest, prey transport, egg).

monticola (Packard). Alaska, Canada, Maine to Ga. Ecology: Nests in sand or gravel road,
makes 11 to 15 or more cells per nest, stores 3-5 flies per cell. Prey: Tabanus
lasiophthalmus Macq., T. microcephalus 0. S., Stonemyia tranquilla (O. S.), Chrysops
celer O. S., C. venus Phil.; Thereva sp.; preferred prey are tabanid males.
Thyreopus monticola Packard, 1867. Ent. Soc. Phila., Proc. 6: 367. 2.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 163, figs. 43-46 (larva).
Biology: Evans, 1960. N. Y. Ent. Soe., Jour. 68: 127-129, fig. 1 (nest, prey transport, cocoon).
—Pechuman, 1963. N. Y. Ent. Soc., Jour. 71: 218-219 (nest, prey).
pallidus Fox. Mont., Oreg.
Crabro pallidus Fox, 1895. Amer. Ent. Soc., Trans. 22: 163. 2, d.
pleuralis Fox. Nebr., Colo., Wyo., Alta., B. C., Wash., Oreg. Predator: Philanthus zebratus
nitens (Bks.).
Crabro pleuralis Fox, 1895. Amer. Ent. Soc., Trans. 22: 162. °, ¢.
tenuis Fox. Mich. to Colo. and Wash., Alta.
Crabro tenuis Fox, 1895. Amer. Ent. Soc., Trans. 22: 166. 2, d.
villosus Fox. Calif. (Los Angeles Co.).
Crabro villosus Fox, 1895. Amer. Ent. Soc., Trans. 22: 158. 9.

1676 Hymenoptera in America North of Mexico

Genus ENOPLOLINDENIUS Rohwer

The genus occurs only in the New World. Nothing is known of the biology, but morphological
characters of the female suggest that these are ground-nesting species.

Revision: Pate, 1942. Rev. de Ent. 13: 386-421 (New World spp.).

Genus ENOPLOLINDENIUS Subgenus ISKUTANA Pate

Enoplolindenius subg. Iskutana Pate, 1942. Rev. de Ent. 13: 390.
Type-species: Enoplolindenius (Iskutana) georgia Pate. Orig. desig.
robertsoni (Rohwer). Ga., La., Tex., Kans., Nebr., Iowa, III.
Lindenius robertsoni Rohwer, 1920. Ent. Soc. Wash., Proc. 22: 57. d.
Enoplolindenius (Iskutana) georgia Pate, 1942. Rev. de Ent. 13: 393. 6.
Enoplolindenius (Iskutana) ponca Pate, 1942. Rev. de Ent. 13: 395. °.

Genus ENOPLOLINDENIUS Subgenus ENOPLOLINDENIUS Rohwer

Lindenius subg. Enoplolindenius Rohwer, 1911. U.S. Natl. Mus., Proce. 40: 562.
Type-species: Lindenius (Enoplolindenius) clypeatus Rohwer. Orig. desig.

clypeatus (Rohwer). Tex. (Brownsville).
Lindenius (Enoplolindenius) clypeatus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 562. ¢.

Genus ECTEMNIUS Dahlbom

In the North American fauna a number of species nest in the soft pith of living or dead stems
or twigs; the nests of these species consist usually of a linear series of cells separated from each
other by partitions of pith particles. Other species make burrows in decaying wood. None of our
species has been reported as nesting in the ground, as do members of several extralimital sub-
genera, but morphological characters of the females suggest that members of the subgenus
Protothyreopus nest in the ground.

Genus ECTEMNIUS Subgenus PROTOTHYREOPUS Ashmead

Protothyreopus Ashmead, 1899. Canad. Ent. 31: 170.
Type-species: Crabro rufifemur Packard. Orig. desig.

Nothing is known of the biology of this subgenus, but the species probably nest in the ground.

dilectus (Cresson). Transit. and U. Austr. Zones in most of U.S. except west of Sierra Nevada
and Cascade ranges.
Crabro dilectus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 478. 2, d.
Crabro bigeminus Patton, 1879. Canad. Ent. 11: 218. °, d.
Crabro (Protothyreopus) megacephalus Rohwer, 1908. Ent. News 19: 249. 2. Preoce.
Crabro (Protothyreopus) dilectiformis Rohwer, 1909. Ent. News 20: 146. °.
Crabro (Protothyreopus) crassiceps Mickel, 1916. Amer. Ent. Soc., Trans. 42: 425. °.

Taxonomy: Pate, 1946. Notulae Nat. 171: 2 (synonymy).
rufifemur rufifemur (Packard). Transit. and U. Austr. Zones east of Rockies. Another subsp.

occurs in Mexico.
Crabro rufifemur Packard, 1866. Ent. Soc. Phila., Proc. 6: 81. 9, d.

Genus ECTEMNIUS Subgenus CLYTOCHRYSUS Morawitz
Crabro subg. Clytochrysus Morawitz, 1864. Acad. Imp. Sci. St. Petersburg, Bul. 7: 453.
Type-species: Crabro chrysostomus Lepeletier and Brulle. Desig. by Richards, 1935.
Several species of this subgenus are known to nest in decaying wood of logs or stumps. The
burrows may have a number of branches each ending in one or two cells.

lapidarius (Panzer). U. S. and Canada, transcont. in Canad., Transit. and U. Austr. Zones;
Palaearctic also. Ecology: Nests in rotting stumps or logs, stores 2-16 prey per cell,
makes 1-2 cells at end of each branch of the burrow and as many as 16 cells per nest.

Superfamily SPHECOIDEA 1677

Prey: Syritta pipiens (L.), Chrysogaster sp., Toxomerus occidentalis Curr.,
Sphaerophoria contigua Macq., Mesograpta marginata (Say), Parapenium sp., Paragus
tibialis (Fall.); Syrphidae seem to be the preferred prey, but occasionally Anthomyiidae
are used in Europe.

Crabro lapidarius Panzer, 1804. Faunae Ins. German., heft 90, pl. 12. d.

Crabro sinuatus Fabricius, 1804. Systema Piezatorum, p. 310.

Crabro cinctus Spinola, 1806. Insectorum Liguriae, v. 1, p. 104. 2, ¢. This is a questionable
synonym.

Crabro chrysostomus Lepeletier and Brulle, 1834. Soe. Ent. France, Ann. 3: 704. ?. Preoce.

Crabro comptus Lepeletier and Brulle, 1834. Soe. Ent. France, Ann. 3: 705. 3.

Crabro xylurgus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 139. 2, 3.

Crabro interstinctus Smith, 1851. Zoologist 9: exxvi. d.

Crabro obscurus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 418. 2, ¢.

Crabro gracilissimus Packard, 1866. Ent. Soe. Phila., Proc. 6: 78. d.

Crabro denticulatus Packard, 1866. Ent. Soc. Phila., Proc. 6: 97. 3.

Crabro effossus Packard, 1866. Ent. Soe. Phila., Proc. 6: 104. 6.

Crabro papagorum Viereck, 1908. Amer. Ent. Soe., Trans. 33: 401. 9.

Taxonomy: Pate, 1946. Notulae Nat. 171: 2-3 (synonymy).

Biology: Michener, 1971. Kans. Ent. Soc., Jour. 44: 405-407, 1 fig. (nest, prey).

ruficornis ruficornis (Zetterstedt). Canada and U. S., transcont. chiefly in Transit. Zone;

Mexico (Guerrero); also Palaearctic. Prey: Syrphus ribesii (L.) in N. Amer. Another
subsp. occurs in Taiwan.

Crabro ruficornis Zetterstedt, 1838. Ins. Lappon. 1: 4438. d.

Crabro aurilabris Herrich-Schaeffer, 1841. Faunae Ins. German., h. 179, p. 12. 6.

Crabro nigrifrons Cresson, 1865. Ent. Soc. Phila., Proc. 4: 482. d.

Crabro contiguus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 484. °.

Crabro septentrionalis Packard, 1866. Ent. Soc. Phila., Proc. 6: 110. 9.

Crabro (Solenius) planifrons Thomson, 1870. Opuse. Ent., v. 2, p. 173. 2, d.

Crabro hector Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 147, pl. 9, fig. 8. @.

Crabro (Clytochrysus) longipalpis Verhoeff, 1892. Ent. Nachr. 18: 70. 9. ‘

Crabro vestor Ashmead, 1899. Canad. Ent. 31: 173. Lapsus for hector Cam.

Crabro chipsanii Matsumura, 1911. Hokkaido Imp. Univ., Faculty Agr., Jour. 4: 102. °.

Crabro lineatotarsis Matsumura, 1911. Hokkaido Imp. Univ., Faculty Agr., Jour. 4: 103. @.

Taxonomy: Pate, 1946. Notulae Nat. 171: 2 (synonymy).

Biology: Krombein, 1936. Ent. News 47: 95 (prey).

yosemite Pate. Calif. (Mariposa Co.).
Ectemnius (Clytochrysus) yosemite Pate, 1946. Notulae Nat. 171: 3. 9.

Genus ECTEMNIUS Subgenus METACRABRO Ashmead

Metacrabro Ashmead, 1899. Canad. Ent. 31: 169.
Type-species: Crabro Kollari Dahlbom. Orig. desig.
Solenius subg. Lophocrabro Rohwer, 1917 (1916). Conn. State Geol. and Nat. Hist. Survey,
Bul. 22: 667.
Type-species: Crabro singularis Smith. Orig. desig.

In Europe cephalotes (Oliv.) has been reported to nest in both sound lumber and in decaying
trunks.

cephalotes (Olivier). Ont., Que., Conn., N. Y., N. J., Pa., Md., Ill.; Europe. Ecology: In Europe

nests both in sound lumber and in decaying logs; several females use a common
entrance but presumably have their own cells. Probably adventive in N. Amer. Prey: In
Europe stores many species of muscoid Diptera, occasionally Tabanidae and Syrphidae,
and one specimen of Nematinae was found in a cell.

Crabro tibialis Olivier, 1792. Encycl. Meth. Ins., v. 6, p. 513.

Crabro cephalotes Olivier, 1792. Encycl. Meth. Ins., v. 6, p. 513.

Crabro floralis Olivier, 1792. Encycl. Meth. Ins., v. 6, p. 517.

Crabro geniculatus Olivier, 1792. Encycl. Meth. Ins., v. 6, p. 517.

1678 Hymenoptera in America North of Mexico

Crabro cephalotes Panzer, 1799. Faunae Ins. German., h. 6, pl. 62, fig. 16. Preoce. This is a
questionable synonym.

Crabro striatus Lepeletier and Brulle, 1834. Soe. Ent. France, Ann. 3: 707. 2, d.

Crabro ornatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 709. °.

Blepharipus striatulus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 737. 6.

Ceratocolus striatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 744. 3. Preoce.

Crabro Lindenius Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 143. 2, ¢.

Crabro Shuckardi Dahlbom, 1838. Exam. Crabron. Scand., p. 98. 2, 6.

Crabro interruptus Dahlbom, 1845. Hym. Europaea, v. 1, p. 418. 2, ¢. N. name for
Shuckardi.

Crabro Fargeii Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 410. N. name for Ceratocolus
striatus.

Crabro Lindensis Inchbald, 1859. Ent. Weekly Intelligencer 6: 199. Lapsus.

Crabro aciculatus Provancher, 1882. Nat. Canad. 13: 108. 9, 6.

Crabro (Solenius) ruthenicus Morawitz, 1892. Soc. Ent. Rossica, Horae 26: 174. 2.

Crabro Lindenii Dalla Torre, 1897. Cat. Hym., v. 8, p. 621. Lapsus for lindensis Inchb.

Taxonomy: Pate, 1945. Brooklyn Ent. Soce., Bul. 40: 41-43 (synonymy). —van der Vecht, 1961.
Zool. Verhandl. Rijksmus. Natuurlijke Hist. Leiden, No. 48, pp. 70-71 (identity of
cephalotes Oliv.).

Biology: Hamm and Richards, 1926. Ent. Soc. London, Trans., pp. 306-307, 321-322 (nest, prey
in Europe; misdet. as quadricinctus F.).
maculosus (Gmelin). Canada and U. S. in Transit. Zone east of 100th meridian. Parasite:

Crabrovidia ectemnii Fain. Prey: Tubifera arbustorwm (L.).

Crabro maculatus Fabricius, 1781. Spec. Ins., v. 1, p. 470. Secondary homonym in Vespa.

Vespa (Crabro) maculosa Gmelin, 1790. In Linnaeus, Syst. Nat., Ed. 13, v. 1, p. 2761. N.
name.

Crabro singularis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 417. d.

Crabro frigidus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 419. 2.

Crabro quadrangularis Packard, 1866. Ent. Soc. Phila., Proce. 6: 85. 2.

Crabro 14-maculatus Packard, 1866. Ent. Soe. Phila., Proc. 6: 87. 6.

Crabro oblongus Packard, 1866. Ent. Soc. Phila., Proc. 6: 88. 9.

Crabro trapezoideus Packard, 1866. Ent. Soc. Phila., Proc. 6: 89. d.

Crabro quadrangulus Cresson, 1928. Amer. Ent. Soc., Mem. 5: 55. Lapsus.

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1025 (prey).
Genus ECTEMNIUS Subgenus HYPOCRABRO Ashmead

Hypocrabro Ashmead, 1899. Canad. Ent. 31: 168.
Type-species: Crabro 10-maculatus Say. Orig. desig.
Pseudocrabro Ashmead, 1899. Canad. Ent. 31: 169.
Type-species: Crabro chrysarginus of Lepeletier. Orig. desig.
Xestocrabro Ashmead, 1899. Canad. Ent. 31: 169.
Type-species: Crabro 6-maculatus Say. Orig. desig.
Xylocrabro Ashmead, 1899. Canad. Ent. 31: 169.
Type-species: Crabro stirpicola Packard. Orig. desig.
Ectemnius subg. Apoctemnius Leclercq, 1950. Rev. Franc. d’Ent. 17: 200.
Type-species: Ectemnius (Apoctemnius) domingensis Leclercq. Orig. desig.

Some of our species nest in the soft pith of living or dead stems or twigs and make a linear se-
ries of cells separated by partitions of pith particles. Other species nest in decaying trunks,
stumps or limbs.

alpheus Pate. Calif., Wash., Utah.
Ectemnius (Hypocrabro) alpheus Pate, 1946. Notulae Nat. 171: 5. 2, do.
arcuatus (Say). Transcont. in Transit. and Austr. Zones. Ecology: Nests in logs. Prey: Musca
domestica L.
Crabro arcwatus Say, 1837. Boston Jour. Nat. Hist. 1: 377. 9.
Crabro Packardii Cresson, 1865. Ent. Soe. Phila., Proe. 4: 477. 2, d.
Crabro honestus Cresson, 1865. Ent. Soe. Phila., Proce. 4: 485. “2” = 6.

Superfamily SPHECOIDEA 1679

Crabro villosifrons Packard, 1866. Ent. Soc. Phila., Proc. 6: 84. °.
Crabro (subgenus?) nokomis Rohwer, 1908. Ent. News 19: 251. 9.
Solenius (Hypocrabro) nokonis (!) Rohwer, 1917. U. S. Natl. Mus., Proe. 53: 242.

Taxonomy: Pate, 1946. Notulae Nat. 171: 9-10 (synonymy).

Biology: Rau and Rau, 1918. Wasp studies afield, pp. 94, 96 (nest, prey).
besseyae (Rohwer). Calif., Colo., Utah, N. Mex.
Crabro (Xylocrabro?) besseyae Rohwer, 1908. Ent. News 19: 419. 2.
centralis (Cameron). Tex., N. Mex., Ariz.; Mexico south to Colombia and Trinidad.
Crabro centralis Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 146. 2.

continuus continuus (Fabricius). Canada and U. S., transcont. in Canad., Transit. and U.
Austr. Zones; Palaearctic Region. Ecology: Nests in borings in tree root or rotten
branch. Parasite: Macronychia aurata (Coq.). Prey: Archytas aterrimus (Desv.),
Winthemia sp., Tachinidae sp.; Sarcophaga sp., Sarcophagidae sp.; Pollenia rudis (F.),
Phaenicia ? sp., Calliphoridae sp.; Muscidae sp.; in Europe muscoid Diptera are also the
preferred prey but Syrphidae are used occasionally. Another subsp. occurs in the
Canary Islands.

Crabro continuus Fabricius, 1804. Systema Piezatorum, p. 312.

Crabro 6-maculatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 341. 9.
Preoce.

Solenius punctatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 720. d.

Ceratocolus punctatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 749. 2, 6.
Preoce.

Crabro fuscitarsis Herrich-Schaeffer, 1841. Faunae Ins. German., h. 181, pl. 7. @.

Crabro impressus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 401. N. name for Ceratocolus
punctatus Lep. and Br.

Crabro sulphureipes Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 415. d.

Crabro (Ectemnius) fuscitarsus Schenck, 1857. Nassau. Ver. f. Naturk., Jahrb. 12: 70.
Lapsus.

Crabro vagatus Smith, 1869. Entomologist 4: 208. 2.

Crabro granulatus Walker, 1871. List Hym. Egypt, p. 26. ¢.

Crabro rugoso-punctatus Taschenberg, 1875. Ztschr. f. Naturw. 45: 385. d.

Crabro validus De Stefani, 1884. Nat. Sicil., v. 3, p. 218. ¢.

Crabro vagans Fokker, 1887. Tijdschr. v. Ent. 30: xx. Lapsus.

Xylocrabro-slossonae Ashmead, 1902. Ent. News 13: 5. ¢6. Nom. nud.

Crabro sayi Cockerell, 1910. Entomologist 43: 61. N. name for Crabro sexmaculatus Say.

Crabro bisexrmaculatus Viereck, 1910 (1909). In Smith, N. J. State Mus. Ann. Rpt. 1909, p.
681. N. name for Crabro sexmaculatus Say.

Crabro hispanicus Kohl, 1915. K. K. Naturhist. Hofmus. Wien, Ann. 29: 81. °.

Solenius (Hypocrabro) giffardi Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 242. 2, d.

Taxonomy: Pate, 1946. Notulae Nat. 171: 10 (synonymy). —Evans, 1964. Amer. Ent. Soc.,
Trans. 90: 295, fig. 114 (larva).

Biology: Peckham and Peckham, 1905. Wasps Social and Solitary, pp. 97-101, 1 fig. (nest,
prey). —Reinhard, 1929. The Witchery of Wasps, pp. 228-231, 4 figs. (nest, prey, parasite).
—Krombein, 1961. Brooklyn Ent. Soc., Bul. 56: 65 (prey). —Krombein, 1964. Brooklyn Ent.
Soe., Bul. 58: 120 (nest, prey, cocoon, life cycle). There is also an extensive European
literature.

decemmaculatus decemmaculatus (Say). U. and L. Austr. Zones east of Rocky Mts.; Mexico.
Crabro 10-maculatus Say, 1823. West. Quart. Rptr., v. 2, p. 78.
Crabro chrysargyrus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 711. @.
Crabro chrysarginus Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 114. Lapsus.
Crabro chrysargurus Dahlbom, 1845. Hym. Europaea, v. 1, p. 386. Lapsus.
Crabro collinus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 420. ¢.
Crabro aurifrons Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 420. 2.
Crabro (Hoplocrabro) novanus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 564. 2, d.

Taxonomy: Pate, 1946. Notulae Nat. 171: 8 (synonymy).

1680 Hymenoptera in America North of Mexico

decemmaculatus tequesta Pate. Cent. and south. Fla.
Ectemnius (Hypocrabro) 10-maculatus tequesta Pate, 1946. Notulae Nat. 171: 9. 2, 6.
excavatus banksi (Rohwer). Md. to Ga., Ind., Kans., Mo., Tex. south to Mexico (Yucatan).
Ecology: Nests in rotten log.
Crabro banksi Rohwer, 1909. Ent. News 20: 147. °.
Ectemnius (Hypocrabro) ravinus Leclereq, 1968. Soc. Ent. France, Ann. (n. s.) 4 (2): 325.
OF

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1026 (nest).

excavatus excavatus (Fox). Fla.
Crabro excavatus Fox, 1892. Ent. News 3: 10. 2, d (in part misdet.).

odyneroides (Cresson). Colo., N. Mex., Ariz.; Mexico (Guerrero, Durango).
Crabro odyneroides Cresson, 1865. Ent. Soc. Phila., Proc. 4: 481. d.
Crabro ariel Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 147, pl. 9, fig. 7, a, b. 2, d.
paucimaculatus (Packard). N. Y. to Fla., Ill., Tex. Ecology: Nests in green Hibiscus stems
along river bank, chewing through side of stem to reach central pith cavity, makes
linear series of 1-18 cells, stores 7-31 prey per cell. Parasite: Vidia cooremani Baker;
Megaselia aletiae (Comst.); Eustalomyia vittipes (Zett.); Macronychia aurata (Coq.),
Ptychoneura aristalis (Coq.); Tetrabaeus americanus (Brues); Perilampus canadensis
Cwfd.; Diomorus zabriskiei Cr. Prey: Dolichopus ovatus Lw., D. sp.; Melanagromyza
diantherae (Mall.), Agromyzidae sp.; Notiphila carinata Lw., N. erythrocera Lw., N. sp.,
Hydrellia spp., Scatella picea (Wlkr.); Eumetopiella rufipes (Macq.), Chaetopsis sp.,
Otitidae sp.; Camptoprosopella angulata Shew.; Lonchaea polita Say; Sepedon armipes
Lw., S. sp., Dictya texensis Curr.; Leptocera richardsi Sabr.; Coenosia atrata Wlkr.,
Lispe albitarsis Stein; preferred prey are Agromyzidae and Ephydridae. The common
name is the hibiscus wasp.
Crabro paucimaculatus Packard, 1866. Ent. Soc. Phila., Proc. 6: 90. °.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 294-295, fig. 113 (larva).

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 1026 (nest). —Krombein, 1964. Biol.
Soe. Wash., Proc. 77: 73-88, figs. 1-11 (nest, prey, egg, cocoon, life cycle, parasites).
rufipes ais Pate. Fla. Ecology: Nests in rose canes, makes linear series of up to 3 cells per
nest, stores 20-30 prey per cell. Prey: Euxesta nitidiventris Lw.; Aciura insecta Lw.,
Paroxyna sorocula (Wied.); Chloropidae spp.; Otitidae and Trypetidae are the preferred
prey.
Ectemnius (Hypocrabro) texanus ais Pate, 1946. Notulae Nat. 171: 12. 2, d.
Biology: Morse and Kerr, 1957. Fla. Ent. 40: 77-80, 1 fig. (nest, prey).
rufipes rufipes (Lepeletier and Brulle). N. Y. to north. Fla., west to Iowa, Kans. and Tex.
Ecology: Nests in twigs.
Ceratocolus rufipes Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 741. 2.
Crabro texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 227. 9.
satan Pate. N. Mex. to Calif.
Ectemnius (Hypocrabro) satan Pate, 1946. Notulae Nat. 171: 10. d.
scaber rufescens Krombein. Cent. and south Fla.
Ectemnius (Hypocrabro) scaber rufescens Krombein, 1954. Amer. Ent. Soc., Trans. 80: 24.
Ones
scaber scaber (Lepeletier and Brulle). N. J. to Fla., La., Tex., Okla. Ecology: Nests in pine.
Solenius scaber Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 715. d.
sonorensis (Cameron). Colo., N. Mex., Ariz., south. Calif.; Mexico (Baja California, Sonora,
Guerrero).
Crabro sonorensis Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 144, pl. 9, fig. 4. °.
Crabro montivagus Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 145, pl. 9, figs. 5, a, b.
OErOr
Crabro imbutus Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 108. °.
Crabro (Solenius ?) ferrugineipes Rohwer, 1908. Ent. News 19: 250. 6.

Superfamily SPHECOIDEA 1681

spiniferus (Fox). Colo., Nev., Ariz., Calif. to Wash. Ecology: Nests in stems of Sambucus,

Foeniculum and Eriogonum, makes linear series of 1-6 cells per nest, stores 2-10 prey
per cell. Parasite: Senotainia trilineata (Wulp), Macronychia sp., Amobia floridensis
(Tns.); Megaselia sp.; Diomorus zabriskiei Cr.; Monodontomerus sp. Prey:
Sphaerophoria sp.; Ogcodes eugonatus Lw.; Hylemya sp.; preferred prey are
Acroceridae. Predator: Cymatodera ovipennis LeC.

Crabro spiniferus Fox, 1895. Amer. Ent. Soc., Trans. 22: 148. 9, 6.

Solenius (Pseudocrabro) conspiciendus Mickel, 1918 (1917). Nebr. Univ. Studies 17: 324. 9,
é.

Biology: Bechtel and Schlinger, 1957. Ent. News 68: 225-232 (nest, prey, parasites, predator).
—Parker and Bohart, 1966. Pan-Pacific Ent. 42: 94 (nest, parasites).
stirpicola (Packard). Canada and U.S. in Transit. and U. Austr. Zones east of 100th meridian.
Ecology: Nests in pith in twigs of Ailanthus, blackberry, sumac, Sambucus, and in old
railroad tie, stores 6-27 prey per cell. Parasite: Diomorus zabriskiei Cr.; Habritys latro
Wall.; Perilampus canadensis Cwfd.; Amobia sp. Prey: Simulium jenningsi Mall.; Solva
pallipes (Lw.); Oxycera maculata Oliv.; Condylostylus sipho (Say), C. sp., Asyndetus
sp.; Mesograpta marginata (Say); Euxesta notata (Wied.); Amphicnephes pullus
(Wied.), Rivellia pallida Lw., R. steyskali Namba; Euaresta bella (Lw.); Lonchaea
polita Say ?, L. sp.; Lyciella pictiventris (Mall.), Sapromyza umbrosa Lw.; Agromyza
parvicornis Lw., A. sp., Melanagromyza burgessi (Mall.), M. virens (Lw.), M. sp.;
Chyromya sp.; Cordilura fuscipes Zett., Phorbia sp., Anthomyia sp.; Coenosia sp. near
antennalis Stein; Phormia regina (Meig.), Lucilia sp., Calliphora vomitoria (L.);
Ravinia derelicta (Wlkr.), Sarcodexia sp.; Hyalomyodes triangulifer (Lw.),
Cryptomeigenia eumyothyroides (Tns.), Paradidyma singularis (Tns.), Sitophaga
calosomoides (Tns.), Lespesia sp., Elfia johnsoni (Coq.), Tachinidae sp.; although
stirpicola preys upon a wide range of Diptera, the preferred prey are clearly
acalyptrate and calyptrate Cyclorrhapha.
Crabro stirpicola Packard, 1866. Ent. Soc. Phila., Proc. 6: 111. 2, ¢.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 109, figs. 85-90 (larva). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 165 (larva).

Biology: Packard, 1869. Guide Study Ins., p. 158 (nest). —Cresson, 1878. Psyche 2: 189 (nest,
parasite). —Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 46-52, pl. 1,
fig. 5, pl. 11, figs. 6, 7 (nest, prey, cocoon, life cycle). —Rau and Rau, 1918. Wasp Studies
Afield, pp. 90-94, figs. 21, 22 (nest, prey, life cycle). —Rau, 1922. Acad. Sci. St. Louis,
Trans. 24: 20-21 (nest, prey, life cycle, parasite). —Rau, 1928. Acad. Sci. St. Louis, Trans.
25: 381-387, fig. 50 (nest, prey, life cycle). —Krombein, 1960. Ent. News 71: 63-68 (nest,
prey, egg, life cycle, cocoon, parasites).

trifasciatus (Say). Chiefly Transit. Zone of Canada and U.S. east of Cascade and Sierra
Nevada ranges.
Crabro trifasciatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 342.

Genus ECTEMNIUS Subgenus ECTEMNIUS Dahlbom

Crabro subg. Ectemnius Dahlbom, 1845. Hym. Europaea, v. 1, p. 389.
Type-species: Crabro guttatus of Dahlbom. Desig. by Ashmead, 1899.
Crabro subg. Mesocrabro Verhoeff, 1892. Ent. Nachr. 18: 70.
Type-species: Crabro guttatus Vander Linden. Desig. by Pate, 1937.
atriceps (Cresson). South. Canada and U.S., chiefly in Transit. Zone. Ecology: Nests in logs.
Crabro atriceps Cresson, 1865. Ent. Soc. Phila., Proc. 4: 483. 9.
Crabro brunneipes Packard, 1866. Ent. Soc. Phila., Proc. 6: 102. d.
Crabro foxti Kineaid, 1900. Ent. News 11: 356. d.

Taxonomy: Pate, 1946. Notulae Nat. 171: 14 (synonymy). —Evans, 1957. Amer. Ent. Soc.,
Trans. 83: 109, figs. 91-94 (larva).
corrugatus (Packard). Canad. and Transit. Zones in Alaska, Canada, U. S.
Crabro pauper Packard, 1866. Ent. Soc. Phila., Proc. 6: 95. ¢.
Crabro corrugatus Packard, 1866. Ent. Soc. Phila., Proc. 6: 107. °.
Crabro (Cuphopterus) operus Rohwer, 1908. Ent. News 19: 247. °.

1682 Hymenoptera in America North of Mexico

Crabro (Xestocrabro) drymocallidis Rohwer, 1908. Ent. News 19: 255. 6.

Taxonomy: Pate, 1946. Notulae Nat. 171: 14 (synonymy).
dives (Lepeletier and Brulle). Transcont. in Canad. and Transit. Zones of Canada and U. S.;

also Palaearctic. Ecology: Nests in log in U. S., in Europe in decayed or rotten logs, and
soft pith of canes and stems. Prey: Syrphidae and Tachinidae in Europe. Predator:
Philanthus pulcher D. T.

Solenius dives Lepeletier and Brulle, 1834. Soe. Ent. France, Ann. 3: 716. 2, d.

Solenius octonotatus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 719. d.

Solenius alatulus Dahlbom, 1838. Exam. Crabron. Seand., p. 85. 2, 6.

Crabro pictipes Herrich-Schaeffer, 1841. Faunae Ins. German., h. 181: pl. 5.

Solenius octavonotatus Lepeletier, 1845. Hist. Nat. Ins. Hym., p. 125. Emend.

Crabro auratus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 398. N. name for dives.

Crabro montanus Cresson, 1865. Ent. Soe. Phila., Proc. 4: 484. ?. Preoce.

Crabro cristatus Packard, 1866. Ent. Soe. Phila., Proce. 6: 101. 6.

Crabro cubiceps Packard, 1866. Ent. Soc. Phila., Proc. 6: 105. 2.

Crabro (Xestocrabro) heraclei Rohwer, 1908. Ent. News 19: 253. 6.

Crabro montivagans Strand, 1917. Archiv f. Naturgesch. 82: 98. N. name for montanus.

Taxonomy: Pate, 1945. Brooklyn Ent. Soc., Bul. 40: 38-40 (synonymy). —Pate, 1946. Notulae
Nat. 171: 14 (synonymy).

Biology: Barth, 1907. Wis. Nat. Hist. Soc., Bul. 5: 251 (nest). —Richards, 1944. Roy. Ent. Soc.
London, Proc., ser. A, 9: 1384 (nest, prey in Europe). —Pate, 1945. Brooklyn Ent. Soc., Bul.
40: 40 (refs. to European lit.).

proletarius (Mickel). Colo., N. Dak.
Crabro parvulus Packard, 1866. Ent. Soc. Phila., Proc. 6: 108. 2. Preoce.
Crabro (Xestocrabro) proletarius Mickel, 1916. Amer. Ent. Soc., Trans. 42: 426. ¢.

Genus LESTICA Billberg
So far as known all species in this genus prey upon adult Lepidoptera, usually Microlepidop-
tera or small Noctuidae, but butterflies may be used occasionally.
Taxonomy: Pate, 1947. Notulae Nat. 185: 13 (key to subg.).

Genus LESTICA Subgenus LESTICA Billberg

Lestica Billberg, 1820. Enum. Ins., p. 107.
Type-species: Crabro subterraneus Fabricius. Desig. by Rohwer, 1911.
Hypothyreus Ashmead, 1899. Canad. Ent. 31: 171.
Type-species: Crabro subterraneus Fabricius. Orig. desig.
The typical subgenus does not occur in North America. Members of this subgenus are
ground-nesting species.

Genus LESTICA Subgenus SOLENIUS Lepeletier and Brulle

Solenius Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 713.
Type-species: Solenius interruptus Lepeletier and Brulle. Desig. by Internatl.
Comn. Zool. Nomencl., Op. 1015, 1974.

Members of this subgenus nest in decaying wood or in soft pith of plant stems.

Taxonomy: Court and Menke, 1968. Bul. Zool. Nomencl. 24: 357-358 (petition to set aside
type-species designation of Sphex vagus Westwood, 1839, and to designate Solenius
interruptus Lep. and Br. as type-species).

cinctella (Fox). Nev., Calif., Oreg.
Crabro cinctellus Fox, 1895. Amer. Ent. Soc., Trans. 22: 136. °.

Taxonomy: Leclercq, 1951, Soc. Ent. Belg., Bul. et Ann. 87: 171. 3.

Superfamily SPHECOIDEA 1683

confluenta (Say). South. Canada and U.S. in Transit. and Austr. Zones; Mexico (Chihuahua).

Ecology: Nests in decaying logs, in borings in wooden posts and in pith of catalpa stem,
stores 6 prey per cell. Prey: Adults of moth spp.

Solenius interruptus Lepeletier and Brulle, 1834. Soc. Ent. France, Ann. 3: 716. 2. Preoce.
in Crabro.

Crabro confluentus Say, 1837. Boston Jour. Nat. Hist. 1: 376. °, d.

Crabro dubius Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 417. N. name.

Crabro confluens Leconte, 1859. Ent. of N. Amer. (Thos. ae p. 758. Lapsus.

Crabro bellus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 481.

Crabro atrifrons Cresson, 1865. Ent. Soe. Phila., Proce. 4: is ee

Crabro eburneus Taschenberg, 1875. Ztschr. Gesam. Naturw. Halle 45: 383. é.

Crabro cinctibellus Viereck, 1908. Amer. Ent. Soc., Trans. 33: 401. 2, do.

Crabro opwana Rohwer, 1908. Ent. News 19: 248. ¢.

Crabro (Solenius) townsendi Rohwer, 1911. U.S. Natl. Mus., Proe. 40: 563. 9.

Crabro (Solenius) planaris Mickel, 1916. Amer. Ent. Soc., Trans. 42: 427. d.

Solenius seamansi Carter, 1925. Canad. Ent. 57: 135. ¢.

Taxonomy: Pate, 1947. Notulae Nat. 185: 13-14 (synonymy).

Biology: Peckham and Peckham, 1905. Wasps, Social and Solitary, pp. 102-105, 1 fig. (nest,
prey). —Rau, 1922. St. Louis Acad. Sci., Trans. 24: 19-20 (nest). —Rau, 1928. Acad. Sci. St.
Louis, Trans. 25: 378 (nest, cocoon, life cycle).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 20, figs. O, R (male
genitalia).

producticollis (Packard). Transcont. in Canada and U.S. in Transit. and U. Austr. Zones.
Crabro producticollis Packard, 1866. Ent. Soc. Phila., Proc. 6: 76. d.

Crabro 4-maculatus Provancher, 1882. Nat. Canad. 12: 102. 2. Typ. err. ? Preoce.
Crabro 4-punctatus Provancher, 1883. Faune Ent. Canad., Hym., p. 653. 3d. Preoce.

NomInA NupDA IN CRABRONINAE
glauconotatus Harris. Mass.

Crabro glauconotatus Harris, 1835. In Hitchcock, Rpt. Geol., Mineral., Bot., Zool., Mass., p.
588.

Family MELLINIDAE

This small family contains only two subfamilies, Xenosphecinae and Mellininae, which appear
to be more closely related to one another than either is to any other family of Sphecoidea. The
former subfamily is known only from the southwestern deserts of North America. The latter is
Holaretic and Neotropical in distribution. Both subfamilies nest in the ground and prey upon
Diptera.

SUBFAMILY XENOSPHECINAE
Genus XENOSPHEX Williams

Xenosphex Williams, 1954. Wasmann Jour. Biol. 12: 97.
Type-species: Xenosphex xerophila Williams. Orig. desig.
Revision: Parker, 1966. Pan-Pacific Ent. 42: 190-195, 8 figs.

boharti Parker. Calif. (Inyo Co.).
Xenosphex boharti Parker, 1966. Pan-Pacific Ent. 42: 191, figs. 1, 2, 6, 7. d, 2.

timberlakei Williams. Southeast. Calif., northwest. Ariz., south. Nev. Ecology: Nests in sand.

Prey: Lordotus miscellus Coq.
Xenosphex timberlakei Williams, 1955. Wasmann Jour. Biol. 13: 313. °.

Biology: Parker, 1966. Pan-Pacifie Ent. 42: 194-195 (nest, prey transport).

xerophilus Williams. South. Calif. and Nev., northwest. and south. Ariz.
Xenosphex xerophila Williams, 1954. Wasmann Jour. Biol. 12: 99, figs. 1-6. d (2 misdet.).

Taxonomy: Williams, 1955. Wasmann Jour. Biol. 13: 313. 9.

1684 Hymenoptera in America North of Mexico

SUBFAMILY MELLININAE

Taxonomy: Maidl and Klima, 1939. Hym. Cat., Pars 8, Sphecidae, v. 1, pp. 30-43 (world
catalog). —Evans, 1959. Amer. Ent. Soe., Trans. 85: 145-147, figs. 23-30 (larva).

Genus MELLINUS Fabricius

Mellinus Fabricius, 1790. Skrivt. Naturhist. Selsk., h. 1, p. 226.
Type-species: Vespa arvensis Linnaeus. Desig. by Curtis, 1836.
Millimus Gimmerthal, 1836. Soc. Imp. Nat. Moscow, Bul. 9: 449. Lapsus or emend.

Nothing is known of the biology of the North American species. The European arvensis (L.)
nests gregariously in the ground, makes multicelled nests with each cell on a branch off the main
burrow, preys primarily on muscoid Diptera and stores 4-9 flies per cell.

Revision: Fox, 1894. Ent. News 5: 201-203 (N. Amer. spp.).

Taxonomy: Siri and Bohart, 1974. Pan-Pacifie Ent. 50: 169-176, 16 figs. (synopsis of world
spp.).
abdominalis Cresson. Nebr., Colo., Wyo., Mont.
Mellinus abdominalis Cresson, 1882 (1881). Amer. Ent. Soc., Trans. 9: Proc., p. xxxix. @,
3.
Mellinus abdominalis var. personatus Fox, 1894. Ent. News 5: 202. °.
bimaculatus Packard. Maine, N. H., Mass., N. Y., Pa., N. J., Ohio, Mich.; Mexico (Jalisco,
Oaxaca).
Mellinus bimaculatus Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p.
68. Nom. nud.
Mellinus bimaculatus Packard, 1867. Ent. Soc. Phila., Proc. 6: 419. 2.
Mellinus wolcotti Smith, 1908. Ent. News 19: 299. °.
imperialis Bohart. Calif. (Imperial Co.); Mexico (Sonora).
Mellinus imperialis Bohart, 1968. Pan-Pacific Ent. 44: 235, fig. 14. d, ?.
rufinodus Cresson. Mont., Colo., S. Dak., Tex., N. Mex., Ariz., Utah; Mexico (Chihuahua,
Durango, Guerrero, Mexico, Aguascalientes).
Mellinus rufinodus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 475. 2, d.

Family NYSSONIDAE

All members of this large cosmopolitan family nest in the ground. They are commonly called
sand wasps.

Taxonomy: Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 404 (key to tribes). —Pate, 1938. Amer.
Ent. Soc., Trans. 64: 119-120 (key to tribes). —Evans and Lin, 1956. Amer. Ent. Soc.,
Trans. 82: 35-66, 13 pls. (larvae). —Evans, 1959. Amer. Ent. Soc., Trans. 85: 149-156, figs.
36-42, 60, 62, 63, 71-73 (larvae). —Evans, 1964. Amer. Ent. Soc., Trans. 90: 253-273, figs.
26-58 (larvae).

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, 526 pp., 215 figs., 47
tabs.

SUBFAMILY ALYSSONINAE

Revision: Handlirsch, 1895. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 104: 830-839 (world
spp.).

Genus ALYSSON Panzer

Alysson Jurine, 1801. Intell. Blatt. Litt.-Ztg. Erlangen, v. 1, p. 164. Name suppressed by
Internat]. Comn. Zool. Nomencl., Op. 135, 1939.
Alysson Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 169.
Type-species: Pompilus spinosus Panzer. Desig. by Morice and Durrant, 1915.
Alyson Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 195.
Type-species: Pompilus spinosus Panzer. Monotypic.

Superfamily SPHECOIDEA 1685

These wasps are primarily Holarctic in distribution, but a few species occur in the Ethiopian
and Oriental Regions. They nest gregariously, often in cool, moist sand, usually make multicellu-
lar nests, and prey primarily on nymphal and adult leafhoppers although spittle bugs and
planthoppers are used occasionally.

Revision: Fox, 1894. Ent. News 5: 86-89 (N. Amer. spp.).
conicus Provancher. N. B., Ont., N. H., N. Y., Md., D. C., Va., Mich.
Alyson(!) conicus Provancher, 1889. Addit. Corr. Faune Ent. Canada Hym., p. 271. 2.
flavomaculatus Cameron. N. Mex. (Santa Fe Mts.).
Allyson(!) flavomaculatus Cameron, 1901. Amer. Ent. Soc., Trans. 27: 314. 6.
guignardi Provancher. Transcont., Que. and Ont. to N. C. and Mo., west to Mich., Iowa, Calif.
and N. Mex. Predator: Dioctria bawmhaueri Meig.
Alyson(!) Guignardi Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 271. 2,
3

Alyson(!) petiolatus Cameron, 1902. Amer. Ent. Soc., Trans. 28: 374. ¢.
Alyson(!) interstitialis Cameron, 1902. Amer. Ent. Soc., Trans. 28: 375. d.
melleus Say. U. and L. Austr. Zones east of 100th meridian. Ecology: Nests in damp sand or

sandy loam, makes 1-5 cells per nest, stores 3-23 prey per cell. Parasite: Phrosinella
fulvicornis (Coq.) ? Prey: Aceratagallia sp., Agallia constricta Van D., Agalliopsis
novella (Say), Balclutha sp., Chlorotettix sp., Ciminius hartii Ball, Colladonus
clitellarius (Say), Deltocephalus flavicosta Stal, Draeculacephala antica (Wlkr.), D.
mollipes (Say), D. paludosa B. and C., D. portola Ball, D. spp., Empoasca fabae (Harr.),
Exitianus exitiosus Uhl., Graminella nigrifrons (Fbs.), G. pallidula Osb., G. sp.,
Hortensia similis (Wlkr.), Keonolla dolobrata (Ball), Macrosteles fascifrons (Stal),
Neokolla hieroglyphica (Say), Paraphlepsius irroratus (Say), Sanctanus sp.,
Scaphytopius sp., Tylozygus bifidus (Say), Cicadellinae spp.; Delphacodes basivitta (Van
D.), Delphacidae sp.; both nymphs and adults are stored; cicadellids are preferred prey
and delphacids are used rarely.

Alyson(!) melleus Say, 1837. Boston Jour. Nat. Hist. 1: 380. 2.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 41, figs. 1-7 (larva). —Evans,
1959. Amer. Ent. Soc., Trans. 85: 150, fig. 60 (larva).

Biology: Hartman, 1905. Tex. Acad. Sci., Trans. 7: 56-57 (nest, prey transport). —Rau and
Rau, 1918. Wasp studies afield, pp. 140-144, fig. 33 (nest, prey). —Evans, 1966. Compar.
ethology and evolution of sand wasps, pp. 17-28, figs. 9-20 (nest, prey, mating, egg, life
cycle, parasite ?). —Evans, 1968. Ent. Soc. Amer., Ann. 61: 1343 (prey). —Kurezewski and
Kurezewski, 1971. Kans. Ent. Soc., Jour. 44: 335 (prey).

oppositus Say. Transit. and U. Austr. Zones east of Rocky Mts., N. B. and Ont. to Ga. and
Tenn., west to Mich,, Iowa and Colo. Prey: Draeculacephala mollipes (Say).
Alyson(!) oppositus Say, 1837. Boston Jour. Nat. Hist. 1: 380. 2, d.
Alyson(!) oppositus var. a Say, 1837. Boston Jour. Nat. Hist. 1: 380.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, p. 28 (prey).
Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 20, figs. F-H (male
genitalia).

radiatus Fox. Colo., Nev., Calif., Wash.
Alyson(!) radiatus Fox, 1894. Ent. News 5: 87. 2, 6.

striatus Fox. D. C., N. Y.,
Alyson(!) striatus Fox, 1894. Ent. News 5: 88. 6.

triangulifer shawi Bradley. Nev., B. C., Calif.
Alysson shawi Bradley, 1920. Amer. Ent. Soc., Trans. 46: 130. ¢.

triangulifer triangulifer Provancher. Ont. and Que. south to Md., Mich. and Ky. west to N.
Dak. and Colo., Idaho, Alaska.
Alyson(!) triangulifer Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 272. 6.

Genus DIDINEIS Wesmael
Didineis Wesmael, 1852. Acad. Roy. Belg., Bul. 19: 109.

1686 Hymenoptera in America North of Mexico

Type-species: Alyson(!) lunicornis of Vander Linden. Monotypice.

The European lwnicornis (F.) makes a unicellular nest in the ground and preys upon nymphal
and adult Cicadellidae and Delphacidae.

Revision: Fox, 1894. Ent. News 5: 126-128 (N. Amer. spp.). —Malloch and Rohwer, 1930. We
S. Natl. Mus., Proc. 77 (14): 1-7, 8 figs. (N. Amer. spp.).
dilata Malloch and Rohwer. Wis., Nebr.
Didineis dilata Malloch and Rohwer, 1930. U. S. Natl. Mus., Proc. 77 (14): 6. 6.
latimana Malloch and Rohwer. N. H. to Va., Ill., Iowa, Mo.
Didineis latimana Malloch and Rohwer, 1930. U. S. Natl. Mus., Proc. 77 (14): 4. 2, 6.
nodosa Fox. Colo., Utah, Idaho, Wash. to Calif.; Mexico (Baja california).
Didineis nodosa Fox, 1894. Ent. News 5: 127. ¢.
Didineis sanctacrucae Bradley, 1920. Amer. Ent. Soc., Trans. 46: 129. 9.
Didineis nodosa var. clypeata Malloch and Rohwer, 1930. U. S. Natl. Mus., Proce. 77 (14): 6.
3.
peculiaris Fox. Iowa, Mont., Calif.; Mexico (Baja California).
Didineis peculiaris Fox, 1894. Ent. News 5: 128. 92, d.
stevensi Rohwer. N. Dak. (Sheldon).
Didineis stevensi Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 98. °.
texana (Cresson). Mostly U. and L. Austr. Zones, Pa. and N. J. to Fla., west to Kans., Mo. and
east. Ariz.; Mexico (Nuevo Leon). Prey: Cixius stigmatus Say.
Alyson(!) texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 226. d.

Biology: Strandtmann, 1945. Ent. Soe. Amer., Ann. 38: 312 (prey).
vierecki Rohwer. Kans., N. Mex.
Didineis crassicornis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 204. 3d. Preoce.
Didineis vierecki Rohwer, 1911. Ent. Soc. Wash., Proc. 13: 4. N. name.

SUBFAMILY NYSSONINAE

The wasps of this subfamily are cleptoparasites in the nests of other ground-nesting aculeates.
The more primitive genera have as their hosts wasps belonging to the closely allied Gorytinae.
Two of the more highly specialized genera appear to have as hosts Larrinae (Larridae) and Cer-
cerinae (Philanthidae), and a third may have an andrenid bee as its host.

Revision: Cresson, 1882. Amer. Ent. Soc., Trans. 9: 273-284. —Fox, 1896. N. Y. Ent. Soc.,
Jour. 4: 10-16. —Pate, 1938. Amer. Ent. Soc., Trans. 64: 121-127 (key to gen.).

Taxonomy: Maid] and Klima, 1939. Hym. Cat., Pars 8, Sphecidae i, pp. 115-150 (world
catalog).

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 83-90, figs. 46-51
(compar. behavior and host relationships).

Genus NYSSON Latreille

Nysso Latreille, 1796. Precis Caract. Gen. Ins., p. 125. No species included. Printer’s error
for Nysson?
Nysson Latreille, 1802-1803. Hist. Nat. Crust. Ins., v. 3, p. 340.
Type-species: Crabro spinosus Fabricius. Desig. by Shuckard, 1837.
Nyssonus Rafinesque, 1815. Analyse Nature ou Tabl. Univers, Palermo, p. 124. Emend.

Taxonomy: Menke, Bohart and Richards, 1974. Bul. Zool. Nomencl. 30: 217-218 (petition to
suppress Nysso Latr., 1796, designate Sphex spinosus Forst. as type-species, and place
Nysson Latr., 1802-1803, and spinosus Forst. on Official Lists of Generic and Specific
Names).
argenticus Bohart. South. Calif. and Ariz.

Nysson argenticus Bohart, 1968 (1967). Pan-Pacific Ent. 43: 315, figs. 1, 10, 13, 22. d, @.
aridulus Bohart. South. Calif.

Nysson aridulus Bohart, 1968 (1967). Pan-Pacific Ent. 43: 317, figs. 9, 26. d, 2.

Superfamily SPHECOIDEA 1687

bakeri Bohart. South. Calif.
Nysson bakeri Bohart, 1968 (1967). Pan-Pacifie Ent. 43: 318, figs. 7, 27. 6, °.

chumash Pate. Calif.
Nysson (Nysson) chumash Pate, 1940. Notulae Nat. 63: 1. d.
compactus Cresson. Wash.
Nysson compactus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 278. 2, d.
daeckei Viereck. Mass., N. Y., N. J., Pa., Mich., Iowa, Alta. Host: Gorytes canaliculatus Pack.,

Hoplisoides nebulosus (Pack.).
Nysson daeckei Viereck, 1904. Amer. Ent. Soc., Trans. 30: 238. 6.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 42, figs. 8-12 (larva). —Evans,
1959. Amer. Ent. Soe., Trans. 85: 150 (larva).

Biology: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 43 (host). —Evans, 1966. Compar.
ethology and evolution of sand wasps, pp. 44, 46, 64-65, fig. 36 (behavior, host, egg).
euphorbiae Bohart. South. N. Mex., Ariz., Calif.; Mexico (Sinaloa).
Nysson euphorbiae Bohart, 1968 (1967). Pan-Pacifie Ent. 43: 319, figs. 2, 25. gd, 2.
fidelis Cresson. Wis., Colo., Mont., Wash., Oreg. Host: Gorytes canaliculatus Pack.
Nysson fidelis Cresson, 1882. Amer. Ent. Soc., Trans. 9: 282. 2, d.

Biology: Barth, 1907. Wis. Nat. Hist. Soc., Bul. 5: 145, 147-148 (host).
gagates Bradley. B. C., N. Y., Miss.
Nysson (Brachystegus) gagates Bradley, 1920. Amer. Ent. Soc., Trans. 46: 123. ¢.
hesperus Bohart. B. C., Oreg., Calif., Nev., Idaho, Wyo.
Nysson hesperus Bohart, 1968 (1967). Pan-Pacifie Ent. 43: 320, figs. 8, 14, 18, 23. 5, 9.
lateralis Packard. Chiefly Transit. Zone east of Rockies. Host: Gorytes canaliculatus Paek.?
Nysson laterale Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p. 68.

Nom. nud.
Nysson laterale Packard, 1867. Ent. Soc. Phila., Proc. 6: 440. ¢.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, p. 65 (host ?).
neorusticus Bohart. Calif., Nev., Oreg., Wash., Utah, Wyo.
Nysson neorusticus Bohart, 1968 (1967). Pan-Pacific Ent. 43: 321, figs. 4, 12, 24. d, 9.
pumilus Cresson. Nev., Calif. Host: Hoplisoides hamatus (Handl.)?
Nysson pumilus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 283. 3.
Nysson (Brachystegus) pumilis(!) Bradley, 1920. Amer. Ent. Soc., Trans. 46: 125. 2.

Biology: Powell and Chemsak, 1959. Pan-Pacifie Ent. 35: 200 (host ?).
recticornis Bradley. Wash., Idaho, Calif.
Nysson (Nysson) recticornis Bradley, 1920. Amer. Ent. Soc., Trans. 46: 127. ¢.
rufiventris Cresson. Colo., Mont.
Nysson rufiventris Cresson, 1882. Amer. Ent. Soc., Trans. 9: 283. °.
Plenoculus punctatus Ashmead, 1897. Psyche 8: 338. °.
rufoflavus Bohart. Calif. (Mt. Diablo).
Nysson rufoflavus Bohart, 1968 (1967). Pan-Pacifie Ent. 43: 322, figs. 5, 21. d.
rusticus rusticus Cresson. Colo., Idaho, Wash., Oreg., Calif. Host: Hoplisoides hamatus

(Handl.)?
Nysson rusticus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 282. 2, d.

Biology: Powell and Chemsak, 1959. Pan-Pacific Ent. 35: 200 (host ?). —Evans, 1970. Mus.

Compar. Zool., Bul. 140: 494 (host ?).
rusticus sphecodoides Bradley. Calif. (Claremont).

Nysson (Nysson) sphecodoides Bradley, 1920. Amer. Ent. Soc., Trans. 46: 126. ¢.
schlingeri Bohart. Calif.

Nysson schlingeri Bohart, 1968 (1967). Pan-Pacifie Ent. 43: 323, figs. 6, 15, 16, 20. d, 2.
simplicicornis Fox. N. Y., Pa., Md., D. C., Va., W. Va., Mich., Iowa, Nebr., Mo.

Nysson simplicicornis Fox, 1896. N. Y. Ent. Soe., Jour. 4: 15. ¢.

Brachystegus maculipes Mickel, 1916. Amer. Ent. Soc., Trans. 42: 400. °.

1688 Hymenoptera in America North of Mexico

Nysson (Nysson) minimus Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 408. 2, (¢ misdet.).
Preoce.
Nysson (Nysson) kaskaskia Pate, 1938. Amer. Ent. Soc., Trans. 64: 130. N. name.
subtilis Fox. N. Y., N. J., Pa., Md., D. C., W. Va.
Nysson subtilis Fox, 1896. N. Y. Ent. Soc., Jour. 4: 14. ¢.
timberlakei Bohart. South. Calif., Nev., N. Mex.
Nysson timberlakei Bohart, 1968 (1967). Pan-Pacific Ent. 43: 324, figs. 3, 19, 28. 6, 2.
trichrus (Mickel). Que., N. Y., N. J., Pa., D. C., Va., Iowa, Nebr., Kans., Tex., Ala.
Nysson nigripes Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 269. “3” =
2. Preoce.
Brachystegus trichrus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 400. 9.
Nysson (Nysson) melanopus Pate, 1938. Amer. Ent. Soc., Trans. 64: 130. N. name.
tristis Cresson. B. C., Wash., Oreg., Calif.
Nysson tristis Cresson, 1882. Amer. Ent. Soc., Trans. 9: 281. ¢.

Genus HYPONYSSON Cresson

Hyponysson Cresson, 1882. Amer. Ent. Soc., Trans. 9: 273.
Type-species: Hyponysson bicolor Cresson. Monotypic.

Revision: Pate, 1938. Amer. Ent. Soc., Trans. 64: 131-135, fig. 23. (N. Amer. spp.).
bicolor Cresson. Idaho, Wash., Oreg., Calif.
Hyponysson bicolor Cresson, 1882. Amer. Ent. Soc., Trans. 9: 284. 2.
raui (Rohwer). Fla., Ala., Tenn., Mo., Ark., Okla., Tex. Host: Calliopsis nebrascensis Cwfd.?
Nysson (Hyponysson) raui Rohwer, 1917. U. S. Natl. Mus., Proc. 53: 176. 9.

Biology: Rau, 1922. Acad. Sci. St. Louis, Proe. 24: 18 (host ?).

Genus SYNNEVRUS Costa

Synnevrus Costa, 1859. Fauna Regn. Napoli, Imen. Acul., Nyssonid., p. 16.
Type-species: Synnevrus procerus Costa. Monotypic.
Synneurus Gerstaecker, 1867. Naturforsch. Gesell. Halle 10: 79. Emend.

These wasps occur in the Holarctic Region.

aequalis (Patton). Mass. south to Fla., La., Wis., IIl., Iowa, S. Dak.
Nysson aequalis Patton, 1879. Canad. Ent. 11: 212. 6.
aurinotus (Say). Ga., Ohio, Ind., Ill, Wis., Minn., Iowa, S. Dak., Nebr., Tex., Colo.
Nysson aurinotus Say, 1837. Boston Jour. Nat. Hist. 1: 368.
Nysson Freyi-Gessneri Handlirsch, 1887. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 95:
355. 6.
Nysson angularis Smith, 1908. Nebr. Univ. Studies 8: 340. 3.
Nysson (Nysson) marlatti Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 408. °.
intermedius (Viereck). Tex., N. Mex., Ariz., Calif.
Nysson intermedius Viereck, 1907. Amer. Ent. Soc., Trans. 33: 409. @.
Nysson (Nysson) coyotero Pate, 1940. Notulae Nat. 63: 3. 2, 6.
maderae Bohart. Ariz. (Santa Rita Mts.).
Synneurus(!) maderae Bohart, 1968. Pan-Pacific Ent. 44: 233, fig. 13. 3.
plagiatus (Cresson). U. S., east of Rockies. Host: Sphex ichnewmoneus (L.)?
Nysson plagiatus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 276. 2, d.

Biology: Ristich, 1953. Canad. Ent. 85: 380 (host ?).
Genus EPINYSSON Pate

Nysson subg. E'pinysson Pate, 1935. Ent. News 46: 250.
Type-species: Nysson basilaris Cresson. Orig. desig.

Authenticated hosts for this genus are all in the genus Hoplisoides (Gorytinae). Epinysson oe-
curs only in the New World.

Superfamily SPHECOIDEA 1689

Taxonomy: Bradley, 1920. Amer. Ent. Soc., Trans. 46: 122-123 (key to spp.).

albomarginatus (Cresson). Nev.
Nysson albomarginatus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 278. 2, 6.
arentis Bohart. South. Calif., Nev., Ariz.
Epinysson arentis Bohart, 1968. Pan-Pacifie Ent. 44: 229, figs. 16-18. d, 2.
basilaris basilaris (Cresson). Va., S. C., Ga., Fla.
Nysson basilaris Cresson, 1882. Amer. Ent. Soe., Trans. 9: 281. 9.
basilaris tuberculatus (Handlirsch). N. H., Conn., N. Y., N. J., Pa., Md., D. C., Va., S. C., Wis.,
N. Dak. Host: Hoplisoides nebulosus (Pack.).
Nysson tuberculatus Handlirsch, 1887. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 95: 363.
3.
Nysson tramosericus Viereck, 1904. Amer. Ent. Soc., Trans. 30: 237. 2.
Nysson (Brachystegus) opulentus var. dakotensis Rohwer, 1921. U. S. Natl. Mus., Proce. 59:
ANNI 6)

Taxonomy: Evans, 1959. Amer. Ent. Soe., Trans. 85: 150-151, figs. 36-38 (larva; misdet. as
opulentus Gerst.).

Biology: Evans, 1959. Amer. Ent. Soc., Trans. 85: 151 (host; misdet. as opulentus Gerst.).
—Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 42-44, figs. 34-35
(behavior, host, egg).

bellus (Cresson). Kans., Okla., N. Dak., Colo., Mont., Tex., N. Mex., Ariz., Calif. Host:
Hoplisoides tricolor (Cr.)?
Nysson bellus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 280. 2.
Nysson clarconis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 204. 6.

Biology: Evans, Lin and Yoshimoto, 1954. Ent. News 65: 10 (behavior in host? nest).
desertus Bohart. South. Calif., N. Mex.; Mexico (Sinaloa).
Epinysson desertus Bohart, 1968. Pan-Pacifie Ent. 44: 231, figs. 19, 20. d, 2.
guatemalensis hoplisivora (Rohwer). D. C. to Fla. Host: Hoplisoides costalis (Cr.). Typical
guatemalensis (Roh.) occurs in Cent. America.
Nysson (Brachystegus) hoplisivora Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 96. 2.

Biology: Reinhard, 1925. Wash. Acad. Sci., Jour. 15: 172-177 (host, egg, larval behavior, life
cycle, cocoon). —Reinhard, 1929. The witchery of wasps, pp. 262-271, text fig. (host, larval
behavior, cocoon).
mellipes (Cresson). N. H. to Fla., La., Minn., Iowa, N. Dak., Colo., Mont., Calif., B. C.

Nysson mellipes Cresson, 1882. Amer. Ent. Soc., Trans. 9: 279. 2, 6.

Nysson submellipes Viereck, 1904. Amer. Ent. Soc., Trans. 30: 237. 6.
metathoracicus (Smith). Nebr. (Sioux Co.).

Brachystegus metathoracicus Smith, 1908. Nebr. Univ. Studies 8: 338. 9.
moestus (Cresson). Wash., Calif. Host: Hoplisoides hamatus (Handl.)?

Nysson moestus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 280. ¢.

Nysson (Brachystegus) barberi Rohwer, 1921. U.S. Natl. Mus., Proce. 59: 410. 6.

Biology: Powell and Chemsak, 1959. Pan-Pacifie Ent. 35: 200 (host ?).
opulentus (Gerstaecker). Canada and U.S. in U. Austr., and Sonor. Zones; Panama?
Nysson opulentus Gerstaecker, 1867. Naturhist. Gesell. Halle, Abhandl. 10: 114. d.
Nysson (Brachystegus) seminole Bradley, 1920. Amer. Ent. Soc., Trans. 46: 125. d.
Nysson (Brachystegus) foxti Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 409. ¢.
Nysson (E\pinysson) maiae Pate, 1938. Amer. Ent. Soc., Trans. 64: 137, fig. 26. ¢.
pacificus (Rohwer). Calif. (Santa Barbara).
Nysson (Brachystegus) pacificus Rohwer, 1917. U. S. Natl. Mus., Proce. 53: 249. ¢.
torridus (Bohart). South. Calif., Nev.
Epinysson torridus Bohart, 1968. Pan-Pacifie Ent. 44: 232, fig. 21. d, 2.

Genus ZANYSSON Rohwer

Nysson subg. Zanysson Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 404.
Type-species: Nysson texanus Cresson. Orig. desig.

1690 Hymenoptera in America North of Mexico

The genus is known only from the New World. Circumstantial evidence suggests that species
of Tachytes may serve as hosts.
Revision: Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 404-407 (N. Amer. spp.).

mexicanus (Cresson). Tex. (Brownsville); Mexico.
Paranysson Mexicanus Cresson, 1882. Amer. Ent. Soc., Trans. 9: 275. 2, 6.
Nysson longispinis Cameron, 1905. Amer. Ent. Soc., Trans. 31: 374. 2.

plesius (Rohwer). N. Y., N. J., Md., D. C., Va., N. C., Fla., Ala., Miss., La., Mo., Kans., Tex.,
Ariz. Host: Tachytes d. distinctus Sm.?, T. chrysocercus Roh.?
Nysson (Zanysson) plesia Rohwer, 1921. U. S. Natl. Mus., Proc. 59: 406. °, 3.
Zanysson matinecoc Pate, 1938. Amer. Ent. Soc., Trans. 64: 163. 2, 6.
Zanysson tonto Pate, 1940. Notulae Nat. 63: 6. d.
Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 89-90 (hosts ?).

texanus fuscipes (Cresson). West. U.S.
Paranysson fuscipes Cresson, 1882. Amer. Ent. Soe., Trans. 9: 274. 2, 6.
Nysson aureobalteatus Cameron, 1901. Amer. Ent. Soc., Trans. 27: 313. 6.

texanus texanus (Cresson). U. 8. Host: Tachytes exornatus Fox?
Nysson texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 223. 2, 6.
Biology: Cockerell, 1903. Entomologist 36: 100 (host ?).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 20, figs. I-K (male
genitalia).

Genus FOXIA Ashmead

Foxia Ashmead, 1898. Ent. News 9: 187.
Type-species: Foxia pacifica Ashmead. Monotypic.

This genus is confined to the New World. Its hosts are unknown.
Revision: Pate, 1938. Amer. Ent. Soc., Trans. 64: 143-153, figs. 1, 6, 7, 13-16, 21, 22 (N. Amer.

spp.).
navajo Pate. Okla., Tex., N. Mex., Ariz., Calif.
Foxia navajo Pate, 1938. Amer. Ent. Soc., Trans. 64: 146. 2, 6.

pacifica Ashmead. Calif.
Foxia pacifica Ashmead, 1898. Ent. News 9: 187. 2, ¢.

secunda (Rohwer). Ariz., Calif.
Nysson (Foxia) secunda Rohwer, 1921. U. S. Natl. Mus., Proce. 59: 407. 6.

Genus METANYSSON Ashmead

Metanysson Ashmead, 1899. Canad. Ent. 31: 326.
Type-species: Nysson solani Cockerell. Orig. desig.
Metanysson subg. Huachuca Pate, 1938. Amer. Ent. Soc., Trans. 64: 185.
Type-species: Metanysson (Huachuca) arivaipa Pate. Orig. desig.
The genus occurs only in the New World. Circumstantial evidence suggests that species of
Cerceris may be the hosts.
Revision: Pate, 1938. Amer. Ent. Soc., Trans. 64: 170-188, figs. 2, 3, 5, 9-12, 18-20 (N. Amer.
spp.).
arivaipa Pate. Ariz. Host: Cerceris graphica Sm.?
Metanysson (Huachuca) arivaipa Pate, 1938. Amer. Ent. Soc., Trans. 64: 186. d.
Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, p. 90 (host ?).
coahuila Pate. Calif., Ariz., Tex. Host: Cerceris conifrons Mick.?
Metanysson (Metanysson) coahuila Pate, 1938. Amer. Ent. Soc., Trans. 64: 183. ?.
Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, p. 90 (host ?).

lipan Pate. Tex. (Hudspeth Co.).
Metanysson (Metanysson) lipan Pate, 1938. Amer. Ent. Soc., Trans. 64: 176. ¢.

Superfamily SPHECOIDEA 1691

solani (Cockerell). N. Mex., Ariz.
Nysson solani Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 294. °.
yavapai Pate. Ariz.
Metanysson (Metanysson) yavapai Pate, 1938. Amer. Ent. Soc., Trans. 64: 178. 2, d.

SUBFAMILY GORYTINAE

This large subfamily of ground-nesting wasps is cosmopolitan in distribution. So far as
recorded the species prey upon nymphal and adult Homoptera.

Revision: Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 316-565, 3 pls. (spp.
of world). —Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc., 47: 517-539 (N. Amer. spp.).
—Rohwer, 1921. U.S. Natl. Mus., Proc. 59: 412 (key to gen.).

Taxonomy: Maid] and Klima, 1939. Hym. Cat., Pars 8, Sphecidae 1, pp. 43-114 (world catalog).

Genus OCHLEROPTERA Holmberg

Ochleroptera Holmberg, 1903. Buenos Aires Mus. Nac. Hist. Nat., An. (8) 2: 487.
Type-species: Ochleroptera oblita Holmberg. Monotypic.

Paramellinus Rohwer, 1912. U.S. Natl. Mus., Proce. 41: 469.
Type-species: Gorytes bipunctatus Say. Orig. desig.

The species of this small genus occur only in the New World.

Taxonomy: Pate, 1947. Brooklyn Ent. Soc., Bul. 42: 65-70.

bipunctata (Say). U. and L. Austr. Zones of U. S. Ecology: Nests in sandy to heavier soil,
makes several cells per nest, stores 6-18 prey per cell. Parasite: Miltogrammini sp. Prey:
Aceratagallia sanguinolenta (Prov.), Coelidia olitoria (Say), Colladonus clitellarius
(Say), Japananus hyalinus (Osb.), Macrosteles fascifrons Stal, M. sp., Orientus ishidae
Mats., Paraphlepsius irroratus (Say), Prescottia lobata (Van D.), Scaphytopius sp.,
Strangania apicalis Osb. and Ball; Clastoptera obtusa Say, Philaenus leucophthalmus
L., P. lineatus L.; Cyrtolobus acutus Van D.; Haplaxius radicis Osb.; Psylla annulata
F.; adults are stored more commonly than nymphs.

Gorytes bipunctatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 338.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 47, figs. 23-29 (larva). —Evans,
1959. Amer. Ent. Soc., Trans. 85: 152 (larva).

Biology: Strandtmann, 1945. Ent. Soc. Amer., Ann. 38: 312 (nest, prey). —Evans, 1966.
Compar. ethology and evolution of sand wasps, pp. 75-77, fig. 42 (nest, prey transport, life
cycle, cocoon, parasite). —Evans, 1968. Ent. Soc. Amer., Ann. 61: 1344 (prey).
—Kurczewski and Kurcezewski, 1971. Kans. Ent. Soc., Jour. 44: 335 (prey).

Genus ARGOGORYTES Ashmead

Argogorytes Ashmead, 1899. Canad. Ent. 31: 324.
Type-species: Gorytes carbonarius Smith. Orig. desig.
Argogorytes subg. Archarpactus Pate, 1937. Amer. Ent. Soc., Mem. 9: 4, 10.
Type-species: Sphex mystacea Linnaeus. Orig. desig. Replacement name proposed
unnecessarily for Arpactus Jurine which was suppressed in Op. 135, 1939,
Internat]. Comn. Zool. Nomencl.

Two Palaearctic species nest in the ground, make 6-9 cells per nest, and store 19-27 nymphal
spittlebugs (Aphrophora) per cell.

nigrifrons (Smith). N. S., south to Ga., Wis., Iowa, Kans., Tex.
Gorytes nigrifrons Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 368. 9.
Gorytes Bollii Cresson, 1872. Amer. Ent. Soc., Trans. 4: 225. 9.
Gorytes (Gorytes) neglectus Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 567. 2.
sapellonis (Baker). N. Mex. (Sapello Canyon).
Gorytes sapellonis Baker, 1907. Invertebrata Pacifica 1: 161. 2.

1692 Hymenoptera in America North of Mexico

Genus DIENOPLUS Fox

Arpactus Jurine, 1807. Nouv. Meth. Class. Hym. Dipt., p. 192. Preoce.
Type-species: Arpactus formosus Jurine. Desig. by Shuckard, 1837.

Harpactus Shuckard, 1837. Essay on Indig. Fossor. Hym., p. 221. Emend.

Harpactes Dahlbom, 1843. Hym. Europaea, v. 1, p. 147. Emend. Preoce.

Dienoplus Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc., 45: 548.
Type-species: Dienoplus pictifrons Fox. Orig. desig.

Most species occur in the Holarctie Region, but a few have been recorded from the Ethiopian
and Oriental Regions. A European species makes 2-15 cells per nest in sandy soil. Recorded prey
are leafhoppers and spittlebugs, both nymphs and adults.

citipes Krombein. Fla., (Orlando, Arcadia). Ecology: Nests in flat sandy areas.
Dienoplus citipes Krombein, 1954. Amer. Ent. Soc., Trans. 80: 20, figs. 9, 11, 13. 6, oF
gyponae (Williams). Kans., N. Mex., Alta. Ecology: Nests in sand, makes more than 2 cells per
nest, stores 4 prey per cell. Prey: Prairiana cinerea (Uhl.) adults and nymph.
Harpactus gyponae Williams, 1914 (1913). Kans. Univ. Sei. Bul. 8: 223. ?.
Biology: Williams, 1914 (1913). Kans. Univ. Sei. Bul. 8: 225-226, pl. 33, fig. 5 (nest, prey
transport).

mendicus (Handlirsch). B. C., Idaho.
Gorytes mendicus Handlirsch, 1893. K. K. Naturhist. Hofmus., Ann. 8: 278. 9.

pictifrons Fox. Idaho, Wash., Calif. Predator: Philanthus pulcher D. T.
Dienoplus pictifrons Fox, 1894 (1893). Acad. Nat. Sci. Phila., Proc., 45: 549. 2, 3.
Harpactus howardi Ashmead, 1899. Ent. News 10: 9. °.

Genus HAPALOMELLINUS Ashmead

Hapalomellinus Ashmead, 1899. Canad. Ent. 31: 300.
Type-species: Gorytes eximius Provancher. Orig. desig.

This small genus occurs only in the desert areas of southwestern North America.

Taxonomy: Bohart, 1971. Biol. Soc. Wash., Proc. 83: 452 (key to spp.).

albitomentosus (Bradley). Tex., N. Mex., Ariz., Calif. Ecology: Nests in level, dry sandy soil,
makes 1-2 cells per nest, one of which may be just a storage cell, and stores 14-15
leafhoppers per cell. Parasite: Senotainia sp. in trilineata (Wulp) complex? Prey:
Stragania robusta Uhl. mostly adults, but a few nymphs.
Gorytes eximius Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 274. ?.
Preoce.
Gorytes (Arpactus) albitomentosus Bradley, 1920. Amer. Ent. Soc., Trans. 46: 121. 2, d.

Biology: Cazier and Mortenson, 1965. Wasmann Jour. Biol. 22: 261-276 (nest, prey, parasite
2).

pulvis Bohart. South. N. Mex., Ariz. and Calif.

Hapalomellinus pulvis Bohart, 1971. Biol. Soc. Wash., Proc. 83: 451. 2, d.
teren Pate. Ariz., Calif.

Hapalomellinus teren Pate, 1946. Ent. News 57: 237. 2.

Genus TRICHOGORYTES Rohwer

Trichogorytes Rohwer, 1912. U. S. Natl. Mus., Proe. 41: 469.
Type-species: Trichogorytes argenteopilosus Rohwer. Orig. desig.
The two known species occur in the southwestern deserts.
Revision: Pate, 1946. Brooklyn Ent. Soc., Bul. 41: 15-17.

argenteopilosus Rohwer. Ariz., Calif.
Trichogorytes argenteopilosus Rohwer, 1912. U. S. Natl. Mus., Proc. 41: 470. °.

Superfamily SPHECOIDEA 1693

cockerelli (Ashmead). N. Mex. (Mesilla). Ecology: Makes up to 2 cells per nest in fine-grained
sand in dunes, stores 8-10 prey per cell. Prey: Exitianus exitiosus (Uhl.), Circulifer
tenellus (Baker), Norvellina sp.; stores mostly adults.
Harpactus cockerelli Ashmead, 1899. Ent. News 10: 10. 2.

Biology: Evans, 1976. Ent. News 87: 33-37, 4 figs. (nest, prey).

Genus GORYTES Latreille

Gorytes Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, Tabl. Meth., p. 180.
Type-species: Mellinus quinquecinctus Fabricius. Monotypic.
Arpactus Panzer, 1805. Faunae Ins. German., heft 98, No. 17.
Type-species: Mellinus quadrifasciatus Fabricius. Monotypic.
Arpactus Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, p. 164. Preoce.
Type-species: Mellinus quadrifasciatus Fabricius. Desig. by Pate, 1937.
Euzonia Stephens, 1829. System. Cat. Brit. Ins., p. 363.
Type-species: Mellinus quinquecinctus Fabricius. Desig. by Pate, 1937.
Hoplisus Lepeletier, 1832. Soc. Ent. France, Ann. 1: 61.
Type-species: Hoplisus quinquecinctus of Lepeletier. Desig. by Westwood, 1839.
Euspongus Lepeletier, 1832. Soc. Ent. France, Ann. 1: 66.
Type-species: Euspongus laticinctus Lepeletier. Desig. by Westwood, 1839.

This genus occurs in the Holarctic and Ethiopian Regions. Prey records include Cicadellidae,
Fulgoridae, Cercopidae and Membracidae. The multicellular nests may contain up to four cells,
and 4-19 prey may be stored per cell.

albosignatus Fox. N. Dak., S. Dak., Nebr., Colo., Mont., Idaho.
Gorytes albosignatus Fox, 1892. Canad. Ent. 24: 152. 2, 3.
angustus (Provancher). Nev., Calif., Wash., B. C.
Hoplisus angustus Provancher, 1895. Nat. Canad. 22: 141.“2” = 6.
Hoplisus angustatus(!) Ashmead, 1899. Canad. Ent. 31: 328.
atricornis Packard. Transcont. in Transit. and U. Austr. Zones. Prey: Aphrophora parallela
(Say); Cyrtolobus tuberosus (Fairm.); adults.
Odynerus(?) atricornis Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p.
68. Nom. nud.
Gorytes atricornis Packard, 1867. Ent. Soc. Phila., Proc. 6: 428. 2, 6.
Gorytes rugosus Packard, 1867. Ent. Soc. Phila., Proc. 6: 427. ¢.
Gorytes decorus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc., 45: 5385. 2, d.
Hoplisus elegantulus Smith, 1908. Nebr. Univ. Studies 8: 346. ¢.

Biology: Pate, 1946. Brooklyn Ent. Soc., Bul. 41: 99 (prey). —Evans, 1966. Compar. ethology
and evolution of sand wasps, p. 67 (prey).
atrifrons Fox. Colo., Nev., Calif.
Gorytes atrifrons Fox, 1892. Canad. Ent. 24: 151. 2, 6.
canaliculatus Packard. Transcont. in Transit. and U. Austr. and Sonor. Zones. Ecology: Nests
both in sand and compacted soil, makes 1-4 cells per nest, stores 6-26 prey per cell.
Parasite: Metopia argyrocephala (Meig.), Senotainia trilineata (Wulp)?, Phrosinella
fulvicornis (Coq.)?, P. fumosa Allen?; Timulla leona (Bl.)?; Nysson fidelis Cr., N.
daeckei Vier., N. lateralis Pack. Prey: Idiocerus apache Ball and Prkr., I. cinctus
DeLong and Caldw., J. lachrymalis Fitch, I. pallidus Fitch, I. perplexus Gill. and Baker,
I. populi suturalis Fitch, J. snow? Gill. and Baker, J. stigmaticalis Lewis, J, spp.,
Macropsis viridis (Fitch), M. sp., Norvellina helenae Ball, Oncopsis sp., Orientus ishidae
Mats., Paraphlepsius sp., Stragania alabamensis Baker; Haplaxius pictifrons Stal;
nymphs and adults.
Gorytes canaliculatus Packard, 1867. Ent. Soc. Phila., Proc. 6: 428. 9.
Gorytes geminus Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 478. °,
3

Gorytes asperatus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc., 45: 534. 2, ¢.

Hoplisus corrugis Mickel, 1918 (1917). Nebr. Univ. Studies 17: 319. 6.

Gorytes (Hoplisus) carrugis(!) Maid] and Klima, 1939. Hym. Cat., Pars 8, Sphecidae, v. 1, p.
101.

1694 Hymenoptera in America North of Mexico

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 43-44, figs. 13-22 (larva).

Biology: Barth, 1907. Wis. Nat. Hist. Soc., Bul. 5: 141-149, 3 figs. (nest, prey, parasite;
observations probably based on both canaliculatus and Hoplisoides nebulosus (Pack.)).
—Krombein, 1964 (1963). Brooklyn Ent. Soc., Bul. 58: 119 (nest, prey). —Evans, 1966.
Compar. ethology and evolution of sand wasps, pp. 58-66, figs. 38, 39 (nest, prey, egg, life
cycle, parasites). —Evans, 1968. Ent. Soc. Amer., Ann. 61: 1344 (prey). —Evans, 1970. Mus.
Compar. Zool., Bul. 140: 494 (nest, prey, parasites). —Kurezewski and Kurczewski, 1971.
Kans. Ent. Soc., Jour. 44: 335-336 (prey). —Alceock, 1973. Wasmann Jour. Biol. 31: 332-333,
figs. 6, 10 (nest, prey transport). —Powell, 1974. Kans. Ent. Soc., Jour. 47: 1-7, 3 figs. (nest,
prey transport, life cycle).

cochisensis Bohart. South. Ariz.
Gorytes cochisensis Bohart, 1971. Biol. Soc. Wash., Proc. 83: 450. 2, d.
deceptor Krombein. N. H. south to Va., west to Ont., Minn., Nebr., Kans. Prey: Spissistylus
constans (Wlk.) adult.
Gorytes (Gorytes) deceptor Krombein, 1958. Ent. Soc. Wash., Proc. 60: 62. 2, d.

Biology: Krombein, 1958. Ent. Soc. Wash., Proc. 60: 64 (prey).

dorothyae dorothyae Krombein. Md., N. C., La.
Gorytes (Gorytes) dorothyae Krombein, 1950 (1949). Elisha Mitchell Sei. Soc., Jour. 65: 269.
on

Taxonomy: Krombein, 1953. Wasmann Jour. Biol. 10: 335. 9.

dorothyae russeolus Krombein. Fla.
Gorytes (Gorytes) dorothyae russeolus Krombein, 1954. Amer. Ent. Soc., Trans. 80: 23. d,
Q

flagellatus Bohart. Calif., Nev., Wash., B. C. Idaho, Wyo. ¥
Gorytes flagellatus Bohart, 1971. Biol. Soc. Wash., Proc. 83: 448. 6, °.
limbellus Bohart. Calif.
Gorytes limbellus Bohart, 1971. Biol. Soc. Wash., Proc. 83: 447. 6, °.
meateei Krombein and Bohart. N. Y., Pa., Md., D. C., Va.
Gorytes (Gorytes) mcateei Krombein and Bohart, 1962. Biol. Soc. Wash., Proc. 75: 15. 9.

nevadensis Fox. Nev., Colo., Idaho.
Gorytes nevadensis Fox, 1892. Canad. Ent. 24: 150. 2, d.
prosopis Bohart. Calif., Oreg.
Gorytes prosopis Bohart, 1971. Biol. Soc. Wash., Proc. 83: 449. °.
provancheri Handlirsch. Wash., Oreg., Calif., Nev., Utah.
Gorytes laticinctus Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 274. d.
Preoce.
Gorytes Provancheri Handlirsch, 1895. Akad. Wiss. Wien, Math.-Nat. KI., Sitzber. 104: 945.
N. name.
simillimus Smith. N. S. to B. C. south to Ga., Ill., Kans. and Colo. Ecology: Nests in sand.
Prey: Gyponana flavolineata (Fitch), G. melanota Spang., G. octolineata (Say), G. spp.,
Scaphoideus productus Osb.; mostly adults but a few nymphs.
Gorytes simillimus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 367. °.
Gorytes ephippiatus Packard, 1867. Ent. Soc. Phila., Proce. 6: 426. d.
Gorytes (Pseudoplisus) gyponacinus Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 568. °, d.

Biology: Rohwer, 1911. U. S. Natl. Mus., Proc. 40: 568 (prey). —Krombein, 1936. Ent. News
47: 93-94 (prey transport). —Krombein, 1952. Amer. Ent. Soc., Trans. 78: 95 (nest, prey).
—Evans, 1966. Compar. ethology and evolution of sand wasps, p. 66 (nest, prey).
—Kurezewski and Kurcezewski, 1971. Kans. Ent. Soc., Jour. 44: 336 (prey).

umatillae Bohart. West. U. S.
Gorytes vicinus Handlirsch, 1893. K. K. Naturhist. Hofmus., Ann. 8: 279. 2. Preoce.
Gorytes wmatillae Bohart, 1971. Biol. Soc. Wash., Proc. 83: 450. N. name.

Genus PSEUDOPLISUS Ashmead
Pseudoplisus Ashmead, 1899. Canad. Ent. 31: 323.

Superfamily SPHECOIDEA 1695

Type-species: Gorytes floridanus Fox. Orig. desig.
Laevigorytes Zavadil, 1948. In Zavadil and Snoflak, Ent. Prirucky Ent. Listu, no. 13, p. 66.
Type-species: Gorytes kohlii Handlirsch. Monotypic.

Most of the species are Nearctic but a few have been described from the Ethiopian and
Palaearctic Regions. The only known prey record is of a spittlebug for one of the Ethiopian spe-
cies.

Taxonomy: Bohart, 1969 (1968). Kans. Ent. Soc., Jour. 41: 494-496 (key to spp. of Phaleratus
Group). —Bohart, 1969. Kans. Ent. Soc., Jour. 42: 392-394 (key to spp. of Montanus,
Abdominalis, Fasciatus, Venustus and Smithii Groups).
abdominalis (Cresson). Colo., Tex.

Gorytes abdominalis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 474. 2, d.
apicalis (Smith). N. J., S. C., Tex., N. Mex., Nebr.

Gorytes apicalis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 369. d.

Gorytes propinquus Cresson, 1868. Amer. Ent. Soc., Trans. 1: 379. “2” = d.
butleri Bohart. Ariz.

Pseudoplisus butleri Bohart, 1969. Kans. Ent. Soc., Jour. 42: 397, figs. 1, 7. d.

californicus Bohart. Calif.
Pseudoplisus californicus Bohart, 1969. Kans. Ent. Soc., Jour. 42: 401, figs. 11, 12. d, °.

catalinae Bohart. Ariz. (Santa Catalina Mts.).
Pseudoplisus catalinae Bohart, 1969. Kans. Ent. Soc., Jour. 42: 395, figs. 18, 17. d, 2.

claripennis Bohart. Ariz. (Santa Rita Mts.).
Pseudoplisus claripennis Bohart, 1969. Kans. Ent. Soc., Jour. 42: 403. d.

divisus (Smith). Va. to Ga., La., Tex., Nebr.
Gorytes divisus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 370. d.
Gorytes bipartitus Handlirsch, 1888. Akad. Wiss. Wien Math.-Nat. K1., Sitzber.-97: 521. °,
3

Pseudoplisus varipunctus Smith, 1908. Nebr. Univ. Studies 8: 348. °.
fasciatus (Fox). Colo., Ariz.

Gorytes fasciatus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc., 45: 589. d.
flavidulus Bohart. South. Calif.

Pseudoplisus flavidulus Bohart, 1969. Kans. Ent. Soc., Jour. 42: 402. d.

imperialis Bohart. Calif. (Imperial Co.).

Pseudoplisus imperialis Bohart, 1969. Kans. Ent. Soc., Jour. 42: 403. d.
nigricomus Bohart. South. Ariz.

Pseudoplisus nigricomus Bohart, 1969. Kans. Ent. Soc., Jour. 42: 398, figs. 5, 10. d, °.
ocellatus Bohart. South. Calif.; Mexico (Baja California).

Pseudoplisus ocellatus Bohart, 1969. Kans. Ent. Soe., Jour. 42: 401. d, 9.

oraclensis Bohart. South. Ariz.
Pseudoplisus oraclensis Bohart, 1969 (1968). Kans. Ent. Soe., Jour. 41: 498, figs. 4, 8. d, °.
phaleratus (Say). Canada and U. S. in Transit. and Austr. Zones; P. E. I. west to N. Dak. and
Colo., south to Fla. and Ariz., south in Mexico to Guerrero.
Odynerus(?) flavicornis Harris, 1835. In Hitchcock, Rpt. Geol. Mineral. Bot. Zool. Mass., p.
68. Nom. nud.
Gorytes phaleratus Say, 1837. Boston Jour. Nat. Hist. 1: 367.
Gorytes fulvipennis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 367. °.
Gorytes modestus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 473. d.
Gorytes rufo-luteus Packard, 1867. Ent. Soc. Phila., Proc. 6: 425. 2, 6.
Gorytes flavicornis Packard, 1867. Ent. Soc. Phila., Proc. 6: 429. 2, d.
Gorytes alticola Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 81, pl. 5, fig. 21. d, 9.
Gorytes alpestris Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 83, pl. 5, fig. 22. d, 2.
Gorytes subaustralis Viereck, 1908 (1907). Amer. Ent. Soc., Trans. 33: 398. 2.
Gorytes papagorum Viereck, 1908 (1907). Amer. Ent. Soc., Trans. 33: 400. 2, d.

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 21, figs. H-K (male
genitalia).

1696 Hymenoptera in America North of Mexico

rufomaculatus (Fox). Nebr., Kans., Colo., Mont.

Gorytes rufomaculatus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc., 45: 538. ?, d.
smithii floridanus (Fox). Fla.

Hoplisus foveolata Fox, 1890. Ent. News 1: 106. 2. Preoce.

Gorytes floridanus Fox, 1891. Ent. News 2: 196. N. name.
smithii smithii (Cresson). Md., N. C., Fla., Ill., Nebr., Kans., Okla., La.

Gorytes Smithii Cresson, 1880. Amer. Ent. Soc., Trans. 8: Proc., p. xviii. “2” = 6.

Pseudoplisus infumatus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 402. ¢.
tanythrix Bohart. Okla., Tex.

Pseudoplisus tanythrix Bohart, 1969. Kans. Ent. Soc., Jour. 42: 399, figs. 6, 8, 14. d.
venustus (Cresson). Tex., N. Mex., Ariz., Utah, Colo.; Mexico (Chihuahua).

Gorytes venustus Cresson, 1865. Ent. Soc. Phila., Proc. 4: 472. 2, d.

Gorytes (Pseudoplisus) venustiformis Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 568. 3.
werneri Bohart. Ariz., Colo.

Pseudoplisus werneri Bohart, 1969 (1968). Kans. Ent. Soc., Jour. 41: 500, figs. 3, 11, 14. @,

3.

Genus LESTIPHORUS Lepeletier

Lestiphorus Lepeletier, 1832. Soc. Ent. France, Ann. 1: 70.
Type-species: Crabro bicinctus Rossi. Desig. by Internat]. Comn. Zool. Nomencl.,
Op. 675, 1963.
Lestophorus Agassiz, 1847. Nomencl. Zool., p. 208. Emend.
Hypomellinus Ashmead, 1899. Canad. Ent. 31: 299.
Type-species: Gorytes rufocinctus Fox. Orig. desig.
Mellinogastra Ashmead, 1899. Canad. Ent. 31: 300.
Type-species: Gorytes mellinoides Fox. Orig. desig.

Most species are Holarctic but one each is known from the Neotropical and Ethiopian Re-
gions. One European species is reported to prey upon spittlebugs.

Revision: Pate, 1946 (1945). Canad. Ent. 77: 210-213 (N. Amer. spp.).

Taxonomy: van der Vecht, 1961. Bul. Zool. Nomencl. 18: 340-341 (request to suppress Crabro
bicinctus F. and to place Lestiphorus Lep. and Crabro bicinctus Rossi on Official Lists of
Generic and Specific Names).

cockerelli (Rohwer). Mass., N. Y., Mich., S. Dak., Colo.
Gorytes cockerelli Rohwer, 1909. Ent. News 20: 371. 6.
Mellinogastra williamsi Mickel, 1916. Amer. Ent. Soc., Trans. 42: 402. ¢.
mellinoides (Fox). Tex. (? Bosque Co.).
Gorytes mellinoides Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 45: 524. 9.
piceus (Handlirsch). N. Dak., Idaho, Wash., B. C.
Gorytes piceus Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 455. ¢.
Gorytes rufocinctus Fox, 1892. Canad. Ent. 24: 153. 2.

Genus ORYTTUS Spinola

Oryttus Spinola, 1836. Soc. Ent. France, Bul. 5: xxiii.
Type-species: Arpactus concinnus (Rossi). Monotypice.
Agraptus Wesmael, 1852. Acad. Roy. Sci. Belg., Bul. 19: 108.
Type-species: Gorytes concinnus of Vander Linden. Monotypic.
Harpactostigma Ashmead, 1899. Canad. Ent. 31: 299.
Type-species: Hoplisus velutinus Spinola. Orig. desig.
Harpactostigma subg. Arcesilas Pate, 1938. Amer. Ent. Soc., Trans. 64: 60.
Type-species: Gorytes mirandus Fox. Orig. desig.

Most species of this small genus occur in the Holarctic Region, but a few are Ethiopian and

Neotropical. Several prey records are of Fulgoridae, but one European species also uses Cicadel-
lidae.

Revision: Pate, 1938. Amer. Ent. Soc., Trans. 64: 57-77 (New World spp.).

Superfamily SPHECOIDEA 1697

Taxonomy: Bohart, 1968. Biol. Soc. Wash., Proc. 81: 431-438, 17 figs. (synopsis of New World
taxa).
gracilis arapaho (Pate). Colo., Kans., Tex.
Harpactostigma (Arcesilas) arapaho Pate, 1938. Amer. Ent. Soc., Trans. 64: 67. °.
Harpactostigma (Arcesilas) rutilum Pate, 1938. Amer. Ent. Soc., Trans. 64: 69. d.
gracilis gracilis (Patton). U.S. in U. and L. Austr. Zones, east of 100th meridian.
Hoplisus gracilis Patton, 1879. Canad. Ent. 11: 210. 2, 6.
laminiferus (Fox). Calif., Wash., Idaho, Utah. Ecology: Nests in vertical clay bank, stores at
least 5 prey per cell. Prey: Scolops hesperius Uhl. ?, nymphs and adults.
Gorytes ruficornis Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 273. 2, 3.
Preoce.
Gorytes laminiferus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 45: 532. 3.
Hypomellinus flavicornis Baker, 1907. Invertebrata Pacifica 1: 162. 3.
Gorytes (Hoplisus) rufulicornis Maid] and Klima, 1939. Hym. Cat., Pars 8, Sphecidae, v. 1,
p. 102. N. name.

Biology: Gittins, 1958. Pan-Pacific Ent. 34: 142 (nest, prey).

mirandus (Fox). Nev., Calif., Wash.

Gorytes mirandus Fox, 1892. Canad. Ent. 24: 152. 6.

Hoplisoides mirandas(!) Ashmead, 1899. Canad. Ent. 31: 328.
umbonatus (Baker). Calif.

Hoplisoides umbonatus Baker, 1907. Invertebrata Pacifica 1: 163. 3.

Gorytes (Hoplisoides) femoratus Bradley, 1920. Amer. Ent. Soc., Trans. 46: 119. ¢.
yumae Bohart. South. Ariz.

Oryttus ywmae Bohart, 1968. Biol. Soc. Wash., Proce. 81: 436, figs. 4, 9, 13, 17. 6, 9.

Genus PSAMMALETES Pate

Hoplisoides subg. Psammaletes Pate, 1936. Amer. Ent. Soc., Trans. 62: 49.
Type-species: Gorytes bigeloviae Cockerell and Fox. Orig. desig.

This small genus occurs only in the Nearctic Region.

Revision: Pate, 1936. Amer. Ent. Soc., Trans. 62: 49-56.
bigeloviae (Cockerell). N. Mex., Ariz.; Mexico.
Gorytes bigeloviae Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila., Proc. 49:
139. 3.

crucis (Cockerell). Iowa, Nebr., N. Mex.; north. Mexico.
Gorytes crucis Cockerell, 1897. In Cockerell and Fox, Acad. Nat. Sci. Phila., Proc. 49: 140.
2

Hypomellinus venustus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 403. ¢.
Hypomellinus tricinctus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 404. °.
mexicanus (Cameron). N. Y., N. J., Md., D. C., Va., N. C., Ga., La., Ind.; Mexico (Yucatan).
Prey: Ormenoides venusta (Melichar).
Gorytes mexicanus Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 76, pl. 5, fig. 15. °.
Psammaletes pechumani Pate, 1936. Amer. Ent. Soc., Trans. 62: 53. 9.

Biology: Pate, 1946. Brooklyn Ent. Soc., Bul. 41: 99 (prey).
Genus SPHECIUS Dahlbom
Taxonomy: Pate, 1936. Brooklyn Ent. Soc., Bul. 31: 198-200 (key to subg.).
Genus SPHECIUS Subgenus SPHECIUS Dahlbom

Sphecius Dahlbom, 1843. Hym. Europaea, v. 1, p. 154.
Type-species: Sphecius speciosus Dahlbom. Monotypic.
Hogardia Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 288.
Type-species: Hogardia rufescens Lepeletier. Type by tautonymy (=Stizus hogardii
Latreille). ;

1698 | Hymenoptera in America North of Mexico

The typical subgenus occurs only in the New World. These large wasps are commonly called
cicada killers. They frequently nest gregariously and dig burrows which may be as long as 4
feet, have a number of branches each of which may terminate in a cluster of 3 cells. One to as
many as 4 prey may be stored per cell depending upon the size of the prey and the sex to be pro-
vided for.

Revision: Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 264-266 (N. Amer. spp.).
convallis Patton. U. S. west of 100th meridian; Mexico (Baja California). Prey: Diceroprocta
apache (Davis), Tibicen pruinosa (Say).
Sphecius speciosus var. convallis Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 343.

Sphecius raptor Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98: 461. 9, 6.
N. name.

Biology: Krombein, 1951. U. S. Dept. Agr., Monog. 2: 987 (prey). —Krombein, 1958. U. S.
Dept. Agr., Monog. 2, Sup. 1, p. 193 (prey).
grandis (Say). Tenn., Mo., Ark., Tex., Kans., N. Mex., Ariz., Utah, Nev., Wash., Calif., south to
Central America. Parasite: Dasymutilla klugii (Gray)? Prey: Tibicen dealbata (Davis).
Stizus grandis Say, 1823. West. Quart. Rptr. 2: 77.
Stizus fervidus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 223. 9.
Stizus nevadensis Cresson, 1874. Amer. Ent. Soce., Trans. 5: 99. ¢.

Biology: Bradley, 1908. Ent. Soc. Amer., Ann. 1: 129 (sleeping aggregation). —Bradley, 1920.
Ent. News 31: 112-113 (nest, parasite ?). —Evans, 1966. Compar. ethology and evolution of
sand wasps, p. 110 (prey transport). —Aleock, 1975. Jour. Nat. Hist. 9: 561-566, 3 figs.
(male territorial and mating behavior).

hogardii hogardii (Latreille). South Fla.; West Indies. Another subspecies occurs in the
Bahamas..
Stizus Hogardii Latreille, 1809. Gen. Crust. Ins., v. 4, p. 100, pl. 13. 9.
Hogardia rufescens Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 289. 9°. N. name.
speciosus (Drury). U.S. in U. and L. Austr. Zones east of Rockies, south into Mexico. Ecology:
Nests gregariously in light clay to sandy soil, the tunnel with several branches each
with one or more cells and with an average of 15.8 cells per nest, usually stores 1-2 prey
per cell. Parasite: Senotainia trilineata (Wulp), Metopia argyrocephala (Meigen). Prey:
Tibicen canicularis Harr., T. chloromera Wlkr., T. dorsata Say, T. linnei Sm. and
Grossb., T. lyricen DeG., T. marginalis Wlkr., T. pruinosa Say, T. robinsoniana Davis,
T. sayi (Grossb.), T. vitripennis Say. This is commonly known as the cicada killer.
Sphex speciosus Drury, 1778. Illus. Nat. Hist., v. 2, p. 71. 9.
Vespa tricincta Fabricius, 1775. Systema Ent., p. 363.
Stizus vespiformis Latreille, 1818. Tabl. Encycel. et Meth., pt. 24, pl. 382, fig. 6.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 46, figs. 30-36 (larva).

Biology: Riley, 1892. Insect Life 4: 248-252 (nest, prey). —Davis, 1920. Brooklyn Ent. Soc.,
Bul. 15: 128-129 (mating, prey transport). —Reinhard, 1929. The witchery of wasps, pp.
18-60, 4 pls. (nest, prey, cocoon, life cycle, mating, parasite). —Dow, 1942. Ent. Soc. Amer.,
Ann. 35: 310-317 (nest, prey). —Dambach and Good, 1943. Ohio Jour. Sci. 43: 32-41, figs. 1-6
(nest, prey, life history). —Lin, 1963. Behavior 20: 115-133, 6 figs. (territoriality of males).
—Lin, 1964 (1963). Brooklyn Ent. Soc., Bul. 58: 121-123 (fighting of nesting females).
—Lin, 1966. Anim. Behaviour 14: 130-131, pl. 6 (mating). —Evans, 1966. Compar. ethology
and evolution of sand wasps, pp. 92-110, figs. 52-58 (nest, prey, egg, life cycle, parasites).
—Lin, 1967. Science 157: 1334-1335, 1 fig. (sex roles in mating). —Lin, 1972. In Michener
and Lin, Quart. Rev. Biol. 47: 139-140 (communal nest possession by several females).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 18, figs. E-O (male
genitalia).
Genus TANYOPRYMNUS Cameron

Tanyoprymnus Cameron, 1905. Amer. Ent. Soc., Trans. 31: 375.
Type-species: Tanyoprymnus longitarsis Cameron. Monotypic.
Ceratostizus Rohwer, 1921. U.S. Natl. Mus., Proc. 59: 412.

Superfamily SPHECOIDEA 1699

Type-species: Gorytes moneduloides Packard. Orig. desig.

Taxonomy: Pate, 1935. Ent. News 46: 249-250.
moneduloides (Packard). U. S. south into Mexico. Ecology: Nests in vertical sand banks. Prey:
Rhynchomitra microrhina (Wlkr.) nymphs and adult; Scolops sulcipes (Say) adults, S.
sp.? nymph.
Gorytes moneduloides Packard, 1867. Ent. Soe. Phila., Proe. 6: 431. ¢.
Gorytes Belfragei Cresson, 1872. Amer. Ent. Soc., Trans. 4: 224. °.
Tanyoprymnus longitarsis Cameron, 1905. Amer. Ent. Soc., Trans. 31: 376. d.
Biology: Krombein, 1959. Ent. Soe. Wash., Proc. 61: 195 (nest, prey transport). —Evans, 1966.
Compar. ethology and evolution of sand wasps, p. 80 (prey).

Genus ARIGORYTES Rohwer

Arigorytes Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 469.
Type-species: Gorytes Coquillettii Fox. Orig. desig.

This is a small genus occurring only in the deserts of western North America. Nothing is
known of the biology.

Revision: Pate, 1947. Canad. Ent. 79: 51-56.

Taxonomy: Bohart, 1971. Biol. Soc. Wash., Proc. 83: 447 (key to N. Amer. spp.).

coachellae Bohart. Calif., Ariz., Nev., Colo.; Mexico (Baja California).

Arigorytes coachellae Bohart, 1971. Biol. Soc. Wash., Proc. 83: 445. 6, 9.
coquillettii (Fox). Calif.

Gorytes Coquillettii Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 47: 531. °.
insolitus (Fox). Nev., Calif., Oreg.

Gorytes insolitus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 47: 532. 6.

Hoplisoides clavatus Baker, 1907. Invertebrata Pagifica 1: 165. d.
ruficrus Bohart. Calif. at 5000-7000 ft. in central Sierra Mts.

Arigorytes ruficrus Bohart, 1971. Biol. Soc. Wash., Proc. 83: 446. d, 9.
smohalla Pate. Idaho, Wash., Calif.

Arigorytes smohalla Pate, 1947. Canad. Ent. 79: 54. 2, ¢.

Genus XEROGORYTES Bohart

Xerogorytes Bohart, 1976. In Bohart and Menke, Sphecid wasps of the world, p. 517.
Type-species: Arigorytes anaetis Pate. Orig. desig.

This is a monotypic genus occurring only in the southwestern United States. The biology is
unknown.

anaetis (Pate). Ariz., southwest. N. Mex.
Arigorytes anaetis Pate, 1947. Canad. Ent. 79: 55. °.

Genus HOPLISOIDES Gribodo

Hoplisoides Gribodo, 1884. Ent. Soe. Ital., Bol. 16: 276.
Type-species: Hoplisoides intricans Gribodo. Monotypic.
Icuma Cameron, 1905. Entomologist 38: 21.
Type-species: Jcwma sericea Cameron. Monotypic.

This large genus occurs in all of the major zoogeographic regions except the Australian. The
species prefer to nest in sandy soil and make relatively short burrows ending in one or several
cells. The recorded prey are all Homoptera belonging to the Membracidae, Cicadellidae or Ful-
goridae; each species prefers prey of a single family usually.

carinatus Bohart. Ariz.; Mexico (Sonora).
Hoplisoides carinatus Bohart, 1968. Ent. Soc. Wash., Proc. 70: 287. 6, °.

cazieri Bohart. South. Ariz.
Hoplisoides cazieri Bohart, 1968. Ent. Soc. Wash., Proc. 70: 288. 6, @.

1700 Hymenoptera in America North of Mexico

confertus (Fox). Western North America.
Gorytes confertus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 47: 525. 2, d.
Gorytes (Hoplisoides) imperialensis Bradley, 1920. Amer. Ent. Soe., Trans. 46: 117. d.
costalis costalis (Cresson). Alaska, U. S. east of 100th meridian. Ecology: Nests in sand, 1-2
cells per nest, 3-9 prey per cell. Parasite: Amobia aurifrons (Tns.); Nysson hoplisivora
Roh.; Notozus viridicyaneus (Nort.). Prey: Archasia galeata (F.), Ceresa bubalus F., C.
palmeri Van D., Thelia bimaculata F., T. sp., Telamona monticola F., T. tristis Fitch, T.
unicolor Fitch, Glossonota crataegi Fitch, Cyrtolobus arcuatus Emm.?, Vanduzea
arcuata (Say), Platycotis vittata F., Campylenchia latipes Say, Stictocephala borealis
(Fairm.); only adults are used as prey.
Gorytes costalis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 225. °.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 151, figs. 39, 62 (larva).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 19 (nest, prey). —Reinhard, 1925. Wash.
Acad. Sci., Jour. 15: 172-177 (nest, prey transport, egg, life cycle, cocoon, parasite (misdet.
as Pachyophthalmus signatus Meig.)). —Reinhard, 1929. The witchery of wasps, pp.
235-260, 3 figs. (nest, prey transport, life cycle, cocoon, parasite (misdet. as
Pachyophthalmus signatus (Meig.)). —Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 286
(nest, prey). —Krombein, 1959. Ent. Soc. Wash., Proc. 61: 196 (nest, prey, egg, life cycle).
—Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 49-53, fig. 37 (nest,
prey).

costalis pygidialis (Fox). Western U.S. in U. and L. Sonor. Zones; Mexico. Prey: Ceresa sp.
Gorytes pygidialis Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 47: 528. 2, d.

Biology: Mickel, 1918 (1917). Nebr. Univ. Studies 17: 36 (prey).
dentatus (Fox). N. Mex., Ariz., Calif.
Gorytes dentatus Fox, 1893. Canad. Ent. 25: 116. ¢.
denticulatus denticulatus (Packard). Widely distributed in North America including Mexico.
Ecology: Nests in sand. Prey: Deltocephalinae sp. nymph. Another subsp. occurs in
Central and South America.
Gorytes denticulatus Packard, 1867. Ent. Soc. Phila., Proc. 6: 430. °.
Gorytes barbatulus Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 408. ¢.

Biology: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 196 (prey, nest).

diversus (Fox). Calif.
Gorytes diversus Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 47: 527. 9, ¢.
floridicus Bohart. Fla.
Hoplisoides floridicus Bohart, 1968. Ent. Soc. Wash., Proc. 70: 289. d, °.
glabratus Bohart. Ariz., N. Mex., Colo., Tex. Ecology: Nests in sandy soil, stores as many as 98
prey per cell. Prey: Aceratagallia uhleri (Van D.), mostly nymphs and a few adults.
Hoplisoides glabratus Bohart, 1968. Ent. Soc. Wash., Proc. 70: 291. 6, °.

Biology: Bohart and Menke, 1976. Sphecid wasps of world, p. 520 (nest, prey, egg).
hamatus (Handlirsch). Western North America. Ecology: Nests, sometimes gregariously, in

soil varying from heavy clay-loam, to sandy clay to sand, makes 1 cell per nest, stores
7-14 prey per cell. Parasite: Senotainia trilineata (Wulp)?, S. rubriventris Macq.?;
Nysson rusticus Cr.?, Epinysson moestus (Cr.), E. pumilis (Cr.)? Prey: Stictocephala?
sp., Ceresini sp. probably Stictocephala sp., Telamonini sp., Membracidae sp.; preys upon
nymphs only.

Gorytes hamatus Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 403. d.

Gorytes spilographus Handlirsch, 1895. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 104: 895.
2

Hoplisoides arizonensis Baker, 1907. Invertebrata Pacifica 1: 164. 2.
Gorytes (Hoplisoides) adornata Bradley, 1920. Amer. Ent. Soc., Trans. 46: 115. 9.
Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 151-152, fig. 40 (larva).

Biology: Powell and Chemsak, 1959. Pan-Pacific Ent. 35: 195-201 (nest, prey transport, egg,
parasites ?). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 46-49 (nest,

Superfamily SPHECOIDEA 1701

prey). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 494 (nest, prey, parasites). —Bohart
and Menke, 1976. Sphecid wasps of world, p. 520 (parasite).
placidus birkmanni Baker. Southwestern U. S.
Hoplisoides Birkmanni Baker, 1907. Invertebrata Pacifica 1: 166. °.
Hoplisoides pruinosus Baker, 1907. Invertebrata Pacifica 1: 166. 9.
placidus nebulosus (Packard). Eastern U.S. Ecology: Nests in sand, makes 1-3 cells per nest,

stores 4-20 prey per cell. Parasite: Senotainia trilineata (Wulp)?; Dasymutilla v. vesta
(Cr.)?; Nysson opulentus Gerst.?, N. tuberculatus Handl., N. daeckei Vier. Prey:
Microcentrus spp., Palonica virida Ball, P. sp., Ceresa palmeri Van D.?, Ceresini spp.,
Campylenchia latipes Say, Enchenopa binotata Say, Entylia concisa Wlkr., E. sinuata
F., Publilia concava Say, Spissistilus festinus Say, Telamona decorata Ball, Tylopelta
brevis Van D., Vanduzea arcuata (Say), V. triguttata Burm.; both adults and nymphs
are used.

Gorytes nebulosus Packard, 1867. Ent. Soc. Phila., Proc. 6: 424. °.

Gorytes armatus Provancher, 1887. Addit. Corr. Faune Ent. Canada Hym., p. 272. “2” =
é.

Gorytes microcephalus Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97:
405. 6.

Gorytes Pergandei Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 407. 3.

Philanthus Harringtonti Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 278.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 45, figs. 43, 44 (larva).

Biology: Pate, 1946. Brooklyn Ent. Soc., Bul. 41: 99 (prey). —Krombein, 1953. Wasmann Jour.
Biol. 10: 286-287 (nest, prey carriage, parasites ?). —Krombein, 1959. Ent. Soc. Wash.,
Proc. 61: 196 (prey, nest). —Evans, 1966. Compar. ethology and evolution of sand wasps,
pp. 34-46, figs. 28-33, 35 (nest, prey, life cycle, cocoon, parasites). —Bohart and Menke,
1976. Sphecid wasps of world, p. 520 (nest, prey).

placidus placidus (Smith). Fla.
Gorytes placidus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 368. 2, ¢.
Gorytes rufipes Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 369. 9.
projectus Bohart. Calif. (San Joaquin Valley).
Hoplisoides projectus Bohart, 1968. Ent. Soc. Wash., Proc. 70: 290. 5, 2.
punctifrons (Cameron). Western U. S.; Mexico (Sinaloa).
Gorytes punctifrons Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 74, pl. 5, figs. 13, 13a.
2

Gorytes gulielmi Viereck, 1907. Amer. Ent. Soc., Trans. 33: 408. 2.

semipunctatus (Taschenberg). Southern U. S.; Argentina. Probably adventive in U.S.
Hoplisus semipunctatus Taschenberg, 1875. Ztschr. Gesell. Naturwiss. Berlin 45: 367. 2.
spilopterus (Handlirsch). Western U.S. Ecology: Nests in damp sand. Prey: Stictocephala
wickhami Van D., Spissistylus festinus (Say), Campylenchia latipes (Say); preys upon
adults and nymphs.
Gorytes spilopterus Handlirsch, 1888. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 97: 414. 9.
Gorytes maculatus Provancher, 1895. Nat. Canad. 22: 140. °.
Gorytes (Hoplisoides) pogonodes Bradley, 1920. Amer. Ent. Soc., Trans. 46: 114. 6.

Biology: Pate, 1946. Brooklyn Ent. Soc., Bul. 41: 99 (prey). —Evans, 1966. Compar. ethology
and evolution of sand wasps, pp. 55-56 (nest, prey).
splendidulus (Bradley). Western U. S.
Gorytes (Hoplisoides) splendidula Bradley, 1920. Amer. Ent. Soc., Trans. 46: 113. 9, ¢.
tricolor (Cresson). Widely distributed in U. S.; Mexico (Sinaloa). Ecology: Nests in
aggregations in sand, makes 2-4 cells per nest, stores 7-19 prey per cell. Parasite:
Nysson bellus Cr.? Prey: Parabolocratus brunneus Ball, adult females and a few
nymphs.
Gorytes tricolor Cresson, 1868. Amer. Ent. Soc., Trans. 1: 380. “2” = 6.
Gorytes (Hoplisus) helianthi Rohwer, 1911. U.S. Natl. Mus., Proc. 40: 569 °.
Hoplisus rufocaudatus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 401. 6.

1702 Hymenoptera in America North of Mexico

Biology: Evans, Lin and Yoshimoto, 1954. Ent. News 65: 5-11 (nest, prey transport, parasite
?).

SUBFAMILY STIZINAE

All species nest in the ground.
Revision: Handlirsch, 1892. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 101: 25-181, 3 pls. (spp.

of world). —Fox, 1896 (1895). Acad. Nat. Sci. Phila., Proc. 47: 266-268 (N. Amer. spp.).
—Parker, 1929. U. S. Natl. Mus., Proc. 75 (5): 7-11, figs. 1-6 (N. Amer. genera).

Genus STIZUS Latreille

Stizus Latreille, 1802-1803. Hist. Nat. Crust. Ins., v. 3, p. 344.
Type-species: Stizus ruficornis Fabricius. Desig. by Blanchard, 1846 (=Larra
ruficornis Fabricius).
Megastizus Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 344.
Type-species: Stizus brevipennis Walsh. Orig. desig.
Stizolarra Saussure, 1887. Soc. Ent. 2: 9.
Type-species: Sphex vespiformis Fabricius. Desig. by Pate, 1937.
Megalostizus Schulz, 1906. Spolia Hym., p. 199. Emend.

This very large genus occurs in the Holarctic and Ethiopian Regions; it is also present in parts
of the Oriental Region, though apparently absent from Southeast Asia. Several species have
been reported to prey upon grasshoppers, katydids, and, rarely, mantids.

Revision: Dow, 1941. Psyche 48: 171-181, 7 figs. (N. Amer. spp.).
brevipennis Walsh. U. S. east of 100th meridian. Prey: Conocephalus sp. adult.

Stizus brevipennis Walsh, 1869. Amer. Ent. 1: 162. ¢.
Larra Brendeli Taschenberg, 1875. Ztschr. Gesam. Naturw. Halle 45: 361. ¢.

Biology: Williams, 1914 (1913). Kans. Univ. Sci. Bul. 8: 198 (prey).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 19, figs. A-G (male
genitalia).
iridis Dow. Utah, Calif. Ecology: Makes a unicellular cell in talus slope, stores at least 8 prey
per cell. Prey: Trimerotropis pallidipennis (Burm.), T. sparsa (Thom.); adults.
Stizus iridis Dow, 1942 (1941). Psyche 48: 171. 6.

occidentalis Parker. Calif., Ariz.
Stizus occidentalis Parker, 1929. U. S. Natl. Mus., Proce. 75 (5): 9. 2, 6.

texanus Cresson. Tex., Ariz.; Mexico (Chihuahua).
Stizus texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 222. 9, d.

Genus STIZOIDES Guerin
Revision: Gillaspy, 1963. Mus. Compar. Zool., Bul. 128: 369-391, 1 pl. (N. Amer. spp.).

Genus STIZOIDES Subgenus STIZOIDES Guerin

Stizus subg. Stizoides Guerin, 1844. Iconogr. Regn. Anim. 7, Ins., p. 438.
Type-species: Larra fasciata Fabricius. Desig. by Parker, 1929.

The typical subgenus does not occur in North America.
Genus STIZOIDES Subgenus TACHYSTIZUS Pate

Tachystizus Pate, 1937. Amer. Ent. Soc., Mem. 9: 638.
Type-species: Crabro tridentatus Fabricius. Orig. desig.
Two species of this subgenus are known to be cleptoparasites of other sphecoid wasps.
foxi Gillaspy. Ariz.; Mexico (Baja California).
Stizoides (Tachystizus) foxi Gillaspy, 1963. Mus. Compar. Zool., Bul. 128: 378, figs. 1, 2, 5, 7,
Se ONCr

Superfamily SPHECOIDEA 1703

renicinctus (Say). Mich., Wis., Ill. south to Tex., west to Alta., B. C. and Calif., D. C., N. C.;
Mexico (Zacatecas). Host: Prionyx atratus (lep.), P. thomae (F.).
Stizus renicinctus Say, 1823. West. Quart. Rptr. 2: 77.
Stizus unicinctus Say, 1824. Amer. Ent. 1: 4. 6. Emend.

Biology: Bradley, 1908. Ent. Soc. Amer., Ann. 1: 129 (sleeping aggregation). — Williams, 1914
(1913). Kans. Univ. Sci. Bul. 8: 230 (host). —Smith, 1915. U.S. Dept. Agr., Bul. 293: 10-11
(host). —Rau and Rau, 1918. Wasp studies afield, pp. 180-193 (host). —Evans, 1966.
Compar. ethology and evolution of sand wasps, p. 129 (host).

Genus BEMBECINUS Costa

Bembecinus Costa, 1859. Fauna Regn. Napoli, Imen. Acul., Nyssonid., p. 4.
Type-species: Bembecinus meridionalis Costa. Monotypic.

Stizomorphus Costa, 1859. Fauna Regn. Napoli, Imen. Acul., Nyssonid., p. 7.
Type-species: Vespa tridens Fabricius. Monotypic.

Gorystizus Pate, 1937. Amer. Ent. Soc., Mem. 9: 29.
Type-species: Vespa tridens Fabricius. Orig. desig.

Lavia Rayment, 1953. Victorian Nat. 70: 123. Nom. nudum.

The genus occurs in all major zoogeographic regions and on many islands. Five species groups
are recognized but the few North American taxa are all assigned to the Tridens Group. Many
species nest in sandy soil in compact colonies, frequently made up of many individuals. All spe-
cies whose biology is known prey upon leafhoppers and some other Homoptera, and practice
progressive provisioning.

Revision: Krombein and Willink, 1951 (1950). Amer. Midland Nat. 44: 699-713 (N. Amer.

spp.).

Biology: Evans, 1955. Behaviour 7: 295-302 (comparative behavior).
bishoppi Krombein and Willink. Tex.

Bembecinus bishoppi Krombein and Willink, 1951 (1950). Amer. Midland Nat. 44: 710. 9,
3

moneduloides (Smith). Fla. (St. Johns Bluff).
Larra moneduloides Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 346. 2, (6 misdet.).

Taxonomy: Willink, 1957. Ann. and Mag. Nat. Hist. (12) 9: 702-704.
nanus floridanus Krombein and Willink. S. Fla.
Bembecinus nanus floridanus Krombein and Willink, 1951 (1950). Amer. Midland Nat. 44:
706. 3.
nanus nanus (Handlirsch). N. J. to north. Fla., west to Tex., Iowa, east. Nebr. Prey:
Graphocephala versula (Say), Fulgoridae spp.
Stizus nanus Handlirsch, 1892. Akad. Wiss. Wien, Math.-Nat. KI., Sitzber. 101: 61. 9, d.

Biology: Shappirio, 1946. Ent. News 57: 229-230 (prey).
nanus strenuus (Mickel). Nebr., S. Dak., Wyo., Tex.
Stizus strenuus Mickel, 1918 (1917). Nebr. Univ. Studies 17: 331. 9, 6.
neglectus (Cresson). Nebr., Kans., Tex., La., Miss. Ecology: Colony nests in hard-packed sand
containing pebbles, makes 1-2 cells per nest, stores 10-15 prey per cell, practices
progressive provisioning. Prey: Gyponana octolineata (Say), Texananus excultus (Uhl.),
Xerophloea majesta Laws., all adults; Scolops sp. nymphs; preys principally upon adult
Cicadellidae.
Monedula neglecta Cresson, 1872. Amer. Ent. Soc., Trans. 4: 222. 2.
Stizus xanthochrous Handlirsch, 1892. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 101: 69.
é.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 47, figs. 37-42 (larva).

Biology: Evans, 1955. Behaviour 7: 287-295, 4 figs. (mating, nest, prey, egg, cocoon, life
cycle).

1704 Hymenoptera in America North of Mexico

quinquespinosus (Say). Nebr., Colo. and south. Calif. to Panama. Ecology: Nests in large

colonies in sand, makes unicellular nest. Prey: Ciminius hartii (Ball), Stirellus bicolor
(Van D.), Exitianus sp., Cuerna lateralis (F.), C. sp., Draeculacephala noveboracensis
(Fitch), Carneocephala sagittifera (Uhl.), Acinopterus angulatus Laws., Cicadellidae sp.;
adults and a few nymphs.

Nysson 5-spinosus Say, 1823. West. Quart. Rptr. 2: 78.

Stizus godmani Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, pl. 5, fig. 8. 2, d.

Stizus lineatus Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 102. ¢.

Stizus flavus Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 103. ¢.

Stizus flavus var. subalpinus Cockerell, 1898. Davenport Acad. Nat. Sci., Proc. 7: 142. 3.

Nysson cressoni Cameron, 1904. Amer. Ent. Soc., Trans. 30: 95. 9.

Taxonomy: Willink, 1957. Ann. and Mag. Nat. Hist. (12) 9: 702-704. —Evans, 1959. Amer.
Ent. Soc., Trans. 85: 152-153, figs. 71-73 (larva).

Biology: Rohwer, 1909. Colo. Univ. Studies 6: 246-247 (colony; prey misdet.). —Rodeck, 1931.
Colo.-Wyo. Acad. Sci., Jour. 1: 61 (prey, nest). —Strandtmann, 1945. Ent. Soc. Amer., Ann.
38: 312-313 (nest, prey transport). —Evans, 1966. Compar. ethology and evolution of sand
wasps, pp. 134-141, fig. 76 (nest, prey). —Evans, 1968. Ent. Soc. Amer., Ann. 61: 1344
(prey).

wheeleri Krombein and Willink. Ariz.
Bembecinus wheeleri Krombein and Willink, 1951 (1950). Amer. Midland Nat. 44: 709. 9°,
é.

SUBFAMILY BEMBICINAE

All species nest in the ground, usually in quite friable soil. The more primitive species are soli-
tary nesters, practice mass provisioning and prey upon Hemimetabola, whereas the more spe-
cialized species nest in large aggregations, have developed progressive provisioning behavior in-
dependently in some genera and prey largely upon advanced Holometabola.

This subfamily has undergone considerable evolution in the New World where a number of
genera occur. Bembix is the only genus of worldwide distribution and it occurs in all major
zoogeographic regions.

Revision: Parker, 1917. U. S. Natl. Mus., Proc. 52: 1-155. —Parker, 1929. U. S. Natl. Mus.
Proc. 75 (5): 11-181.

Taxonomy: Bohart and Horning, 1971. Calif. Ins. Survey, Bul. 13: 1-49, 74 figs., 18 maps
(Calif. spp.).

Genus BICYRTES Lepeletier

Bicyrtes Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 53.
Type-species: Bicyrtes Servillii Lepeletier. Monotypic.
Bembidula Burmeister, 1874. Acad. Nac. Cien. Cordoba, Bol. 1: 122.
Type-species: Monedula discisa Taschenberg. Desig. by Parker, 1917.
Dumonela Reed, 1894. Univ. Chile, An. 85: 608.
Type-species: Monedula sericea Spinola. Orig. desig.

Wasps of this genus make uni- or multicellular nests, usually in fine- to coarse-grained sand.
They prey upon Heteroptera, chiefly Pentatomidae and Coreidae, although Pyrrhocoridae, Scu-
telleridae, Cydnidae, Lygaeidae and Reduviidae may be used occasionally; nymphs are preferred
to adults. The egg is laid on the first bug placed in the cell.

affinis (Cameron). South. Ariz.; Mexico (Guerrero).
Bembidula affinis Cameron, 1897. Ann. and Mag. Nat. Hist. (6) 19: 371. 2.
capnoptera (Handlirsch). Ga., Fla., La., Tex., Kans., N. Mex., Ariz., Calif.; Mexico (Baja
California). Ecology: Nests in soil, stores 5-7 prey per cell. Prey: Thyanta pallidovirens
accerra (McAtee) nymphs and adults.
Bembidula capnoptera Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98:
ASTOR se
Bembidula capnoptera var. mesillensis Cockerell, 1898. Davenport Acad. Nat. Sci., Proe. 7:
142. 3.

Superfamily SPHECOIDEA 1705

Bicyrtes annulata Parker, 1917. U.S. Natl. Mus., Proc. 52: 67. 6, °.
Bicyrtes tristis Fox, 1923. Calif. Acad. Sci., Proc. (4) 12: 435. d.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 157-158, fig. 5 (sleeping aggregation).
—Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 171-172 (nest, prey).
fodiens (Handlirsch). Southeast. to southwest. U. S.; Mexico, Central America. Ecology: Nests

in fine sand or sandy clay, makes 4-5 cells per completed nest and stores 10-23 prey per
cell. Parasite: Metopia argyrocephala (Meig.); Holopyga ventralis (Say). Prey: Solubea
pugnax F. adults, Mormidea lugens (F.) adults; Homaemus aeneifrons Say nymphs.
Bembidula fodiens Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98: 499. 6,
OF
Bembidula Burmeisteri Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98:
500. °.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 50, figs. 53-55 (larva).

Biology: Rau, 1922. St. Louis Acad. Sci., Trans. 24 (7): 28 (nest). —Evans, 1966. Compar.
ethology and evolution of sand wasps, pp. 167-170, 173 (nest, prey transport, egg, cocoon,
life cycle, parasites).

insidiatrix (Handlirsch). Mass., N. J., Fla., Ky., Tex., N. Mex.

Bembidula insidiatrix Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98: 494.

Ores
quadrifasciata (Say). U.S. east of Rocky Mts. in Transit. and Austr. Zones. Ecology: Nests in

sand or heavier soil, usually makes a unicellular nest but occasionally one with 2-3 cells,
stores 4-14 prey per cell. Parasite: Senotainia rubriventris Macq., S. trilineata (Wulp),
S. vigilans Allen, S. sp.; Holopyga ventralis (Say). Prey: Aplomerus sp., Zelus sp.;
Lygaeidae sp.; Acanthocephala femorata (F.), Anasa tristis (DeG.), Archimerus
alternatus (Say), A. calcarator (F.), Chariestrus antennator (F.), Leptoglossus clypealis
Heid., L. oppositus Say, L. phyllopus (L.); Acrosternum hilare (Say), A. sp., Banasa
dimidiata (Say), Brochymena arborea Say, B. cariosa Stal, B. carolinensis Westw., B.
quadripustulata (F.), Chlorochroa uhleri Stal, Dendrocoris humeralis Uhl., Edessa
florida Barb., Euschistus tristigmus Say, E. spp., Menecles incertus Say, Murgantia
histrionica Hahn, Nezara viridula (L.), N. sp., Podisus sp., Stethaulax marmoratus Say,
Thyanta custator F., T. sp.; Homaemus aeneifrons Say, Tetyra bipunctata H.-S.;
Cyrtomenus sp.; Pentatomidae and Coreidae are preferred prey and only nymphs are
used.

Monedula quadrifasciata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app., p.
336. 3, .

Monedula sallei Guerin, 1844. Iconogr. Regne Anim., v. 7, p. 437.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 49, figs. 45-52 (larva). —Evans,
1964. Amer. Ent. Soc., Trans. 90: 265 (larva).

Biology: Hartman, 1905. Tex. Univ., Bul. 65: 32-36, fig. 14 (nest, prey, parasites). —Parker,
1917. U. S. Natl. Mus., Proc. 52: 134 (nest, prey). —Rau and Rau, 1918. Wasp studies
afield, pp. 41-43 (nest, prey). —Smith, 1923. Ent. Soc. Amer., Ann. 16: 238-246 (nest, prey,
parasite). —Davis, 1926. N. Y. Ent. Soc., Jour. 34: 89-90 (nest, prey). —Rau, 1934. Canad.
Ent. 66: 260 (prey transport). —Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 287-288
(nest, prey). —Krombein, 1955. Ent. Soc. Wash., Proc. 57: 152-157, 4 figs. (nest, prey
transport, egg, cocoon, life cycle, parasite). —Krombein, 1958. Ent. Soc. Wash., Proce. 60:
105-106 (nest, prey). —Krombein, 1959. Ent. Soc. Wash., Proc. 61: 196 (nest, prey).
—Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 146-160, figs. 80-90
(nest, mating, prey transport, life cycle, egg, cocoon, parasites). —Kurezewski and
Kurezewski, 1971. Kans. Ent. Soc., Jour. 44: 336 (prey).

variegata (Olivier). Tex.; Mexico to Argentina, West Indies.
Bembex variegata Olivier, 1789. Encycl. Meth., Ins., v. 4, p. 292.
Monedula sericea Spinola, 1851. In Gay, Hist. Fis. Pol. Chile, Zool., v. 6, p. 315.
Bembex guiana Cameron, 1912. Timehri (3) 2: 431. 9.

1706 Hymenoptera in America North of Mexico

ventralis (Say). South. Canada, U. S.; north. Mexico. Ecology: Nests in sand or sandy gravel,

sometimes in small aggregations, usually makes a unicellular cell but occasionally one
with 2-3 cells, stores 3-18 prey per cell. Parasite: Senotainia vigilans Allen, S. trilineata
(Wulp). Prey: Anasa tristis (DeG.); Banasa dimidiata Say, Cosmopepla bimaculata
Thom., Elasmostethus cruciatus Say, Euschistus euschistoides Voll., E. tristigmus Say,
E. variolarius Beauv., Menecles incertus Say, Mormidia lugens F., Thyanta
pallidovirens accerra McAtee, Trichopepla semivittata Say; only nymphs are used and
Pentatomidae are the preferred prey.

Monedula ventralis Say, 1824. In Keating, Narr. Long’s 2nd Exped. v. 2, app., p. 337. 6.

Bicyrtes Servillii Lepeletier, 1845. Hist. Nat. Ins., Hym., v. 3, p. 53. 2.

Monedula parata Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 416. @.

Bembidula meliloti Rohwer, 1908. Ent. News 19: 376. d.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 50, figs. 56-58 (larva).

Biology: Hartman, 1905. Tex. Univ., Bul. 65: 36-39 (nest, prey). —Parker, 1917. U.S. Natl.
Mus., Proc. 52: 132-133 (nest, prey, life cycle). —Evans, 1966. Compar. ethology and
evolution of sand wasps, pp. 160-167, figs. 91-93 (nest, prey transport, egg, life cycle).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 19, figs. I-L (male
genitalia).
viduata (Handlirsch). Utah, Ariz., Tex.; north. Mexico. Ecology: Nests in coarse gravel. Prey:

Coreidae nymphs.
Bembidula viduata Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. KI., Sitzber. 98: 491. 9.

Bicyrtes gracilis Parker, 1917. U.S. Natl. Mus., Proc. 52: 68. 3.
Biology: Alcock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 160 (nest, prey).

Genus MICROBEMBEX Patton

Microbembex Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 364.
Type-species: Bembex monodonta Say. Orig. desig.

The North American species are scavengers, bringing in as food for their larvae a wide
variety of dead arthropods. Two South American species have been reported as paralyzing live
beetles and other insects, but these observations need confirmation. The egg is laid in the empty
cell and progressive provisioning is practiced.

argyropleura Bohart. South. Calif., Ariz., Nev., Utah; Mexico (Baja California). Ecology: Nests
in dense aggregations in sand dune blowouts, lays egg in empty cell, practices
progressive provisioning. Parasite: Senotainia sp. near rufiventris (Coq.)? Prey: Dead
arthropods including Diptera, Hymenoptera, Coleoptera, Heteroptera, Araneae.
Microbembex argyropleura Bohart, 1970. Pan-Pacifie Ent. 46: 203. 6, 9.

Biology: Alcock, 1975. Southwest. Nat. 20: 337-339, figs. 1, 2 (nest, prey, parasite).

aurata Parker. West. Tex. to south. Calif.
Microbembex aurata Parker, 1917. U. S. Natl. Mus., Proc. 52: 121. d, 2.

Taxonomy: Gillaspy, 1963. Proc. Ent. Soc. Wash. 65: 230.
californica Bohart. Cent. Calif. to Wyo., Utah and N. Mex.; Mexico (Baja California). Prey:

Dead arthropods.
Microbembex californica Bohart, 1970. Pan-Pacific Ent. 46: 202. d, 9.

Biology: Goodman, 1970. Pan-Pacific Ent. 46: 207-209 (nest, “prey”).
hirsuta Parker. N. Mex., Tex.
Microbembex hirsuta Parker, 1917. U.S. Natl. Mus., Proc. 52: 122. 6, °.

monodonta (Say). South. Canada and U. S. east of Rocky Mts.; Mexico, Central America.
Ecology: Nests in large aggregations in loose dry to moist firm sand, makes a
unicellular nest, places egg upright in empty cell, practices progressive provisioning.
Parasite: Senotainia trilineata (Wulp), S. rufiventris (Coq.)?; Exoprosopa fascipennis
(Say); Dasymutilla bioculata (Cr.), D. sp.; Parnopes f. fulvicornis Cam.?, P. f. atlanticus
Krom.? Prey: Dead arthropods belonging to Phalangida, Araneida, Ephemeroptera,

Superfamily SPHECOIDEA 1707

Orthoptera, Psocoptera, Hemiptera, Neuroptera, Trichoptera, Lepidoptera, Coleoptera,
Diptera, Hymenoptera; live paralyzed prey of other wasps may be stolen. Predator:
Small red ants.
Bembex monodonta Say, 1824. In Keating, Narr. Long’s 2nd. Exped., v. 2, app., p. 335. ¢.
Microbembex monodonta occidentalis Johnson and Rohwer, 1908. Ent. News 19: 375.
Microbembex tarsalis Rohwer, 1914. U.S. Natl. Mus., Proc. 47: 516. d, @.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 55, figs. 79-84 (larva). —Evans,
1964. Amer. Ent. Soc., Trans. 90: 266 (larva).

Biology: Hartman, 1905. Tex. Univ., Bul. 65: 21-26 (nest, prey). —Parker, 1917. U.S. Natl.
Mus., Proc. 52: 134-141 (mating, nest, prey, life cycle, cocoon, parasite, predator). —Stoehr,
1917. Nat. Canad. 43: 113-119 (mating, nest, prey). —Rau and Rau, 1918. Wasp studies
afield, pp. 39-41, fig. 7 (nest, prey). —Mickel, 1924. Ent. News 35: 236-242 (parasite).
—Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 288 (prey, life cycle). —Krombein, 1958.
Amer. Ent. Soc., Trans. 84: 166 (parasite ?). —Evans, 1966. Compar. ethology and
evolution of sand wasps, pp. 361-388, figs. 177-190 (sleeping burrows, nest, prey, egg, life
cycle, parasites ?).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 19, figs. H, Q, R, pl. 20,
figs. A, B (male genitalia).
nigrifrons (Provancher). U.S. and Mexico west of 100th meridian. Ecology: Nests in sand.

Parasite: Dasymutilla cassandra Mick., D. gloriosa (Sauss.); Parnopes f. fulvicornis
Cam. Prey: Many kinds of dead arthropods.

Bembex nigrifrons Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 415. 6, °.

Microbembex monodonta neomexicana Johnson and Rohwer, 1908. Ent. News 19: 375.

Microbembex monodonta deltaensis Johnson and Rohwer, 1908. Ent. News 19: 375. 6.

Biology: Bohart and MacSwain, 1940. Pan-Pacifie Ent. 16: 92-93 (parasite; misdet. as awrata
Prkr.). —Alcock and Ryan, 1973. Pan-Pacific Ent. 49: 144-148, 1 fig. (prey, mating
behavior). —Aleock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 161-162 (nesting
behavior).

rufiventris Bohart. Calif.
Microbembex rufiventris Bohart, 1970. Pan-Pacific Ent. 46: 204. dg, 2.

Genus STICTIA Illiger

Monedula Latreille, 1802-1803. Hist. Nat. Crust. Ins., v. 3, p. 345. Preoce.
Type-species: Vespa signata Linnaeus. Desig. by Latreille, 1810.
Stictia Illiger, 1807. Jn Rossi, Fauna Etrusca, Ed. 2, v. 2, p. 131. N. name for Monedula
Latreille.

These wasps occur only in the New World where most species are restricted to the tropical
areas. They occasionally nest in large aggregations in sandy soil. The nest is unicellular. They
practice progressive provisioning and prefer horseflies (Tabanidae) as prey. The egg is placed in
the empty cell in carolina (F.) and on the first fly brought into the cell in our other species.

carolina (Fabricius). N. J. and Pa. south to Fla., west to Ill., Kans., and N. Mex. Ecology:
Occasionally nests in very large aggregations in sand; the egg is placed in the empty
cell; as many as 63 prey may be supplied. Parasite: Senotainia trilineata
(Wulp), Miltogrammini spp.; Dohrniphora cornuta (Big.). Prey: Psorophora ciliata (F.);
Chrysops dimmocki Hine, Chlorotabanus crepuscularis Beq., Hybomitra hinei wrighti
Whit., Tabanus abdominalis F., T. americanus Forst., T. atratus F., T. a. var.
nantuckensis Hine, T. bishoppi Stone, T. cheliopterus Rond., T. coarctatus Stone, T.
endymion O.S8., T. fumipennis Wied., T. i. imitans Wlkr., T. lineola F., T. melanocerus
lacustris Stone, T. mularis Stone, T. nigripes Wied., T. nigrescens Beauv., T. petiolatus
Hine, T. sparus Whit., T. stygius Say, T. sulcifrons Macgq., T. trijunctus Wlkr., T. vittiger
schwardti Phil., T. spp.; Odontomyia cincta Oliv.; Volucella mexicana Macq,., V. nigra
Greene; Graphomya maculata Scop., Musca domestica L., Orthellia caesarion Meig.,
Stomoxys calcitrans L.; Amobia erythrura (Wulp), Sarcophaga bullata Prkr., S.
ochracea Ald., S. sarracenioides Ald., S. spp.; Calliphora vomitoria L., Cochliomyia

1708 Hymenoptera in America North of Mexico

macellaria F.; Tachinidae spp.; Melampsalta calliope (Wlkr.); Atalopedes campestris
Boisd.; the last two prey species (cicada and skipper) were stored only in a large
aggregation where there was considerable competition for prey. Predator: Solenopsis
geminata F. This wasp is commonly called the “horse guard” because it preys so
commonly on horseflies.

Bembex carolina Fabricius, 1793. Ent. System., v. 2, p. 249.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 53, figs. 72-78 (larva). —Evans,
1964. Amer. Ent. Soe., Trans. 90: 268, fig. 55 (larva).

Biology: Bryant, 1870. Amer. Ent. 2: 87 (prey capture). —Ashmead, 1894. Psyche 7: 61 (prey
capture). —Hartman, 1905. Tex. Univ., Bul. 65: 27-29, figs. 7, 15 (nest, prey capture, egg,
life cycle, cocoon). —Hine, 1906. La. State Crop Pest Comn. Cir. 6: 20-27 (nest, prey).
—Hine, 1907. La. Agr. Expt. Sta., Bul. 93: 18-15 (nest, prey). —Krombein, 1958. Ent. Soc.
Wash., Proc. 60: 106-107 (nest, prey, parasite). —Krombein, 1959. Ent. Soc. Wash., Proc. 61:
196-197 (nest, prey, life cycle, cocoon). —Evans, 1966. Compar. ethology and evolution of
sand wasps, pp. 223-243, figs. 119-133 (mating, nest, egg, prey capture and transport, life
cycle, parasites). —Lin, 1971. Tex. Jour. Sci. 23: 275-283, 3 figs. (mating, nest, egg, prey,
life cycle, cocoon).

signata signata (Linnaeus). South. Fla., south. Calif; West Indies, tropical Mexico, Central and

South America. Ecology: Nests in large aggregations in sand along coasts and rivers;
the egg is placed on the first fly brought into the cell. Parasite: Villa sp.? Prey:
Tabanus truquii Bell., T. sp., Diachlorus curvipes (F.), Lepiselaga crassipes F.,
Chrysops costatus F.; Hedriodiscus dorsalis (F.); Allograpta sp., Eristalis sp., Volucella
sp.; Musca domestica L., Morellia scapulata (Big.); Cochliomyia macellaria (F.),
Phaenicia cluvia (Wlkr.); Sitophaga sp., Tachinidae sp.; Sarcophaga spp.; Tabanidae are
the preferred prey. Another subsp. occurs in Peru.

Vespa signata Linnaeus, 1758. Syst. Nat., Ed. 10, v. 1, p. 574.

Apis vespiformis DeGeer, 1773. Mem. Hist. Ins., v. 3, p. 570, pl. 28, figs. 3, 4.

Monedula insularis Dahlbom, 1845. Hym. Europaea, v. 1, pp. 186, 494. °.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 243-247 (nest, prey,
egg, scavengers).
vivida (Handlirsch). Tex. (Cameron Co.); Mexico (Tamaulipas, Veracruz, Yucatan). Ecology:
Nests in hard-packed beach sand in small aggregations; the egg is placed on the first fly
brought into the cell. Parasite: Miltogrammini sp. Prey: Tabanus texanus Hine, T. s.
schwarati Phil., T. vittiger guatemalensis Hine, T. spp., Leucotabanus itzarum Beq.;
Cochliomyia macellaria (F.); Sarcophaga effrenata Wlkr.
Monedula vivida Handlirsch, 1890. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 99: 101. °.

Taxonomy: Evans, 1957. Ent. News 68: 77. ¢. —Evans, 1959. Amer. Ent. Soe., Trans. 85: 153
(larva). —Evans, 1964. Amer. Ent. Soc., Trans. 90: 268-269 (larva).

Biology: Evans, 1957. Ent. News 68: 76-77 (nest, prey transport). —Evans, 1966. Compar.
ethology and evolution of sand wasps, pp. 247-252, figs. 134-137 (male territoriality, nest,
prey, egg, parasite).

Genus BEMBIX Fabricius

Bembix Fabricius, 1775. Systema Ent., Char. Gen., p. xxili. No species.
Type-species: Bembex rostrata of Fabricius. Desig. by Latreille, 1810.
Bembyx Fabricius, 1775. Systema Ent., p. 361. Lapsus.
Bembex Fabricius, 1776. Gen. Ins., p. 122. Emend.
Apobembex Pate, 1937. Amer. Ent. Soc., Mem. 9: 9.
Type-species: Bembex oculata of Latreille. Orig. desig.
Epibembex Pate, 1937. Amer. Ent. Soc., Mem. 9: 26.
Type-species: Apis rostrata Linnaeus. Orig. desig.

This large genus occurs in all major zoogeographic regions and is the only representative of
the subfamily in the Old World. These wasps nest in a variety of soils ranging from loose sand
subject to blowing to hard-packed soil, and almost all species practice progressive provisioning.

Superfamily SPHECOIDEA 1709

The more primitive species lay the egg on the first prey brought into the cell and the most ad-
vanced species place the egg in the empty cell. The nest may have a simple unicellular structure
or it may contain up to 5 cells; accessory burrows are made by some of the more advanced spe-
cles.

Taxonomy: Evans and Matthews, 1968. Ent. Soe. Amer., Ann. 61: 1284-1299, 26 figs.
(synopsis of N. Amer. spp., keys, characters of species groups).

Biology: Evans, 1957. Studies on compar. ethology Bembix, 248 pp., 52 figs., 29 tables.
SPECIES GROUP BELFRAGEI

These primitive species nest in a variety of soil types, make multicellular nests, and place the
egg on the first fly brought into the cell.

belfragei Cresson. South cent. U.S. Ecology: Nests in large aggregations in coarse to fine

sand, makes a 2-celled nest with each cell at the end of a branch off the main burrow,
each with an accessory branch, provides 20-28 prey per cell. Parasite: Senotainia sp. in
trilineata (Wulp) complex; Dasymutilla pyrrhus (Fox)? Prey: Tabanidae spp.;
Systoechus vulgaris Lw.; Eristalis agrorum F., E. latifrons Lw., E. tenax L., Volucella
fasciata Macq.; Orthellia caesarion Meig., Stomoxys calcitrans L.; Cochliomyia
macellaria (F.), Phaenicia caeruleiviridis Macq., Phormia regina Meig.; Sarcophaga
bullata Prkr., S. combicis Tns., S. lherminieri R.-Desv., S. prohibita Ald., S. querula
Wilkr., S. uncata Wulp, S. ventricosa Wulp, S. spp.; Acroglossa hesperidarum Will.,
Archytas apicifera Wlkr., A. aterrima R.-Desv., Belvosia semiflava Ald., Biomya
neomexicana Tns.?, Bonnetia comta Fall., Ceracia dentata Coq., Copecrypta nitens
Wied., Euphorocera floridensis Tns., Gonia sequax Will., Microphthalma disjuncta
Wied., Peleteria sp., Phorocera claripennis Macq., Prosenoides sp., Sturmia sp.,
Winthemia quadripustulata F., Tachinidae sp.

Bembex(!) Belfragei Cresson, 1873. Amer. Ent. Soc., Trans. 4: 220. d.

Bembex(!) cressonis Handlirsch, 1893. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 102: 792.
Owner.

Bembex(!) insignis Handlirsch, 1893. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 102: 793.

ee
Bembix cressonii Dalla Torre, 1897. Cat. Hym., v. 8, p. 503. Lapsus.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 61, figs. 98-101 (larva).

Biology: Evans, 1957. Studies on compar. ethology Bembix, pp. 59-78, figs. 13-18 (mating,
nest, prey, egg, life cycle). —Evans, 1966. Compar. ethology and evolution of sand wasps,
pp. 315-317, fig. 161 (nest, prey, parasite).

frommeri Bohart. Calif. (Inyo Co.).
Bembix frommeri Bohart, 1970. Pan-Pacifie Ent. 46: 201. ¢.
gillaspyi Evans and Matthews. South. Calif. deserts.
Bembizx gillaspyi Evans and Matthews, 1968. Ent. Soc. Amer., Ann. 61: 1290, figs. 8-9. ¢,
De

rugosa Parker. Ariz.
Bembix rugosa Parker, 1917. U. S. Natl. Mus., Proc. 52: 95. 9.

stenobdoma Parker. West. Tex. to south. Calif.

Bembix stenebdoma(!) Parker, 1917. U.S. Natl. Mus., Proc. 52: 79. In key.
Bembix stenobdoma Parker, 1917. U. S. Natl. Mus., Proc. 52: 94. 6.

U-scripta Fox. Tex. to south. Calif.; Mexico, Guerrero and Morelos to Baja California. Ecology:
Nests occasionally in large aggregations in soil varying from loose sand to coarse,
compact sandy gravel, makes 2-5 cells per nest, places egg on first prey brought into
nest, hunts prey and stores nest only at dusk, provides up to 40 prey per cell. Prey:
Hermetia aurata Bell; Tabanus texanus Hine; Chromolepida pruinosa (Coq.); Apiocera
haruspex O.S.; Efferia sp., Erax cressoni Hine, E. tuberculatus Coq., Psilocurus
nudisculus Lw., P. modestus Will., P. puellus Brom., Saropogon sp., Stenopogon ebyi
Brom., Asilidae spp.; Aphoebantus sp. near hirsutus Coq., A. spp., Desmatoneura
argentifrons Will., Lordotus g. gibbus Lw., L. g. striatus Paint., Phthiria sulphurea Lw.,
P. sp., Poecilanthrax lucifera F., Villa flavipilosa Cole, V. parvicornis Lw., V.

1710 Hymenoptera in America North of Mexico

salebrosus Paint., V. sp.; Volucella fraudulenta Will., V. mexicana Macq., V. unipuncta
Curr., V. spp., Syrphidae spp.; Acrosticta mexicana Cole, A. sp.; Limnophora sp.,
Mydaea sp., Phyllogaster cordyluroides Stein; Sarcophaga johnsoni Ald., S. spp.,
Senotainia kansensis Tns.; Archytas marmoratus Tns., Chaetogaedia sp. near analis
Wulp, Goniochaeta plagioides Tns., Olenochaeta kansensis Tns., Phorocera
tachinomoides Tns., P. sp., Promasiphya confusa Ald., Ptilodexia sp., Tachinophyto sp.
near vanderwulpi Tns., Xenoppia monela Reinh.

Bembizx dentilabris Handlirsch, 1893. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 102: 794.
9. A nomen oblitum.

Bembex(!) U-scripta Fox, 1895. Acad. Nat. Sci. Phila., Proc., p. 362. ¢, 2. The valid name
for this taxon chosen by the first revisers.

Bembiz arcuata Parker, 1917. U. S. Natl. Mus., Proc. 52: 81. 6, 9°.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 155, figs. 41, 63 (larva). —Evans, 1961
(1960). Psyche 67: 45-61 (possible syn. of arcuata).

Biology: Evans, 1957. Studies on compar. ethology Bembix, pp. 104-108, figs. 27-28 (nest,
prey). —Evans, 1961 (1960). Psyche 67: 45-61, 2 figs. (male sun dance, mating, nest, prey,
egg, life cycle). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 317-321,
figs. 162-164 (nest, prey).

SPECIES GROUP AMOENA

Members of this group make simple nests with one or more accessory (false) burrows, place
the egg on the first prey brought into the cell, and do not level the mound of excavated earth at
the nest entrance.

amoena Handlirsch. West. Canada and U.S. Ecology: Nests in colonies in soil varying from
sandy loam to coarse sandy gravel, makes 1- or 2-celled nest provided with 1-2
accessory burrows, probably stores more than 30 large prey per cell. Parasite:
Dasymutilla creusa var. bellona (Cr.); Parnopes edwardsii Cr. ?; Physocephala texana
Will. ?; Miltogrammini sp.; Exoprosopa dorcadion O.8., Villa melasoma (Wulp). Prey:
Anoplodonta nigrirostris Lw., Hedriodiscus varipes Lw.; Hybomitra captonis Mart., H.
fulvilateralis Macq., H. liorhina Phil., H. opaca Coq., H. osburni Hine, H. phaenops O.
S.; Poecilanthrax sackeni Coq., Systoechus fumipennis Paint., Villa alternata Say, V.
eumenes (O.S.), V. fulviana nigricauda Lw., V. harveyi Hine, V. lateralis Say, V.
sinuosa jaennickeana O.S.; Thereva sp.; Asilus sp., Cyrtopogon glarealis Mel.,
Promachus sp.; Chrysotoxum ypsilon Will., Eristalis anthophorinus Fall., E. barda Say,
E. latifrons Lw., Eupeodes volucris 0. S., Helophilus hybridus Lw., Metasyrphus
lapponicus Zett., M. meadii Jones, Scaeva pyrastri L., Stenosyrphus pullulus Snow,
Syrphus jonesi Fluke, S. opinator O.S., S. ribesii L., S. vitripennis Meig., Volucella
esuriens (F.); Tetanocera vicina Meig.; Helina punctata R.-Desv., Limnophora
magnipunctata Mall., Lispe brevipes Ald., Mydaea persimilis Mall., Phaonia monticola
Mall., Pyrellia cyanicolor Zett.; Calliphora vicina R.-Desv., C. vomitoria L.,
Cynomyopsis cadaverina R.-Desv., Eucalliphora lilaea Wlkr., Lucilia illustris Meig.,
Melanodexiopsis sp., Phormia regina Meig., Protophormia terraenovae R.-Desv.;
Sarcofahrtia montanensis Prkr., Sarcophaga spp., Macronychia sp.; Acroglossa
hesperidarum Will., Arctophyto sp., Argenteopalpus signiferus Wlkr., Bonellimyia
subpolita Brks., Eumegaparia flaveola Coq., Fabriciella brevirostris Tot., F. rostrata
Tot., Gonia porca Will., G. spp., Melanodexia sp., Melinocera flavicornis Br., Mericia
alberta Curr., M. ampela Wlkr., M. arcuata Tot., M. bicarina Tot., Microphthalma
disjuncta Wied., Paramuscopterysx sp., Peleteria iterans Wlkr., P. neotexensis Brks.,
Ptilodexia sp., Rhachogaster algens Wied., Siphosturmiopsis sp., Spallanzania sp.,
Trochilodes skinneri Coq.

Bembex(!) amoena Handlirsch, 1893. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 102: 769.

Cpe

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 270-271, fig. 53 (larva).

Biology: Evans, 1957. Studies on compar. ethology Bembix, p. 59 (nesting site, prey).
—Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 269-288, figs. 149-154
(mating, nest, prey, egg, life cycle, parasite).

Superfamily SPHECOIDEA 1711

sayi Cresson. Gulf Coast, Miss. Valley, and Great Plains States west to Colo. and N. Mex.;
north. Mexico. Ecology: Nests in small numbers (3-10) in loose to compacted sand,
makes a 1-3 celled nest with an accessory burrow, stores 14 or more flies per cell.
Parasite: Amobiopsis confundens Tns., Senotainia rubriventris Macq., S. opiparis
Reinh., S. sp. in trilineata (Wulp) complex. Prey: Stratiomyidae sp.; Chlorotabanus:
crepuscularis Beq., Chrysops flavida Wied., C. spp., Hybomitra hinei wrighti Whit.,
Silvius quadrivittatus Say, Tabanus cheliopterus fronto O.8., T. fratellus Will., T.
lineola F., T. melanocerus lacustris Stone, T. nigripes Wied., T. quinquevittatus Wied.;
Exoprosopa fasciata Macq., E. fascipennis noctula Wied., Geron sp., Phthiria sp.,
Poecilanthrax lucifer F., Systoechus solitus Wlkr., S. vulgaris Lw., Villa cypris Meig.,
V. flavicostalis Paint., V. lateralis Say, V. molitor Lw.; Furcifera punctipennis Wied.;
Ablautus nigronotum Wilcox, Erax tabascens Bks., E. sp., Mallophorina laphroides
Wied., Asilidae sp.; Allograpta obliqua Say, Eristalis agrorum F., E. albifrons Wied., E.
tenax L., Microdon rufipes Macq., Syrphus sp., Volucella pusilla Macgq., V. sp.; Musca
domestica L., Orthellia caesarion Meig.; Phaenicia caeruleiviridis Macq.; Sarcophaga
ventricosa Wulp; Archytas analis F., Belvosia slossonae Coq., Fabriciella actinosa
Reinh., F. egula Reinh., F. latigena Tot., Gonia sequax Will., Juriniopsis sp.,
Paradidyma singularis Tns., Phorocera claripennis Macq., Prosenoides flavipes Coq.,
Tachinidae sp.

Bembex(!) sayi Cresson, 1865. Ent. Soe. Phila., Proc. 4: 467. 2.
Bembix latifrons Parker, 1917. U. S. Natl. Mus., Proc. 52: 116. 9.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 60, figs. 94-97 (larva).

Biology: Rohwer, 1909. Univ. Colo. Studies 6: 245 (nest, prey). —Hungerford and Williams,
1912. Ent. News 23: 247 (nest, prey). —Evans, 1957. Studies on compar. ethology Bembiz,
pp. 48-59, figs. 11-12 (mating, nest, prey, egg, life cycle, parasite). —Evans, 1966. Compar.
ethology and evolution of sand wasps, pp. 288-298, figs. 155-158 (nest, prey, parasites).
—Aleock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 154, fig. 1 (nest).

SPECIES GROUP CINEREA

These species nest in soil with a high saline content, make simple multicellular nests, and place
the egg on the first fly brought into the cell.

cinerea Handlirsch. N. J. to Fla. west to Tex. Ecology: Nests in very large aggregations on
salt flats in heavy black soil with high salt content, makes 1-2 cells per nest, stores 11-17
medium sized prey per cell. Parasite: Dasymutilla cypris (Bl.), D. vesta sappho (Fox).
Prey: Eulalia evansi James, Hedriodiscus dorsalis F.; Tabanus lineola F., T.
nigrovittatus Macq., T. vittiger bellardii Szil., T. spp.; Anthrax anale Say; Eristalis
albifrons Wied.; Dimecoenia austrina Coq.; Lispe'sp. near sordida Ald., Phyllogaster
sp.; Phaenicia cluvia Wlkr.; Gymnopsoa texana Tns.; Cistogaster sp.

Bembex(!) cinerea Handlirsch, 1893. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 102: 837. ¢,
Me
Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 59, fig. 109 (larva).

Biology: Krombein and Evans, 1954. Ent. Soc. Wash., Proc. 56: 235 (nest). —Krombein and
Evans, 1955. Ent. Soc. Wash., Proc. 57: 233-234 (nest, prey, parasite). —Evans, 1957.
Studies on compar. ethology Bembix, pp. 78-94, figs. 19-23 (male sun dance, mating, nest,
prey, egg, life cycle, parasite).

hinei Parker. La., Tex., coastal only. Ecology: Nests in hard packed sand beach above high-tide
mark, occasionally in large aggregations, makes at least 5 cells per nest, stores 7-13 prey
per cell. Prey: Eulalia cincta Latr.; Tabanus lineola F., T. nigrovittatus Macq.;
Eristalis albifrons Wied.
Bembizx hinei Parker, 1917. U. S. Natl. Mus., Proc. 52: 86. 6, 2.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 156 (larva).

Biology: Hine, 1906. La. State Crop Pest Comn., Cir. 6: 27 (nesting site, prey; misdet. as
belfragei Cr.). —Strandtmann, 1953. Kans. Ent. Soc., Jour. 26: 48-49 (nest, prey). —Evans,
1957. Studies on compar. ethology Bembix, pp. 94-104, figs. 24-26 (nest, prey transport,
egg, life cycle).

1712 Hymenoptera in America North of Mexico

SPECIES GRoUP AMERICANA

These species make simple multicellular nests, place the egg on the first prey brought into the
nest, and, so far as known, level the mound of excavated earth at the nest entrance.

americana comata Parker. Pacific Coast of N. Amer. to Mexico. Ecology: Nests in sand or

powdery soil, makes 1-3 cells per nest, provides 16-44 prey per cell. Parasite: Parnopes
edwardsii (Cr.); Physocephala affinis Will. Prey: Apatolestes hera 0. S.; Thereva
niveipennis Kroeb.; Hydrophorus gratiosus Ald., H. sp.; Eristalis arbustorum L., E.
latifrons Lw., E. tenax L., Heliophilus sp., Melanostoma sp., Mesograpta geminata Say,
M. marginata Say, Metasyrphus subsimis Fluke, Sphaerophoria cylindrica Say, S.
sulphuripes (Thom.), Syrphus sp., Toxomerus sp.; Ceroxys latiuscula Lw.; Coenosia
tigrina F., Helina bispinosa Mall., H. procedens Wlkr., Hylemya cilicrura Rond., Lispa
tentaculata DeG., Muscina assimilis Fall., Musca domestica L., Ophyra leucostoma
Weid., Pegomyia duplicata Mall.; Spaziphora cincta (Lw.).; Calliphora terraenovae
Macq., Phaenicia sericata Meig., Phormia regina Meig., Pollenia rudis F.; Sarcophaga
rapax Wlkr., S. spp., Senotainia trilineata (Wulp); Bonnetia comta (Fall.), Dexodes
cinerea Tns., Tachinomyia similis Will. Typical americana F. occurs in the West Indies.

Bembix comata Parker, 1917. U.S. Natl. Mus., Proc. 52: 100. 6, °.

Bembix comata var. nevadensis Rodeck, 1934. Amer. Mus. Novitates 692: 1. d.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 59 (larva).

Biology: Parker, 1925. Ent. Soc. Wash., Proc. 27: 189-195 (nest, prey capture and transport,
cocoon). —Bohart and MacSwain, 1940. Pan-Pacific Ent. 16: 16 (parasite). —Evans, 1957.
Studies on compar. ethology Bembix, pp. 36-48, figs. 8-10 (mating, nest, prey transport,
egg, parasite). —Alcock, 1973 (1972). Psyche 79: 158-164 (nest construction). — Alcock,
1973. Wasmann Jour. Biol. 31: 331-332 (prey).

americana hamata Fox. Calif., Santa Cruz to San Miguel Islands only.
Bembix hamata Fox, 1923. Psyche 30: 6. d.
Bembix hamata lucida Fox, 1923. Psyche 30: 7. d.
Bembizx sanctae-rosae Cockerell, 1940. South. Calif. Acad. Sci., Bul. 38: 135.

americana nicolai Cockerell. Calif., San Nicolas Island only.
Bembex(!) nicolai Cockerell, 1938. Pan-Pacifie Ent. 14: 150. 6, 2.

americana spinolae Lepeletier. N. Amer. except Pacific Coast States. Ecology: Nests in soil
varying from open sand to coarse soil, makes a unicellular nest, stores 16-24 prey per
cell. Parasite: Physocephala texana Will.; Phrosinella fulvicornis (Coq.)?, Opsidia
gonioides Coq., Senotainia vigilans Allen, S. trilineata (Wulp), Miltogrammini spp.;
Exoprosopa fascipennis (Say); Parnopes chrysoprasinus Sm.?, P. edwardsii (Cr.)?;
Dasymutilla bioculata (Cr.)?; Macrosiagon flavipenne LeC. Prey: Eulalia cincta Oliv.,
Odontomyia hoodiana Big., O. virgo Wied.; Chrysops aestuans abaestuans Phil., C.
flavida Wied., C. furcatus Wlkr., C. lugens Wied., C. mitis 0. S., C. niger Macgq., C.
noctifer pertinax Will., C. pudicus O. S., Hybomitra osburni Hine, H. phaenops 0. S.,
Silvius quadrivittatus Say, Tabanus atratus L., T. coffeatus Macq., T. daeckei Hine, T.
insuetus O.S., T. lasiopthalmus Macq., T. phaenops O.S., T. pumilus Macgq,, T.
quinquevittatus Wied., T. vittiger schwardti Phil.; Conophorus painteri Prid.,
Exoprosopa pueblensis Jaen., Phthiria sulfurea Lw., Sparnopolius brevirostris Macgq., S.
fulvus Wied., Systoechus fumipennis Paint., S. vulgaris Lw., Villa agrippina O.S., V.
alternata Say, V. fulviana Say, V. lateralis Say, V. sinuosa jaennickiana O. S.;
Furcifera rufiventris Lw., Psilocephala haemorrhoidalis Macq., Thereva cingulata
Kroeb.; Asilus sp., Atomosia melanopogon Herm., Lasiopogon cinereus Cole, Tolmerus
callidus Will.; Chrysotoxum ventricosum Lw., Dasysyrphus amalopsis O. S., Eristalis
arbustorum L., E. sp., Eupeodes volucris 0. S., Helophilus latifrons Lw., H. obscurus
Lw., Melanostoma sp., Metasyrphus astutus Fluke, M. palliventris Curr., Microdon
lanceolatus Adams, Paragus bicolor F., Scaeva pyrastri L., Sphaerophoria sp., Syrphus
opinator 0. S., S. ribesii L., Xylota bigelowi Curr.; Rivellia sp., Penocera quadrilineata
Mel., Sepedon fuscipennis Lw., Tetanocera plumosa Lw.; Cordilura latifrons Lw.,
Hydrophoria divisa Meig., Hylemya sp.; Calythea separata Mall., Helina cinerella
Wulp, H. latifrons Zett., H. troene Wlkr., Musca domestica L., Phaonia deleta Stein,

Superfamily SPHECOIDEA 1713

Pyrellia serena Meig., Quadrularia annosa Zett., Siphona irritans L., Spilogona sp.,
Stomoxys calcitrans L.; Eucalliphora lilaea Wlkr., Lucilia illustris Meig., L. sp.,
Phaenicia sericata Meig., Phormia regina Meig., Pollenia rudis F.; Eumacronychia sp.,
Sarcophaga derelicta Wlkr., S. importuna Wlkr., S. planifrons Ald., S. querula Wlkr.,
Senotainia flavicornis Tns., S. trilineata (Wulp); Bonellimyia glauca Brooks, Cuphocera
stricklandi Curr., Dinera sp., Microphthalma disjuncta Wied., Nearchus duplaris
Reinh., Paradidyma singularis Tns., Peleteria confusa Curr., Ptilodexia sp.,
Siphoplagia spinulosa Big., Trochilodes skinneri Coq., Winthemia quadripustulata F.

Bembex(!) spinolae Lepeletier, 1845. Hist. Nat. Ins. Hym., v. 3, p. 227. 6.

Bembex(!) similans Fox, 1895. Acad. Nat. Sci. Phila.; Proce. 47: 358. 6, 2.

Bembex(!) connexus Fox, 1895. Acad. Nat. Sci. Phila., Proce. 47: 360. d, °.

Bembex(!) primaaestate Johnson and Rohwer, 1908. Ent. News 19: 378. @.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 58, fig. 108 (larva).

Biology: Walsh and Riley, 1869. Amer. Ent. 1: 126 (nest, prey). —Peckham and Peckham,
1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 58-76 (nest, prey, parasite, life cycle). —Barber,
1915. Ent. Soc. Wash., Proc. 17: 187-188 (parasite). —Parker, 1917. U.S. Natl. Mus., Proc.
52: 127-131 (mating, nest, prey transport, life cycle, cocoon). —Rau and Rau, 1918. Wasp
studies afield, pp. 37-38 (nest, prey, parasite ?). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24
(7): 29, fig. 8 (nest). —Krombein, 1936. Ent. News 47: 95 (prey, parasite). —Evans, 1957.
Studies on compar. ethology Bembix, pp. 17-36, figs. 6-7 (mating, nest, prey capture and
transport, egg, life cycle, parasites). —Evans, 1966. Compar. ethology and evolution of
sand wasps, pp. 311-315, fig. 160 (nest, prey, parasites).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 21, figs. M-P (male
genitalia).
cameroni Rohwer. Southwest. U.S.; centr. Mexico. Ecology: Nests in alluvial soil with sand on

surface and frequently silt or loam beneath surface. Prey: E'senbeckia delta Hine; Geron
sp., Phthiria sulphurea Lw.; Eristalis latifrons Lw.; Musca domestica L., Orthellia
caesarion Meig.; Peleteria sp., Plagioprospherysa parvipalpis Wulp.

Bembyzx(!) cameroni Rohwer, 1912. U. S. Natl. Mus., Proce. 41: 467. 3.

Bembix festiva Parker, 1929. U. S. Natl. Mus., Proce. 75 (5): 118. 6.

Bembix rohweri Maidl and Klima, 1948 (1944). Jn Lohrmann, Muenchen. Ent. Gesell., Mitt.
34: 424. N. name for cameroni Roh. which is not preocc.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 310-311 (nest,
prey).
nubilipennis Cresson. Miss. Valley, Great Plains and southwest. States; Mexico (San Luis
Potosi). Ecology: Nests in large aggregations in hard-packed soil, makes 1-5 cells per
nest, stores up to 48 prey per cell. Parasite: Parnopes chrysoprasinus Sm.;
Miltogrammini spp. Prey: Hedriodiscus truquii (Bell.), Nemotelus trinotatus Mel.,
Stratiomys jamesi Steysk., S. meigenti Wied., S. nigriventris Lw.; Tabanus sulcifrons
Macq, T. trimaculatus Beauv.; Anthrax anale Say, A. irrorata Say, Bombylius io Will.,
Exoprosopa emarginata Macq., E. fasciata Macq., Poecilanthrax lucifera (F.),
Sparnopolius brevirostris Macq., Villa spp.; Neorhynchocephalus sackenii (Will.);
Atomosia puella (Wied.), Diogmites misellus Lw., D. umbrinus Lw., Erax sp. near
tuberculatus Coq., Proctacanthella cacopiloga (Hine); Eristalis arbustorwm (L.), E.
tenax (L.); Chrysomyza demandata (F.), Tetanops luridipennis Lw.; Paracantha culta
(Wied.); Musca domestica L., Orthellia caesarion (Meig.), Stomoxys calcitrans (L.);
Bufolucilia silvarum (Meig.), Lucilia illustris (Meig.), Phaenicia caeruleiviridis (Macq.),
P. sericata (Meig.), Pollenia rudis (F.); Sarcophaga assidua Wlkr., S. derelicta Wlkr., S.
helicis Tns., S. lherminieri R.-Desv., S. rapax Wlkr., S. sinuata Meig., S. ventricosa
Wulp, S. sp.; Aplomya theclarum (Seud.), Archytas apiciferus Wlkr., A. metallicus
(R.-Desv.), Gymnoclytia occidua (Wlkr.), Hyalomyodes triangulifera (Lw.), Phorocera
claripennis Macq., P. tachinomoides Tns., Trichopoda pennipes (F.).
Bembex(!) nubilipennis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 218. d, @.
Bembix nubilosa Parker, 1929. U.S. Natl. Mus., Proce. 75 (5): 140. @.

Taxonomy: Evans, 1959. Amer. Ent. Soc., Trans. 85: 42 (larva).

1714 Hymenoptera in America North of Mexico

Biology: Parker, 1910. Ohio Nat. 10: 163-165 (nest, prey, parasite ?). —Rau and Rau, 1918.
Wasp studies afield, pp. 9-37, figs. 2-6 (sun dance, nest, prey, life cycle, cocoon). —Rau,
1922. St. Louis Acad. Sci., Trans. 24 (7): 29-30 (sun dance, nest). —Rau, 1935. Psyche 41:
243-244 (nesting site). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp.
298-308, fig. 159 (sun dance, mating, nest, prey, egg, life cycle, parasites).

SPECIES GROUP TEXANA

These species make unicellular nests and place the egg in an erect position in the empty cell.
False burrows are frequently constructed, and one species removes the prey remains before
bringing in fresh prey.

melanaspis Parker. Southwest. U. S.; Mexico (Baja California). Ecology: Makes unicellular
nest in sand; lays egg in empty cell. Prey: Tabanidae, Therevidae, Syrphidae, Conopidae,
Calliphoridae; Coenagrionidae.
Bembix melanaspis Parker, 1917. U.S. Natl. Mus., Proc. 52: 109. 6, 2.

Biology: Aleock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 160-161, fig. 2 (nest, prey).

texana Cresson. Southeast. and south. States. Ecology: Nests in loose to hard-packed sand,

sometimes in large aggregations, makes a unicellular nest sometimes with one or two
accessory burrows, places egg in empty cell, usually removes old prey remains from cell
before bringing in fresh prey. Parasite: Dasymutilla pyrrhus (Fox)? Prey: Hedriodiscus
trivittatus Say, Odontomyia sp., Stratiomyidae sp.; Chlorotabanus crepuscularis Begq.,
Chrysops dimmocki Hine, C. pudica O. S., Hybomitra hinei wrighti Whit., Tabanus
abdominalis F., T. bishoppi Stone, T. coarctatus Stone, T. endymion O. S., T.
fuscicostatus Hine, T. gracilis Wied., T. lineola F., T. melanocerus lacustris Stone, T.
mularis Stone, T. nigripes Wied., T. quinquevittatus Hine, T. sparus milleri Whit., T.
trijunctus Wlkr.; Villa sp.; Meromacrus acutus F., Microdon fulgens Wied., Tubifera
sp.; Micropezidae sp.; Cochliomyia macellaria F., Phaenicia caeruleiviridis Macq.;
Exorista larvarum L., Tachinidae spp.

Bembiz fasciata Fabricius, 1804. Systema Piezatorum, p. 224. Preocc. This is a
questionable synonym.

Bembex(!) texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 219. °.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 62, figs. 105-107 (larva).

Biology: Evans, 1957. Studies on compar. ethology Bembix, pp. 135-141, fig. 37 (nest, egg,
prey, life cycle). —Krombein, 1958. Ent. Soc. Wash., Proc. 60: 107-110 (nest, prey).
—Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 323-335, figs. 165-170
(male sun dance, nest, prey, parasite 2).

troglodytes Handlirsch. Southwest. States north to Kans.; north. Mexico. Ecology: Nests in

fine-grained damp sand along watercourse, not markedly gregarious, makes unicellular
nest frequently with an accessory burrow, places egg in empty cell, stores 21-26 prey
per cell. Parasite: Timulla leona (Bl.)?; Exoprosopa fascipennis (Say)? Prey: Eulalia
communis James, Hedriodiscus truquii (Bell.), Hoplitimyia constans Lw., Stratiomys
meigenii Wied.; Chrysops sequax Will., C. wiedemanni Kroeb., Tabanus sp.; Exoprosopa
fascipennis (Say), E. iota (O. S.), Sparnopolius sp., Systoechus vulgaris Lw., Villa
chimaera (O. S.). V. salebrosa Paint.; Chrysogaster nitida Wied., Eristalis tenax L.;
Musca domestica L.; Orthellia caesarion Meig., Stomoxys calcitrans L.; Callitroga
macellaria F., Phormia regina Meig.; Amobia sp., Amobiopsis aurata Coq., Sarcophaga
derelicta Wlkr., S. lherminieri R.-Desv., S. opifera Coq., S. rapax Wlkr., S. ventricosa
Wulp, Senotainia flavicornis Tns., S. litoralis Allen, S. trilineata (Wulp), S. sp.;
Achaetoneura sp., Copecrypta nitens Wied., Gymnoclytia occidua Wlkr., Phorocera
claripennis Macq.

Bembex(!) troglodytes Handlirsch, 1893. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 102:
SPAY), Ci, Se

Bembix helianthopolis Parker, 1917. U.S. Natl. Mus., Proce. 52: 113. 6, @.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 61, figs. 102-104 (larva).

Biology: Evans, 1957. Studies on compar. ethology Bembix, pp. 116-135, figs. 29-36 (male sun
dance, mating, nest, egg, prey, life cycle, parasites?). —Evans, 1966. Compar. ethology and
evolution of sand wasps, pp. 335-337 (nest, prey).

Superfamily SPHECOIDEA 1715

SPECIES GROUP PRUINOSA

The two species of this group make deep, complex unicellular nests, place the egg flat in the
empty cell, and remove prey remains from the cell.

occidentalis Fox. West. and southwest. U. S.; north. Mexico. Ecology: Makes complex

unicellular nest in sand dunes, sometimes in large aggregations, lays egg in empty cell,
cleans nest frequently so number of prey provided is unknown. Parasite: Dasymutilla
sackenii (Cr.); Villa atrata (Coq.), V. melasoma Wulp, Exoprosopa eremita O. S.;
Physocephala affinis Will. Prey: Stratiomys laticeps Lw.; Apatolestes villosulus Big.,
Chrysops sp., Tabanus punctifer 0. S.; Aphoebantus tardus Cogq., A. sp., Villa sinuosa
Wied., V. sp. near agrippina O.8.; Psilocephala costalis Lw.; Apiocera haruspex O.5S.,
A. sp.; Eristalis latifrons Lw., Helophilus latifrons Lw., Lejops lululatus Meig.,
Melanostoma rostratum Big., Metasyrphus meadii Jones; Anacampta latiuscula Lw.;
Musca domestica L., Paregle cinerella Fall.; Calliphora erythrocephala Meig., Lucilia
illustris Meig., Phaenicia sericata Meig., Phormia regina Meig.; Sarcophaga bullata
Prkr., S. sp.; Aphria ocypterata Tns. Predator: Proctacanthus occidentalis Hine.

Bembex(!) occidentalis Fox, 1893. Calif. Acad. Sci., Proc. (2) 4: 10. 6, 9.

Bembex(!) beutenmuelleri Fox, 1901. N. Y. Ent. Soc., Jour. 9: 83. 6.

Bembex(!) obsoleta Howard, 1901. Insect Book, pl. 4, fig. 36.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 58, figs. 85-87 (larva).

Biology: Coquillett, 1895. Ent. Soc. Wash., Proc. 3: 236-237 (nest, prey; misdet. as cinerea
Handl.). —Bohart and MacSwain, 1939. South. Calif. Acad. Sci., Bul. 38: 84-98 (mating,
nest, prey, parasites). —Ross, 1953. Insects close up, pp. 42-43, 5 figs. (nest, prey, cocoon,
parasites). —Evans, 1957. Studies on compar. ethology Bembix, pp. 167-181, figs. 47-49
(mating, nest, prey).

pruinosa Fox. U.S.; Mexico. Ecology: Nests in large aggregations in loose sand, frequently in
dunes, makes a long unicellular nest sometimes with an accessory burrow, places egg in
empty cell, provides 20-30 prey per cell, placing later flies in a long single file. Parasite:
Dasymutilla bioculata (Cr.); Parnopes edwardsti (Cr.)?; Physocephala texana Will.;
Exoprosopa fascipennis (Say), E. arenicola Johns. and Johns., Villa atrata (Cogq.);
Senotainia inyoensis Reinh.? Prey: Anoplodonta nigrirostris Lw., Eulalia cincta Oliv.,
Odontomyia tumida Bks., Stratiomys jamesi Steysk., S. norma Wied.; Chrysops flavida
Wied., Tabanus equalis Hine, T. fulvulus Wied., T. lineola F., T. productus Hine, T.
quinquevittatus Wied., T. sulcifrons Macq., T. trispilus Wied., T. vittiger schwardti Phil.;
Anastoechus barbatus 0. S., Aphoebantus sp., Bombylius medorae Paint., Exoprosopa
fascipennis (Say), E. divisa Coq., Heterostylum robustum O. S., Lepidanthrax
proboscidea Lw., Poecilanthrax lucifera F., P. sackenii Coq., P. willistoni Coq., Villa
alternata Say, V. cypris Meig., V. faustina O.S., V. fulviana nigricauda Lw., V.
lateralis Say, V. molitor Lw.; Psilocephala aldrichii Coq., P. haemorrhoidalis Macq.;
Plagioneurus univittatus Lw.; Asemosyrphus polygrammus Lw., Eristalis albifrons
Wied., E. arbustorum L., E. brousti Will., E. latifrons Lw., E. tenax L., E. vinetorum F.,
Eupeodes volucris O. S., Helophilus latifrons Lw., Platycheirus erraticus Curr., Rhingia
nasica Say, Sericomyia chrysotoxoides Macq., Sphaerophoria robusta Curr., Syrphus
rectus O. S., S. torvus O. S., Xylota metallica Wied.; Ceroxys latiusculus Lw.;
Hydrophoria divisa Meig.; Limnophora narona Wlkr., Lispe cotidiana Snyd., L. nasoni
Stein, Musca autumnalis DeG., M. domestica L., Muscina assimilis Fall., M. dorsilinea
Wulp, Orthellia caesarion Meig., Pyrellia cyanicolor Zett., Stomoxys calcitrans L.;
Bufolucilia silvarum Meig., Callitroga macellaria F., Eucalliphora lilaea Wlkr., Lucilia
illustris Meig., Phaenicia sericata Meig., Phormia regina Meig., Pollenia rudis F.;
Amobia floridensis (Tns.), Sarathromyia femoralis Sch., Sarcophaga derelicta Wlkr., S.
errabunda Wulp, S. lherminieri R.-Desv., S. querula Wlkr.; Achaetoneura archippivora
Riley, Admontia sp., Archytas apicifera Wlkr., Paradidyma affinis Reinh., Peleteria
eronis Curr., Spallanzania sp., Sitophaga sp., Winthemia rufopicta Big.

Bembex(!) pruinosa Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 361. d, °.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 57, figs. 88-93 (larva).

1716 Hymenoptera in America North of Mexico

Biology: Mickel, 1924. Ent. News 35: 236-242 (parasite). —Evans, 1957. Studies on compar.
ethology Bembix, pp. 142-167, figs. 38-46 (hopping dance, mating, nest, prey, egg, life
cycle, parasites). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp.
345-349 (hopping dance, nest, prey capture, parasites).

Genus STICTIELLA Parker

Stictiella Parker, 1917. U. S. Natl. Mus., Proe. 52: 21.
Type-species: Monedula formosa Cresson. Orig. desig.
Microstictia Gillaspy, 1963. Ent. News 74: 196.
Type-species: Monedula femorata Fox. Orig. desig.

The genus is restricted to the Nearctic Region and all but one species occur in America north
of Mexico. The species whose biology is known use adult Lepidoptera as prey and usually prac-
tice mass provisioning, although there is one record suggestive of progressive provisioning in
serrata, a species also known to practice mass provisioning.

Taxonomy: Gillaspy, 1959. Pan-Pacifie Ent. 35: 193 (species groups). —Gillaspy, Evans and
Lin, 1962. Ent. Soc. Amer., Ann. 55: 559-566, 1 fig. (partition into Stictiella and Xerostictia
on morph. characters).

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 559-563, fig. 1 (ethology).
callista Parker. Ariz., N. Mex. Ecology: Nests in sand. Prey: Melipotis indomita (Wlkr.);

Strymon melinus pudica Edw.; Hesperiidae sp.
Stictiella callista Parker, 1917. U.S. Natl. Mus., Proc. 52: 34. d, @.

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 562 (prey). —Alcock and
Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 160 (nest, prey).

corniculata Mickel. Wyo. to Calif.
Stictiella corniculata Mickel, 1918 (1917). Nebr. Univ. Studies 17: 332. 6.

Taxonomy: Gillaspy, 1963. Ent. News 74: 251-252.
divergens Parker. Kans.
Stictiella divergens Parker, 1917. U. S. Natl. Mus., Proc. 52: 55. d.
emarginata (Cresson). Generally distributed throughout the U. S. and south. Canada. Prey:
Euxoa quinquelinea incallida (Sm.).
Monedula emarginata Cresson, 1865. Ent. Soc. Phila., Proc. 4: 468. d, °.
Monedula mamillata Handlirsch, 1890. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 99: 146.
Ge oe

Biology: Bradley, 1908. Ent. Soc. Amer., Ann. 1: 129 (sleeping aggregation). —Gillaspy,
Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 562 (prey).
exigua (Fox). Mont.
Monedula exigua Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 370. °.
femorata (Fox). Fla., Tex.
Monedula femorata Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 368. d.
formosa (Cresson). Kans., Okla., Tex. Ecology: Nests in sand, makes 5-17 cells per nest, stores
6-11 butterflies per cell, practices mass provisioning. Parasite: Acarina sp.; Diptera sp.
Prey: Polites sp., Hylephila phyleus Dru., Atalopedes campestris Boisd., Pyrgus
communis Grt., Thanaos zarucco Luc.; Phyciodes phaon Edw., Libythea larvata Streck.,
L. sp.; Strymon melinus Hbn.
Monedula formosa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 221. 6, 2.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 260, figs. 37-41 (larva).
Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 559-561 (nest, prey,
parasite).
minutula (Handlirsch). Tex.
Monedula minutula Handlirsch, 1890. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 99: 148.
OF
nubilosa Gillaspy. Calif. (Los Angeles).
Stictiella nubilosa Gillaspy, 1963. Ent. News 74: 252. °.

Superfamily SPHECOIDEA LANG,

plana (Fox). Great Plains States.
Monedula plana Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 367. d.
pulchella (Cresson). Colo. to Calif. Ecology: Usually makes unicellular nest in sand but rarely
2 cells, stores as many as 19 moths in a completed cell. Prey: Leptotes marina Reak.;
Loxostege similalis Guen.; Characoma proteella Dyar.
Monedula pulchella Cresson, 1865. Ent. Soc. Phila., Proc. 4: 471. 6, @.
Stictiella melanosterna Parker, 1917. U. S. Natl. Mus., Proe. 52: 30. 6, 2.

Taxonomy: Gillaspy, 1963. Ent. News 74: 252 (synonymy). —Evans, 1964. Amer. Ent. Soc.,
Trans. 90: 260-261, fig. 42 (larva).

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 560, 562-563 (nest, prey,
sleeping aggregation).
serrata (Handlirsch). Fla., Ga., N. C. Ecology: Nests in fine-grained sand, makes unicellular

nest, stores 12 to more than 21 moths per cell, practices both mass and progressive
provisioning. Parasite: Senotainia sp. near rubriventris Macq. Prey: Elasmopalpus
lignosellus (Zell.); Hellula rogatilis (Hulst); Bactra verutana Zell., Eucosminae sp.;
Crambus satrapellus Zink. C. quinquareatus Zell., C. sp., Argynia argentana Martyn;
Jocara sp.

Monedula serrata Handlirsch, 1890. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 99: 143. 3.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soe., Trans. 82: 51, figs. 59-62 (larva).

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 561-562, fig. 1 (nest, prey
transport, egg, life cycle). —Krombein, 1964. Amer. Mus. Novitates 2201: 19-20 (nest, prey
transport, life cycle, cocoon).

speciosa (Cresson). Alta., Ariz., Sask., Colo., N. Mex., Nebr., Kans.
Monedula speciosa Cresson, 1865. Ent. Soc. Phila., Proc. 4: 470. 2.
spinifera (Mickel). Alta., Kans., Nebr.
Stictia spinifera Mickel, 1916. Amer. Ent. Soc., Trans. 42: 418. 6.
Stictiella melampous Parker, 1917. U.S. Natl. Mus., Proce. 52: 43. 6.
tuberculata (Fox). Idaho, Nev., Wash.
Monedula tuberculata Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 366. 3.

Genus GLENOSTICTIA Gillaspy

Glenostictia Gillaspy, 1962. Jn Gillaspy, Evans and Lin, Ent. Soc. Amer., Ann. 55: 563.
Type-species: Monedula pulla Handlirsch. Orig. desig.

This genus occurs only in the Nearctic Region and all species are found in America north of
Mexico. So far as known the members of the genus practice progressive provisioning and prey
upon adult Hymenoptera, Diptera and Hemiptera.

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 563-566 (ethology).
argentata (Fox). Calif.; Mexico. Ecology: Nests in sand dune, practices progressive
provisioning. Prey: Primarily Bombyliidae.
Stictiella argentata Fox, 1923. Calif. Acad. Sci., Proc. (4) 12: 484. d, 9.

Biology: Alcock, 1975. Southwest. Nat. 20: 339 (nest, prey).
bifurcata (Fox). South. Calif.; Mexico (Baja California, Sonora).
Stictiella bifurcata Fox, 1923. Calif. Acad. Sci., Proc. (4) 12: 431. ¢.
Stictiella bifurcata var. albicera Fox, 1923. Calif. Acad. Sci., Proc. (4) 12: 482. ¢.
Stictiella directa Fox, 1923. Calif. Acad. Sci., Proc. (4) 12: 488. ¢, 2.
bituberculata (Parker). Ariz., Calif., N. Mex.
Stictiella bituberculata Parker, 1917. U. S. Natl. Mus., Proe. 52: 36. 6, 9°.
clypeata (Gillaspy). West. Tex. to south. Calif.; Mexico (Chihuahua, Coahuila, Guerrero). Prey:
Apiocera augur O. S.; Tubifera latifrons (Lw.).
Stictiella clypeata Gillaspy, 1959. Pan-Pacifie Ent. 35: 187. 6, @.

Biology: Painter, 1936. Kans. Univ. Sci. Bul. 24: 192 (prey; misdet. as tenwicornis (Fox)).
—Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 563 (prey).

1718 Hymenoptera in America North of Mexico

gilva Gillaspy. Ariz., Calif. Ecology: Nests in dry sand, provisions progressively. Prey:
Aphoebantus interruptus Coq., A. sp.
Glenostictia gilva Gillaspy, 1963. Ent. News 74: 198. 6, ?.

Biology: Evans, 1966. Compar. ethology and evolution of sand wasps, pp. 188-189 (nest,
prey). —Alcock, 1975. Southwest. Nat. 20: 339 (nest, prey).
megacera (Parker). Colo., Utah, Wash.; Mexico.
Stictiella megacera Parker, 1917. U. S. Natl. Mus., Proc. 52: 49. 6, 2.
pictifrons (Smith). Generally distributed from Pa. and Ga. west to Colo., Tex., and Calif. Prey:
Villa sp. in lateralis Say group.
Monedula pictifrons Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 335. 2.
Monedula inermis Handlirsch, 1890. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 99: 144. ¢.
Monedula denverensis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 235. °.

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 563-564 (prey). —Alcock,
1975. Southwest. Nat. 20: 339-340 (prey-hunting behavior).

pulla (Handlirsch). Western U. S. Ecology: Nests in sand, practices progressive provisioning.
Parasite: Miltogrammini sp. Prey: Lepidanthrax sp., Geron sp., Aphoebantus sp.;
Psilocephala aldrichi Coq.; Eupeodes volucris 0. S., Helophilus latifrons 0. S.;
Hylemya cilicrura (Rond.); Senotainia rubriventris (Macq.), Sarcophaga opifera Coq,, S.
tuberosa Pand., S. sp.; Stomatomyia parvipalpis (Wulp).
Monedula pulla Handlirsch, 1890. Akad. Wiss. Wien, Math.-Nat. KL, Sitzber. 99: 149: 9:
Monedula usitata Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 371. ¢ (2 misdet.).

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 52, figs. 63-65 (larva).

Biology: LaRivers, 1942. Pan-Pacifie Ent. 18: 4-8 (nest, prey, parasite). —Gillaspy, Evans and
Lin, 1962. Ent. Soc. Amer., Ann. 55: 564 (nest, prey, sleeping aggregation). —Evans, 1966.
Compar. ethology and evolution of sand wasps, pp. 186-188, fig. 105 (nest, prey).

scitula (Fox). Ariz., Calif., N. Mex., Tex. Ecology: Nests in sandy loam or heavier stony soil,

makes 5 or more cells per nest, may store as many as 40 prey per cell, provisions
progressively. Parasite: Parnopes concinnus Vier.?; Senotainia sp.? Prey: Cicadellidae
sp.; Heteropsylla tecana Cwfd.; Neurocolpus arizonae Knight, Psallus sp.; Procladius
sp. near bellus Lw.; Dicyphoma schaefferi Coq., Zabrachia sp.; Silvius quadrivittatus
Say; Mythicomyia intermedia Mel., Oligodranes sp., Phthiria sp.; Brevitrichia griseola
Coq., Scenopinus sp.; Holopogon phaeonotus Lw., Asilidae sp.; Allograpta obliqua Say,
Mesograpta marginata Say; Euxesta magdalenae Cr., E. nitidiventris Lw.; Trupanea
bisetosa Coq.; Conioscinella sp.; Melanagromyza sp.; Astiosoma sp.; Milichia aethiops
Mall.; Calythea micropteryx Thom., Hylemya platura Meig., Pegomya longimana Pok.;
Haematobia irritans L., Musca domestica L.; Eumacronychia sp.; Siphophyto setigera
Coq.; Apanteles sp.; Torymus sp.; Iridomyrmex pruinosus Rog.; Lindenius sp.;
Tachysphex sp.; Colletes sp.; Perdita exclamans Ckll., P. knulli Timb., P. marcialis
CkIl., P. mentzeliarum CkIL, P. larreae Ckll., P. n. nwmerata CkIl., P. spp.; Dufourea sp.,
Lasioglossum sp.; preferred prey are Perdita bees, but other small Hymenoptera,
Diptera and Hemiptera are also used.

Monedula scitula Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 369. °.

Monedula villosa Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 370. ¢.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 262, figs. 48-52 (larva).

Biology: Gillaspy, Evans and Lin, 1962. Ent. Soc. Amer., Ann. 55: 564-566 (nest, prey,
sleeping aggregation). —Evans, 1966. Compar. ethology and evolution of sand wasps, pp.
189-205, figs. 106-109 (nest, prey transport, egg, cocoon, life cycle, parasite).
tenuicornis (Fox). Calif., Ariz., Tex.

Monedula tenwicornis Fox, 1895. Acad. Nat. Sci. Phila., Proc. 47: 368. 2, 6.
terlinguae Fox. Tex.

Stictiella terlinguae Fox, 1928. Pan-Pacific Ent. 4: 108. d, 9.

Genus XEROSTICTIA Gillaspy
Xerostictia Gillaspy, 1963. Ent. News 74: 187.

Superfamily SPHECOIDEA 1719

Type-species: Xerostictia longilabris Gillaspy. Orig. desig.

The genus contains only one polytypic species occurring in the southwestern deserts of the
Nearctic Region.

longilabris longilabris Gillaspy. Ariz., Calif. Ecology: Makes a multicelled nest in sand dunes,
practices progressive provisioning. Prey: Brachynemurus longipalpis Hag. adults;
Ormenis saucia Van D. adults. Another subspecies occurs in Baja California.
Xerostictia longilabris longilabris Gillaspy, 1963. Ent. News 74: 187, figs. 1-7. 2, d.

Biology: Alcock, 1975. Southwest. Nat. 20: 340-341, figs. 3-5 (nest, prey).

Genus STENIOLIA Say

Steniolia Say, 1837. Boston Jour. Nat. Hist. 1: 367.
Type-species: Bembex(!) longirostra Say. Monotypic.

All but three species occur in western America north of Mexico; two of the extralimital spe-
cies range as far south as Venezuela or Ecuador. The wasps nest in small aggregations, con-
struct shallow unicellular nests and practice progressive provisioning. Preferred prey of most
species are bee-flies (Bombyliidae). The egg is laid on the first prey brought into the nest.
Adults of both sexes form large sleeping aggregations on shrubs or plants, often at some
distance from the nesting site.

Revision: Gillaspy, 1964. Amer. Ent. Soc., Trans. 89: 1-117, 6 pls.

Biology: Evans and Gillaspy, 1964. Amer. Midland Nat. 72: 257-280, 14 figs. (comparative
ethology).

californiensis Gillaspy. Calif.; Mexico (Baja California).
Steniolia californiensis Gillaspy, 1964. Amer. Ent. Soc., Trans. 89: 52, figs. 5, 10, 25, 29, 34,
DUNO ee
dissimilis Fox. Southern Ariz.; Mexico (Sonora, Sinaloa).
Steniolia dissimilis Fox, 1923. Calif. Acad. Sci. Proce. (4) 12: 429. 6, @.
duplicata Provancher. Western Tex. to Calif., Nev.; Mexico (Baja California, Chihuahua,
Coahuila). Ecology: Nests in moderately sandy soil. Parasite: Parnopes concinnus Vier.?
Prey: Villa sp. near alternata Say, V. pallida (Coq.), V. spp., Aphoebantus sp. near
hirsutus Coq.; Paragus tibialis Fall., Mesogramma marginata Lw., Eupeodes volucris
O. S.; Syrphidae spp.; Fannia sp.; Sarcophaga sp.; Calliphoridae sp.; Syrphidae appear
to be the preferred prey.
Steniolia duplicata Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 414. 6, @.
Steniolia edwardsii Patton, 1894. Ent. Soc. Wash., Proc. 3: 45. Nom. nud.
Steniolia meridionalis Fox, 1923. Calif. Acad. Sci., Proc. (4) 12: 430. 6.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 263-264, figs. 34-36 (larva).

Biology: Bradley, 1908. Ent. Soc. Amer., Ann. 1: 129 (sleeping aggregation). —Evans and
Gillaspy, 1964. Amer. Midland Nat. 72: 274-276 (clustering, nest, prey). —Evans, 1966.
Compar. ethology and evolution of sand wasps, p. 205 (parasite).

elegans Parker. Wyo. to Wash., south to western Tex. and southern Calif.; Mexico (Baja
California, Sonora, Coahuila, Aguascalientes, San Luis Potosi, Hidalgo). Ecology: Nests
in powdery, rocky soil. Prey: Anthrax irroratus Say, Poecilanthrax signatipennis (Cole),
Systoechus vulgaris Lw., Villa sinuosa Wied., V. spp.; Holopogon atripennis Back;
Eupeodes volucris 0. S., Microdon coarctatus Lw., Volucella sp.; Bombyliidae are the
preferred prey.
Steniolia elegans Parker, 1929. U.S. Natl. Mus., Proc. 75 (5): 50. ¢.

Taxonomy: Evans and Lin, 1956 Amer. Ent. Soc., Trans. 82: 52, figs. 66-71 (larva).

Biology: Evans and Gillaspy, 1964. Amer. Midland Nat. 72: 276 (clustering, nest, prey
capture). —Evans, 1973. Great Basin Nat. 33: 29-30 (nest, prey transport).
eremica Gillaspy. Southern Calif., Ariz., Nev. Prey: Efferia texana (Bks.).
Steniolia eremica Gillaspy, 1964. Amer. Ent. Soc., Trans. 89: 67, figs. 14, 38, 54, 84. d, @.

Biology: Evans and Gillaspy, 1964. Amer. Midland Nat. 72: 276-277 (prey).

1720 Hymenoptera in America North of Mexico

nigripes Parker. Calif.; Mexico (Baja California). Ecology: Nests in bare desert soil. Prey:
Aphoebantus sp. near tardus Coq., A. hirsutus Coq.
Steniolia nigripes Parker, 1917. U. S. Natl. Mus., Proc. 52: 8. d.

Taxonomy: Evans and Lin, 1956. Amer. Ent. Soc., Trans. 82: 53 (larva).

Biology: Gillaspy, 1951. Pan-Pacific Ent. 27: 167-168 (nest, prey, cocoon).
obliqua (Cresson). Mont. to N. Mex., west to B. C. and Calif. Ecology: Nests in fine-grained,
powdery sandy loam. Parasite: Parnopes chrysoprasinus Sm.?, P. edwardsii (Cr.);
Mutillidae sp.; Taxigramma heteroneura (Meig.)?, Hilarella hilarella (Zett.)? Prey: Villa
sinuosa jaennickeana O. S., V. l. lateralis Say, V. a. alternata (Say), V. a. nigropecta
Cr., V. concessor Coq., Geron sp., Bombylius sp.; Pipiza calcarata Lw.; Lucilia sp.;
Bombyliidae are the preferred prey.
Monedula obliqua Cresson, 1865. Ent. Soc. Phila., Proc. 4: 469. °.
Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 265, fig. 45 (larva).

Biology: Evans and Gillaspy, 1964. Amer. Midland Nat. 72: 266-272, figs. 1-10, 13, 14
(clustering, mating, nest, prey capture and transport, life cycle, cocoon, parasites).
—Evans, 1970. Mus. Compar. Zool., Bul. 140: 494-495 (nest, prey, parasites).

scolopacea albicantia Parker. B. C. to northern Calif., Idaho.

Steniolia albicantia Parker, 1917. U. S. Natl. Mus., Proce. 52: 12. 6.

Biology: Evans and Gillaspy, 1964. Amer. Midland Nat. 72: 261 (clustering).

scolopacea scolopacea Handlirsch. Calif., Nev.; Mexico (Baja California).

Steniolia scolopacea Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98: 510.

Gee.

sulfurea Fox. Calif.

Steniolia sulfurea Fox, 1901. N. Y. Ent. Soc., Jour. 9: 84. 6.
tibialis Handlirsch. Wash. to Calif., Nev., Idaho. Prey: Conophorus nigripennis (Lw.); Syrphus

sp.
Steniolia tibialis Handlirsch, 1889. Akad. Wiss. Wien, Math.-Nat. K1., Sitzber. 98: 513. ¢,
on

Biology: Evans and Gillaspy, 1964. Amer. Midland Nat. 72: 276 (prey).

vanduzeei Gillaspy. Calif., Nev.
Steniolia vanduzeei Gillaspy, 1964. Amer. Ent. Soc., Trans. 89: 62, figs. 13, 37, 538, 83. d, °.

Family PHILANTHIDAE
So far as known all members of this family are ground-nesters.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 79-89, figs. 1-35 (larvae). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 156-157 (larvae). — Menke, 1967. Pan-Pacific Ent. 43: 147-148
(key to genera). —Bohart and Grissell, 1975. Calif. Ins. Survey, Bul. 19: 1-92, 151 figs., 46
maps (Calif. spp. with keys to North American Philanthinae).

Biology: Alcock, 1975. Kans. Ent. Soc., Jour. 48: 532-545, 7 figs. (male mating strategy).
SUBFAMILY PHILANTHINAE

Genus PHILANTHUS Fabricius

Philanthus Fabricius, 1790. Skrivt. Natur. Hist. Selsk., v. 1, p. 224.
Type-species: Philanthus coronatus Fabricius. Desig. by Shuckard, 1837.
Symblephilus Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 171.
Type-species: Philanthus pictus Panzer. Desig. by Pate, 1937.
Simblephilus Jurine, 1807. Nouv. Method. Class. Hym. Dipt., p. 185.
Type-species: Vespa triangulum Fabricius. Desig. by Morice and Durrant, 1914.
Cheilopogonus Westwood, 1834. Zool. Jour. 5: 441.
Type-species: Cheilopogonus punctiger Westwood. Monotypice.
Philianthus Guerin, 1835. Iconogr. Regne Anim., Ins., pl. 71, fig. 8. Lapsus.
Anthophilus Dahlbom, 1844. Hym. Europaea, v. 1, p. 190.

Superfamily SPHECOIDEA 1721

Type-species: Philanthus politus Say. Desig. by Ashmead, 1899.
Chilopogon Kohl, 1896. K. K. Naturhist. Hofmus., Ann. 11: 329. Emend. of Cheilopogonus
Westwood.
Epiphilanthus Ashmead, 1899. Canad. Ent. 31: 294.
Type-species: Philanthus solivagus Say. Orig. desig.
Pseudanthophilus Ashmead, 1899. Canad. Ent. 31: 294.
Type-species: Philanthus ventilabris Fabricius. Orig. desig.
Oclocletes Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 423.
Type-species: Philanthus sanbornii Cresson. Orig. desig.
Ococletes Mickel, 1916. Amer. Ent. Soc., Trans. 42: 407. Emend. or lapsus.

This genus occurs in all major zoogeographic regions except The Neotropical and Australian.
The preferred prey are bees, sometimes honeybees, but occasionally wasps or even Parasitica
may be used, perhaps when there is a shortage of bees.

Revision: Strandtmann, 1946. A Rev. of the N. Amer. Spp. of Philanthus, 126 pp., 8 pls.

albopilosus Cresson. Rocky Mtn. States east to Tex., Ohio, and Ont.; Mexico (Chihuahua).

Ecology: Makes up to 4 cells per nest with 1-6 accessory burrows per nest, stores 6-10
prey per cell. Parasite: Phrosinella fulvicornis Coq. Prey: Ancistrocerus c. catskill
(Sauss.); Aphilanthops frigidus (Sm.); Mimumesa fuscipes (Pack.); Oxybelus
bipunctatus Oliv.; Colletes hyalinus Prov., C. simulans armatus Patt., C. willistoni
Robt.?; Andrena robertsonii D. T.; Agapostemon angelicus Ckll., Dialictus imitatus
(Sm.), D. lineatulus (Cwfd.), D. pilosus (Sm.), Halictus confusus Sm., H. ligatus Say, H.
rubicundus Chr., Lasioglossum leucozonium (Schr.).

Philanthus albopilosus Cresson, 1865. Ent. Soe. Phila., Proce. 5: 91. 6.

Philanthus simillimus Cresson, 1865. Ent. Soc. Phila., Proc. 5: 95. °.

Biology: Evans, 1975. Ent. Soc. Amer., Ann. 68: 888-892, 6 figs. (male behavior, nest, prey,
parasite).
arizonicus Bohart. Ariz. (Maricopa Co.).
Philanthus arizonicus Bohart, 1972. Ent. Soe. Wash., Proc. 74: 397, fig. 1. 3.
barbatus Smith. Idaho and Oreg. to Colo. and N. Mex.; Mexico (Chihuahua to Puebla). Ecology:
Nests in sand dune. Prey: Agapostemon melliventris Cr.; Exomalopsis sidae Ckll.
Philanthus barbatus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 473. 6.
Philanthus albifrons Cresson, 1865. Ent. Soc. Phila., Proc. 5: 101. d, 2.
Philanthus henricus Dunning, 1898. Canad. Ent. 30: 153. °.

Biology: Evans and Lin, 1959. Wasmann Jour. Biol. 17: 128 (nest, prey).

barbiger Mickel. Colo., Nebr.
Philanthus barbiger Mickel, 1916. Amer. Ent. Soc., Trans. 42: 405. 6, @.
bicinctus (Mickel). Colo., Utah, Wyo. Ecology: Nests in aggregations, makes very deep nest in
friable, clay-silt soil, stores 5 prey per cell. Parasite: Metopia argyrocephala (Meig.) ?
Prey: Bombus bifarius Cr., B. occidentalis Gr., B. fervidus (F.), B. melanopygus Nyl., B.
flavifrons Cr., B. mixtus Cr., B. rufocinctus Cr., B. centralis Cr.; preys on bumblebee
workers and males.
Ococletes bicinctus Mickel, 1916. Amer. Ent. Soc., Trans. 42: 407. °.
Ococletes hirticulus Mickel, 1918 (1917). Nebr. Univ. Studies 17: 326. ¢.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 275-276, figs. 65-67 (larva).

Biology: Armitage, 1965. Kans. Ent. Soc., Jour. 38: 89-100, 4 figs. (nest, prey). —Evans, 1970.
Mus. Compar. Zool., Bul. 140: 500 (nest, prey, parasite ?).
bilunatus Cresson. East. U. S. west to N. Dak., Colo., and N. Mex., east. Canad. provinces west

to Alta. Ecology: Nests in flat sand or vertical sand banks, makes 3 or more cells per
nest, stores 7-8 small bees per cell. Prey: Hylaeus m. modestus Say; Halictus ligatus
Say, H. confusus Sm., Lasioglossum leucozonium (Schr.), Dialictus inconspicuus (Sm.),
Augochlora striata Prov.

Philanthus bilunatus Cresson, 1865. Ent. Soc. Phil., Proc. 5: 97. 6.

Philanthus scelestus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxxiii. °.

Philanthus assimilis Banks, 1915. Canad. Ent. 47: 404. 6. Preoce.

Philanthus consimilis Banks, 1923. Canad. Ent. 55: 21. N. name. Preoce.

1722 Hymenoptera in America North of Mexico

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 83, fig. 19 (larva).

Biology: Evans and Lin, 1959. Amer. Midland Nat. 17: 126-127, figs. 4, 5, 9 (nest, prey, egg,
life cycle).
crabroniformis Smith. Rocky Mt. and Pacific Coast States, N. Dak., Alta., B. C. Ecology:
Nests in rather diffuse aggregations in hard stony soil, usually makes only one nest
during lifetime of an individual female containing up to 15 cells, stores 12-24 small prey
per cell. Parasite: Senotainia trilineata (Wulp), Phrosinella pilosifrons Allen, M etopia
argyrocephala (Meig.) ? Prey: Astata bakeri Parker, A. nubecula Cr.; Plenoculus davisi
Fox.; Ectemnius sp.; Hylaeus affinis (Sm.), H. sp.; Andrena albosellata Ckll., A. spp.,
Perdita fallax Ckll.; Agapostemon texanus Cr., Dialictus incompletus (Cwfd.), D.
laevissimus (Sm.), D. spp., Evylaeus cooleyi (Cwfd.), E. niger (Vier.), E. peraltus (Ckll.),
E. synthridis (Ckll.), Halictus confusus (Sm.), H. farinosus (Sm.), H. rubicundus Chr.,
H. tripartitus Ckll., Sphecodes patruelis Ckll., S. sulcatulus Ckll., S. spp.; Apis mellifera
L.; small bees are the preferred prey.
Philanthus crabroniformis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 474. d.
Philanthus flavifrons Cresson, 1865. Ent. Soc. Phila., Proc. 5: 102. 2.
Philanthus sublimis Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxxil. 6.
Philanthus californicus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proe., p. xxxii. d.
Liris magnifica Provancher, 1895. Nat. Canad. 22: 130. d.

Biology: Bohart, 1954. Ent. Soc. Wash., Proc. 56: 26-27 (prey). —Evans, 1970. Mus. Compar.
Zool., Bul. 140: 489-499 (nest, prey). —Aleock, 1974. Jour. Zool. 173: 233-246, 2 figs., 3 pls.
(nest, prey capture and transport, mating, parasite, competition with P. gibbosus (F.)).

crotoniphilus Viereck and Cockerell. Calif., N. Mex., Tex.; Mexico (Coahuila).
Philanthus crotoniphilus Viereck and Cockerell, 1904. N. Y. Ent. Soe., Jour. 12: 145. d.
gibbosus (Fabricius). Transcont. in south. Canada and U. S.; Mexico to El Salvador. Ecology:

Nests in flat or vertical soil of types varying from sandy loam to compacted heavy soil,
makes 7-15 or more cells per nest, stores 8-16 small prey per cell; rarely 2 females may
utilize a joint burrow; 1 female may nest in parental burrow; males frequently occupy
parental burrow for life. Parasite: Metopia argyrocephala (Meig.), Hilarella hilarella
(Zett.), Senotainia trilineata (wulp) complex; Dasymutilla nigripes (F.) ?,
Sphaeropthalma sp. Prey: Crossocerus sulcus (Fox); Colletes tucsonensis Ckll.;
Agapostemon virescens (F.), Augochlora pura (Say), Augochlorella striata (Prov.), A.
sp., Augochloropsis cuprea (Sm.), A. m. metallica (F.), Dialictus euryceps (Ellis), D.
incompletus (Cwfd.), D. imitatus (Sm.), D. laevissimus (Sm.), D. lineatulus (Cwfd.), D.
microlepoides (Ellis), D. obscurus (Robt.), D. pilosus (Sm.), D. rohweri (Ellis), D.
zephyrus (Sm.), D. spp., Halictus confusus Sm., H. ligatus Say, H. tripartitus Ckll.,
Lasioglossum coriaceum (Sm.), L. zonulum (Sm.), Sphecodes arvensis Patt., S. spp.
Predator: Diogmites discolor Lw.; Misumena calycina (L.).

Vespa gibbosus Fabricius, 1775. Systema Ent., p. 370.

Philanthus punctatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app., p. 342. d.

Cheilopogonus punctiger Westwood, 1835. Zool. Jour. 5: 441. d, °.

Anthophilus gibbosus Dahlbom, 1844. Hym. Europaea, v. 1, pp. 192, 497. 6, 2. Preoce.

Anthophilus nodosus Klug, 1846. Akad. Wiss. Berlin. Ber. Verh., p. 42. Lapsus.

Philanthus xanthostigma Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 131, pl. 8, fig.
L2G

Philanthus maculifrons Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 132, pl. 8, fig. 18.

é.

Philanthus punctatus var. cockerelli Dunning, 1897. Ent. News 8: 69. 3.

E. punctatus var. chilopsidis Cockerell, 1898. Davenport Acad. Nat. Sci., Proc. 7: 141. °.
The E. presumably refers to Epiphilanthus inasmuch as this new var. was mentioned in
a list of Philanthidae.

Anthophilus melanaspis Cameron, 1905. Amer. Ent. Soc., Trans. 31: 377. °.

Anthophilus maculiventris Cameron, 1905. Amer. Ent. Soc., Trans. 31: 377. °.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 82, figs. 11-15 (larva).

Superfamily SPHECOIDEA 1723

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 117-124 (nest,
prey). —Rau and Rau, 1918. Wasp studies afield, pp. 109-116 (nest, prey). —Reinhard,
1924. Smithsn. Inst., Ann. Rpt. for 1922, pp. 363-376. —Reinhard, 1929. Witchery of wasps,
pp. 92-140, 2 pls., 5 figs. (nest, prey capture, cocoon, life cycle, parasite, predators).
—Evans and Lin, 1959. Amer. Midland Nat. 17: 124-125, fig. 3 (nest, prey). —Cazier and
Mortenson, 1965. So. Calif. Acad. Sci., Bul. 64: 174-197, figs. 1-4 (nest, prey). —Lin, 1968.
Ent. Soc. Wash., Proc. 70: 10-12 (sleeping behavior). —Evans 1973. Anim. Behaviour 21:
302-308, 3 figs. (burrow sharing, nest transfer). —Barrows and Snyder, 1974 (1973). Ent.
News 84: 314-316 (prey, parasite). —Alcock, 1974. Jour. Zool. 173: 241-246 (nest, prey
transport, competition with P. crabroniformis).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 20, figs. C-E (male
genitalia).
gloriosus Cresson. Alta., Ariz., Kans., Nebr., N. Dak., N. Mex., Tex.; Mexico (Chihuahua).
Philanthus gloriosus Cresson, 1865. Ent. Soc. Phila., Proc. 5: 86. °.
Philanthus insignatus Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 421. @.
inversus Patton. Great Plains, Iowa, and Kans., north to Alta. and N. Dak., Calif.
Philanthus inversus Patton, 1879. U.S. Geol. and Geog. Survey, Bul. 5: 355. 2.
lepidus Cresson. East. U. S. west to Tex. and Colo. Ecology: Nests in large aggregations in
sand banks, makes 1-5 false burrows and 2 or more cells per nest, stores 9-11 small bees
per cell. Parasite: Senotainia trilineata (Wulp) complex? Prey: Pseudopanurgus
andrenoides (Sm.), Augochlora pura (Say), Augochlorella striata (Prov.), Dialictus
apertus (Sandh.), D. cressonii (Robt.), D. inconspicuus (Sm.), D. laevissimus (Sm.), D.
tegularis (Robt.), D. versans (Lov.), D. spp., Evylaeus divergenoides Mitch., E.
macoupinensis (Robt.), Halictus ligatus Say.
Philanthus lepidus Cresson, 1865. Ent. Soe. Phila., Proc. 5: 92. 3.
Philanthus carolinensis Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 422. 9.
Philanthus carolinensis var. reductus Banks, 1921. Ent. Soc. Amer., Ann. 14: 18. ¢, @.

Biology: Evans, 1964. Psyche 71: 142-149, 4 figs. (nest, prey, parasite 7).
levini Bohart. South. Calif.
Philanthus levini Bohart, 1972. Ent. Soc. Wash., Proc. 74: 398, fig. 2. d, 2.
multimaculatus Cameron. Rocky Mt. States, Alta., B. C.; Mexico (Veracruz). Ecology: Nests in
slopes or vertical banks of dry sandy soil or mixture of sand, earth and clay, makes up
to 5 cells per nest, stores up to 8 small bees per cell. Parasite: Dasymutilla dilucida
Mick.?; Senotainia trilineata (Wulp) complex? Prey: Dialictus clematisellus (Ellis), D.
microlepoides (Ellis), D. perparvus (Ellis), D. pruinosiformis (Cwfd.), D. sp., Sphecodes
spp.
Philanthus multimaculatus Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 133, pl. 8, fig.
14. 6.
Philanthus anna Dunning, 1897. Ent. News 8: 68. 6.
Philanthus multiannulatus Dalla Torre, 1897. Cat. Hym., v. 8, p. 488. Lapsus.
Philanthus cleomae Dunning, 1898. Canad. Ent. 30: 152. 2.
Philanthus annae(!) Dunning, 1898. Canad. Ent. 30: 154.
Philanthus subversus Banks, 1915. Canad. Ent. 47: 405. 2.
Philanthus yakima Banks, 1919. Canad. Ent. 51: 85. °.

Biology: Cazier and Mortenson, 1965. So. Calif. Acad. Sci., Bul. 64: 197-202, fig. 1 (nest, prey,
parasites ?). —Alcock, 1975. Amer. Midland Nat. 93: 222-226, 2 figs. (nesting behavior).
—Aleock, 1975. Anim. Behaviour 23: 889-895, 4 figs. (male territoriality and mating).

nasalis Bohart. Calif. (Contra Costa Co.).
Philanthus nasalis Bohart, 1972. Ent. Soc. Wash., Proc. 74: 401, figs. 5, 6. d, °.

neomexicanus Strandtmann. Calif., Ariz., N. Mex.
Philanthus neomexicanus Strandtmann, 1946. A Rev. of the N. Amer. Spp. of Philanthus,

ps olin dene

occidentalis Strandtmann. Calif.
Philanthus occidentalis Strandtmann, 1946. A Rev. of the N. Amer. Spp. of Philanthus,

p. 64. 3, 2.

1724 Hymenoptera in America North of Mexico

pacificus arizonae Dunning. Wyo. to Ariz. and Calif.; Mexico. Ecology: Nests in fine-grained

alluvial sand, usually in a colony of only a few individuals, makes up to 3 cells per nest,
and stores 8-15 quite small prey per nest. Parasite: Senotainia trilineata (Wulp)?, Prey:
Diplazon laetatorius (F.); Chelonus texanus Cr.; Chrysididae sp.; Stenodynerus
valliceps Boh.; Diodontus ater (Mick.), D. gillettei Fox. D. sp., Passaloecus relativus
Fox, Pseninae sp.; Solierella blaisdelli (Bridw.); Lindenius columbianus (Kohl);
Hylaeus sp.; Perdita fallax Ckll., P. interserta ciliata Timb.; Dialictus incompletus
(Cwfd.), D. laevissimus (Sm.), D. ruidosensis (Ckll.), D. tegulariformis (Cwfd.), D. spp.,
Dufourea scabricornis Boh., Evylaeus niger (Vier.), E. synthridis (Ckll.), E. sp.,
Halictus confusus Sm., H. ligatus Say, H. tripartitus Ckll., Sphecodes spp.

Philanthus arizonae Dunning, 1898. Canad. Ent. 30: 155. d.

Anthophilus hirticeps Cameron, 1905. Amer. Ent. Soc., Trans. 31: 376. d.

Philanthus assimilis Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 422. 9.

Biology: Powell and Chemsak, 1959. Kans. Ent. Soc., Jour. 32: 115-120 (nest, prey; misdet. as
politus pacificus Cr.). —Evans, 1970. Mus. Compar. Zool., Bul. 140: 501 (nest, prey,
parasite ?; misdet. as pacificus Cr.).

pacificus pacificus Cresson. Nev.
Philanthus pacificus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proe., p. xxxii. 6.
politus Say. East. U. S. and southeast. Canada. Ecology: Nests in flat or slightly sloping, bare
sandy soil, makes up to 4 cells per nest, stores 9-18 small bees per cell. Prey: Hylaews
m. modestus Say, H. verticalis (Cr.), H. ziziae (Robt.), H. sp.; Calliopsis andreniformis
Sm.,; Augochlorella aurata (Sm.), A. striata (Prov.), Dialictus albipennis (Robt.), D.
cressoni (Robt.), D. imitatus (Sm.), D. lineatulus (Cwfd.), D. p. pilosus (Sm.), D. spp.,
Evylaeus macoupinensis (Robt.), Halictus c. confusus Sm.
Philanthus politus Say, 1824. In Keating Narr. Long’s 2nd Exped., v. 2, app., p. 343.
Philanthus dubius Cresson, 1865. Ent. Soc. Phila., Proc. 5: 96. ¢.
Philanthus texanus Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 422. 2, d.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 84, fig. 20 (larva).

Biology: Evans and Lin, 1959. Amer. Midland Nat. 17: 116-120, figs. 1, 2 (nest, prey
transport).
psyche Dunning. Iowa to Tex., west to N. Mex., Utah, and Mont., Alta.; Mexico (Chihuahua).
Philanthus psyche Dunning, 1896. Ent. News 7: 287. °.
Philanthus punctinudus Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 144. d.
Philanthus hermosus Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 421. 9.

pulcher Dalla Torre. Southwest. States, Wyo., Calif., Alta. Ecology: Nests in large, dense

aggregations in hard-packed clay-loam, makes up to 6 cells per nest, stores 6-14 small to
medium-sized prey per cell. Parasite: Senotainia trilineata (Wulp), Phrosinella
fulvicornis Allen. Prey: Scelio sp.; Chrysura pacifica (Say), Elampus viridicyaneus
Nort., Hedychridium fletcheri Bod., Holopyga ventralis (Say), Omalus aeneus (F.);
Ancistrocerus c. catskill (Sauss.), Stenodynerus papagorum (Vier.), Symmorphus
canadensis (Sauss.); Diodontus argentinae Roh., D. gillettei Fox, D. sp., Mimesa
unicincta Cr., Mimumesa mixta (Fox), Passaloecus cuspidatus Sm., P. relativus Fox, P.
sp.; Plenoculus davisi Fox, Tachysphex aethiops Cr., T. nigrior Fox, T. tarsatus (Say),
T. sp., Trypoxylon aldrichi Sandh.; Belomicrus forbesii Robt., Crabro florissantensis
Roh., Ectemnius dives (Lep. and Br.), Lindenius columbianus (Kohl), Oxybelus
uniglumis (L.); Dienoplus pictifrons Fox; Colletes nigrifrons Tit., Hylaeus basalis
(Sm.), H. conspicuus (Metz), H. ellipticus (Kby.); Andrena melanochroa Ckll.,
Panurginus atriceps (Cr.), P. cressoniellus Ckll., Perdita wyomingensis Ckll.; Dialictus
laevissimus (Sm.), D. ruidosensis (Ckll.), D. spp., Dufourea maura (Cr.), D. seabricornis
Boh., Evylaeus niger (Vier.), Halictus confusus Sm., H. tripartitus Ckll., Sphecodes spp.;
Formicapis clypeata Slad., Hoplitis producta (Cr.), Osmia pentstemonis Ckll., O. spp.,
Stelis lateralis Cr.; Nomada spp.

Philanthus pulchellus Cresson, 1865. Ent. Soe. Phila., Proc. 5: 93. 3. Preoce.

Philanthus pulcher Dalla Torre, 1897. Cat. Hym., v. 8, p. 489. N. name.

Philanthus clarconis Viereck, 1906. Amer. Ent. Soc., Trans. 32: 206. 9.

Superfamily SPHECOIDEA 1725

Biology: Evans, 1966. Great Basin Nat. 26: 35-38 (nest, prey, parasite). —Evans, 1970. Mus.
Compar. Zool., Bul. 140: 496-498 (nest, prey, parasites).
sanbornii Cresson. Ont., New England States south to N. J. and west to Man. and N. Mex.

Ecology: Nests in small aggregations in flat sand covered with grass. Prey:
Agapostemon radiatus (Say), Lasioglossum forbesii (Robt.); Apis mellifera L.; worker
honeybees are the preferred prey.

Philanthus sanborniui Cresson, 1865. Ent. Soe. Phila., Proc. 5: 89. 6, 2.

Philanthus scutellaris Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxxiv. 6.

Philanthus eurynome Fox, 1890. Ent. News 1: 107. @.

Philanthus trumani Dunning, 1897. Ent. News 8: 70. 2.

Philanthus magdalenae Strandtmann, 1946. A Rev. of the N. Amer. Spp. of Philanthus,
p- 39. 2.

Biology: Evans, 1955. Brooklyn Ent. Soc. Bul. 50: 47 (nest, prey). —Evans and Lin, 1959.
Amer. Midland Nat. 17: 128-129 (nest, prey).
schusteri Bohart. South. Calif.
Philanthus schusteri Bohart, 1972. Ent. Soc. Wash., Proc. 74: 402, fig. 4. d, 2.
serrulatae Dunning. Southwest. States; Mexico (Sonora).
Philanthus serrulatae Dunning, 1898. Canad. Ent. 30: 154. 9.
siouxensis Mickel. Great Plains and Rocky Mt. States; Mexico (Chihuahua, Coahuila).
Philanthus siouxensis Mickel, 1916. Amer. Ent. Soc., Trans. 42: 406. 2.
solivagus Say. Northeast. States south to Va. and west to Wis., southeast. Canada. Ecology:
Nests in flat to slightly sloping sand, makes up to 6 or more cells per nest, stores 6-14
prey per cell. Parasite: Senotainia trilineata (Wulp), Phrosinella fulvicornis Coq.? Prey:
Ancistrocerus c. catskill (Sauss.), A. c. albophaleratus (Sauss.), A. a. adiabatus (Sauss.);
Ectemnius continuus (F.), E. lapidarius (Panz.), Lestica i. interrupta (Lep. and Br.);
Colletes americanus Cr.; Andrena asteris Robt., A. nubecula Sm., A. solidaginis Robt.,
A. subaustralis Ckll., A. spp.; Agapostemon virescens (F.), Augochlora pura (Say),
Augochlorella aurata (Sm.), A. striata (Prov.), Augochloropsis cuprea (Sm.), Dialictus
euryceps (Ellis), D. lineatulus (Cwfd.), D. oblongus (Lov.), D. versans (Lov.), Halictus c.
confusus Sm., H. ligatus Say, H. rubicundus (Chr.), Lasioglossum coriaceum (Sm.), L.
leucozonium (Schr.), Sphecodes davisii Robt.; bees are the preferred prey.
Philanthus solivagus Say, 1837. Boston Jour. Nat. Hist. 1: 383. ¢.
Philanthus solidagus(!) Howard, 1901. Insect Book, pl. 3, fig. 31.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 83, figs. 16-18 (larva).

Biology: Ristich, 1956. Ohio Jour. Sci. 56: 271-274 (parasite). —Evans and Lin, 1959. Amer.
Midland Nat. 17: 120-124, figs. 6-8 (nest, prey, parasite).
tarsatus Smith. Colo., Nebr., Tex.
Philanthus tarsatus Smith, 1908. Nebr. Univ. Studies 8: 356. 2.
ventilabris Fabricius. Generally distributed throughout the U.S. and south. Canada. Ecology:
Nests in flat sand, makes accessory burrow, stores up to 8 bees per cell. Prey: Perdita
albipennis Cr., P. sp.; Dialictus inconspicuus (Sm.), D. microlepoides (Ellis), D.
pruinosus (Robt.), D. versatus (Robt.), D. spp., Halictus ligatus Say, Lasioglossum
sisymbrii Ckll., Nomia nevadensis arizonensis Ckll.
Philanthus ventilabris Fabricius, 1798. Sup. Ent. System., p. 268.
Philanthus vertilabris(!) Fabricius, 1804. Systema Piezatorum, p. 303.
Philanthus frontalis Cresson, 1865. Ent. Soc. Phila., Proc. 5: 99. ¢. Preoce.
Liris rugosa Provancher, 1895. Nat. Canad. 22: 130. d.
Epiphilanthus ventralis(!) Ashmead, 1899. Canad. Ent. 31: 296.
Philanthus ventralis(!) Howard, 1901. Insect Book, pl. 3, fig. 33.
Philanthus vertilabris(!) var. completus Banks, 1915. Canad. Ent. 47: 406. d.

Biology: Peckham and Peckham, 1905. Wasps, social and solitary, pp. 166-167 (nest, prey).
—Rau and Rau, 1918. Wasp studies afield, pp. 116-117 (nest, prey). —Evans and Lin, 1959.
Amer. Midland Nat. 17: 127 (nest, prey). —Alcock, 1975. Kans. Ent. Soc., Jour. 48: 541, figs.
6-7 (male mating strategy). —Aleock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 163-164,
fig. 4 (nest, prey).

1726 Hymenoptera in America North of Mexico

ventralis (Mickel). Pacifie Coast States.
Ococletes ventralis Mickel, 1918 (1917). Nebr. Univ. Studies 17: 328. °.
Philanthus strandtmanni Burks, 1951. In Muesebeck, U.S. Dept. Agr., Agr. Monog. 2:
1002. N. name proposed unnecessarily.
zebratus Cresson. N. Dak., Nebr. and Colo. west to B. C. and Calif. Ecology: Nests in coarse
sandy loam, occasionally makes only one nest during a lifetime which may contain up to
17 cells, stores 3-9 large prey per cell. Parasite: Senotainia trilineata (Wulp),
Phrosinella pilosifrons Allen. Prey: Diphyus sp., Dusona sp., Eutanyacra sp.,
Ichneumon sp., Spilichneumon sp.; Ancistrocerus a. adiabatus (Sauss.), Ewodynerus
castigatus (Sauss.), Stenodynerus taos (Cr.), Symmorphus meridionalis Vier.;
Pseudomasaris vespoides Cr.; Ammophila azteca Cam., A. dysmica Menke, A. mediata
Cr., Palmodes carbo Boh. and Menke, Podalonia communis (Cr.), P. luctuosa (Sm.);
Astata nubecula Cr.; Larropsis capax (Fox), Tachysphex aethiops (Cr.), T. sp.; Crabro
latipes Sm., C. pleuralis Fox, Ectemnius sp.; Aphilanthops subfrigidus Dunn., Cerceris
aequalis idahoensis Seul., Eucerceris fulvipes Cr.
Philanthus zebratus Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxxili. 6.
Philanthus basilaris Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxxiii. d, °.
Oclocletes nitens Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 423. °.
Ococletes(!) illustris Mickel, 1918 (1917). Nebr. Univ. Studies 17: 327. °.

Biology: Evans, 1966. Great Basin Nat. 26: 38-39, fig. 2 (nest, prey, parasites). —Evans, 1970.
Mus. Compar. Zool., Bul. 140: 499-500 (nest, prey, parasites).

Genus TRACHYPUS Klug

Trachypus Klug, 1810. Mag. Gesell. Naturf. Freunde Berlin 4: 41.
Type-species: Trachypus Gomesii Klug. Monotypic.
Simblephilus Dahlbom. 1844. Hym. Europaea, v. 1, p. 190.
Type-species: Philanthus petiolatus Spinola. Monotypic.
Philanthocephalus Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 86.
Type-species: Philanthocephalus gracilis Cameron. Desig. by Pate, 1937.
This small genus is essentially Neotropical, but one Mexican species occurs in southern Texas.
These wasps also prey upon bees.
mexicanus Saussure. South. Tex.; Mexico to Costa Rica. Ecology: Nests in flat sand or
limestone rubble, makes multicellular nest, stores up to 6 prey per cell. Prey:
Augochlora sp., Halictus hesperus Sm.; Exomalopsis sp., Melissodes sp.
Trachypus Mexicanus Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 86. 9.
Philanthocephalus mexicanus Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 89, pl. 6,

figs. 5, 6. 2, d. Preoce.
Trachypus annulitarsis Cameron, 1908. Amer. Ent. Soc., Trans. 34: 232. ?.

Biology: Evans, 1964. Kans. Ent. Soc., Jour. 37: 303-304 (nest, prey).
SuBFAMILY APHILANTHOPSINAE
TRIBE APHILANTHOPSINI

This tribe occurs only in the Nearctic Region. Its members prey only upon ants, the more
primitive Aphilanthops on winged queen ants. The other two genera prey upon harvester ant
workers and have developed a complex mechanism at the apex of the abdomen which is used in
prey transport.

Taxonomy: Bohart, 1966. Ent. Soc. Wash., Proc. 68: 158-167 (review of Nearctic spp.).
Biology: Evans, 1962. Behaviour 19: 239-260, 5 figs. (nesting behavior, prey carriage).
Genus APHILANTHOPS Patton

Aphilanthops Patton, 1881. Boston Soc. Nat. Hist., Proc. 20: 401.
Type-species: Philanthus frigidus Smith. Orig. desig.

Superfamily SPHECOIDEA 1727

Winged queen ants of the genus Formica are used as prey. The wasps dealate the queens be-
fore placing them in the brood cell.

Revision: Dunning, 1898. Amer. Ent. Soc. Trans. 25: 19-26.
foxi Dunning. South. Calif. deserts.
Aphilanthops foxi Dunning, 1898. Amer. Ent. Soc., Trans. 25: 21. 6.
frigidus (Smith). Transcont., N. S., Que., Wyo. and Wash. south to Va., Mich., N. Mex., Utah
and Calif. Ecology: Nests gregariously in pebbly sand, makes up to 4 or more cells per
nest, stores 2-3 dealated queen ants per cell. Parasite: Senotainia trilineata (Wulp),
Euaraba tergata (Cog.)? Prey: Winged queens of Formica fusca L., F. pallidefulva
nitidiventris Em., F. neogagates Em.
Philanthus frigidus Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 475. 3.
Aphilanthops bakeri Dunning, 1896. Canad. Ent. 28: 203. ¢.
Nomada (Holonomada) dawsoni Swenk, 1912. Nebr. Univ. Studies 12: 83. ¢.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 81, figs. 1-10 (larva).

Biology: Peckham and Peckham, 1905. Wasps, social and solitary, pp. 167-177, 1 fig. (nest,
prey). —Wheeler, 1913. Jour. Anim. Behavior 3: 374-387 (nest, prey). —Ristich, 1956. Ohio
Jour. Sci. 56: 271-274 (parasite). —Evans, 1962. Behaviour 19: 240-245, fig. la (nest, prey
carriage, parasite ?).

hispidus Fox. Ariz., south. Calif.; Mexico (Baja California).
Aphilanthops hispidus Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 106. 6.
subfrigidus Dunning. N. Y., Colo., Wyo., Mont., Idaho, B. C., Wash., Oreg., Calif., Nev. Ecology:
Nests in compacted soil. Parasite: Senotainia trilineata (Wulp)?, Metopia argyrocephala
(Meig.)? Prey: Winged queens of Formica neogagates Em., F. fusca L. group. Predator:
Philanthus zebratus Cr.
Aphilanthops subfrigidus Dunning, 1898. Amer. Ent. Soc., Trans. 25: 21. d, @.
Aphilanthops elsiae Dunning, 1898. Amer. Ent. Soc., Trans. 25: 23. 9°.

Biology: Evans, 1962. Behaviour 19: 245 (prey). —Evans, 1970. Mus. Compar. Zool., Bul. 140:
496 (nest, prey, parasites ?).

Genus CLYPEADON Patton

Clypeadon Patton, 1897. Ent. News 8: 13.
Type-species: Aphilanthops quadrinotatus Ashmead. Orig. desig.

The eight known species occur in the western deserts of the Nearctic Region. Worker har-
vester ants of the genus Pogonomyrmex are used as prey, and each species apparently preys
upon only one species of ant. The ant is transported to the nest in a special ant clamp formed by
modification of the apex of the abdomen.

californicus (Bohart). Calif., Oreg.
Aphilanthops (Clypeadon) californica Bohart, 1959. Ent. Soc. Amer., Ann. 52: 108. d, @.
dreisbachi (Bohart). Tex., Okla., Colo.; Mexico (Zacatecas, Jalisco, Nayarit, Queretaro, San Luis
Potosi). Prey: Worker ants, Pogonomyrmex barbatus rugosus Em.
Aphilanthops (Clypeadon) dreisbachi Bohart, 1959. Ent. Soc. Amer., Ann. 52: 107. 6, 2.

Biology: Evans, 1962. Behaviour 19: 249 (prey).
evansi Bohart. N. Mex., Ariz., Calif. Ecology: Nests in sand dune, makes up to 5 or more cells
per nest, stores about 7-12 prey per cell. Parasite: Senotainia trilineata (Wulp). Prey:
Pogonomyrmex barbatus rugosus Em. workers.
Clypeadon evansi Bohart, 1966. Ent. Soc. Wash., Proc. 68: 163. d, 2.

Biology: Evans, 1962. Behaviour 19: 248-249, fig. 2 (nest, prey, parasite; det. as Aphilanthops
(Clypeadon) sp. A). —Aleock and Gamboa, 1975. Ariz. Acad. Sci., Jour. 10: 162-163, fig. 3
(prey capture, nest).

haigi (Bohart). Ariz. to west. Tex. Ecology: Nests in small aggregations in sand dunes, makes
up to 3 cells per nest, stores up to 14 prey per cell. Parasite: Miltogrammini sp.? Prey:
Pogonomyrmex barbatus rugosus Em. workers.
Aphilanthops (Clypeadon) haigi Bohart, 1959. Ent. Soc. Amer., Ann. 52: 106. 6, °.

1728 Hymenoptera in America North of Mexico

Biology: Evans, 1962. Behaviour 19: 245-248, figs. 1b, 2, 3, 4b (nest, prey carriage, parasite ?).
laticinctus (Cresson). West. Tex. to east. and south. Calif., north to Colo., Idaho and Oreg.
Ecology: Nests in bare level sand, makes up to 3 cells per nest, stores 15-26 prey per
cell. Parasite: Senotainia sp. near trilineata (Wulp). Prey: Pogonomyrmex occidentalis

Cr. workers.
Philanthus laticinctus Cresson, 1865. Ent. Soc. Phila., Proc. 5: 91. d.
Aphilanthops quadrinotatus Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 7. 2.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 274-275, figs. 63-64 (larva).

Biology: Hicks, 1927. Canad. Ent. 59: 51-55 (nest, prey carriage, cocoon). —Hicks, 1933.
Canad. Ent. 65: 141-144 (nest, prey). —Evans, 1962. Behaviour 19: 250-253 (nest, prey
carriage, parasite).

sculleni (Bohart). Ariz., Colo., N. Mex., Tex.; Mexico (Chihuahua). Prey: Worker ants,
Pogonomyrmex maricopa barnesi Sm.
Aphilanthops (Clypeadon) sculleni Bohart, 1959. Ent. Soe. Amer., Ann. 52: 107. d, 2.

Biology: Evans, 1962. Behaviour 19: 250 (prey).
taurulus (Cockerell). Ariz., N. Mex., west. Tex. Prey: Workers of Pogonomyrmex barbatus
rugosus Em.
Aphilanthops taurulus Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 293. ¢.
Aphilanthops (Clypeadon) phoenix Pate, 1947. Pan-Pacifie Ent. 23: 66. 2.

Biology: Ainslie, 1909. Canad. Ent. 41: 99-100 (prey capture). —Alcock, 1975. Kans. Ent. Soc.,
Jour. 48: 538-541, fig. 5 (male mating strategy).
utahensis (Baker). Southwest. Tex., N. Mex., Ariz., Nev., Utah and south. Calif. in deserts;
Mexico (Sonora, Queretaro). Prey: Pogonomyrmex barbatus (Sm.) workers.
Aphilanthops utahensis Baker, 1895. Canad. Ent 27: 335. d.
Aphilanthops concinnulus Cockerell, 1896. Canad. Ent. 28: 221. d.

Biology: Evans, 1962. Behaviour 19: 250, fig. 4c (prey).

Genus LISTROPYGIA Bohart

Aphilanthops subg. Listropygia Bohart, 1959. Ent. Soc. Amer., Ann. 52: 106.
Type-species: Aphilanthops (Listropygia) bechteli Bohart. Orig. desig.

Only one species is known in this genus. It has an ant clamp at the apex of the abdomen and
preys upon worker harvester ants.

bechteli (Bohart). South. Calif., Ariz. Prey: Workers of Pogonomyrmex californicus Buck.
Aphilanthops (Listropygia) bechteli Bohart, 1959. Ent. Soc. Amer., Ann. 52: 106. 6, °.

Biology: Evans, 1962. Behaviour 19: 253, figs. 4c, 5 (prey).
SUBFAMILY CERCERINAE

This subfamily contains only two genera, Cerceris and Eucerceris. The former contains nu-
merous species and occurs in all major zoogeographic regions. The latter genus has comparative-
ly few species, occurs in the New World only as far south as Panama, and the majority of species
are Nearctic. So far as known the New World species prey upon Coleoptera only as do most of
the Old World species, but a few of the latter are known to prey upon bees, wasps and chalci-
doids.

Biology: Scullen and Wold, 1969. Ent. Soc. Amer., Ann. 62: 209-214 (prey records of spp. in
America north of Mexico). —Evans, 1971. Kans. Ent. Soc., Jour. 44: 518, 520-522 (notes on
comparative behavior).

Genus CERCERIS Latreille

Cerceris Latreille, 1802-1803. Hist. Nat. Crust. Ins., v. 3, p. 367.
Type-species: Philanthus ornatus Fabricius. Desig. by Latreille, 1810 (~Sphex
rybyensis Linnaeus).
Nectanebus Spinola, 1838. Soc. Ent. France, Ann. 7: 489.

Superfamily SPHECOIDEA 1729

Type-species: Nectanebus Fischeri Spinola. Orig. desig.
Diamma Dahlbom, 1844. Hym. Europaea. v. 1, p. 225. Preoce.
Type-species: Diamma Spinolae Dahlbom. Monotypic (=Cerceris binodis Spinola).
Didesmus Dahlbom, 1845. Hym. Europaea, v. 1, p. 502. N. name.
Apiraptrix Shestakov, 1923. Sbornik Jaroslav Gosudarst. Universitet., p. 101.
Type-species: Sphex rybyensis Linnaeus. Orig. desig.
Paracerceris Brethes, 1913. Mus. Nac. Hist. Nat. Buenos Aires, An. 24: 127.
Type-species: Paracerceris tridentifera Brethes. Monotypic.
Bucerceris Minkiewicz, 1934. Polski Pismo Ent. 12: 253.
Type-species: Cerceris bupresticida Dufour. Monotypic.
Stercobata Gussakovskij, 1935. Acad. Sci. U. R. S.S., Tadjikistan, Trav. 5: 445.
Type-species: Cerceris bupresticida Dufour. Monotypic.
Apicerceris Pate, 1937. Amer. Ent. Soc., Mem. 9: 8.
Type-species: Sphex rybyensis Linnaeus. Orig. desig.

Revision: Scullen, 1965. U. S. Natl. Mus., Proc. 116: 333-548, 182 figs. (spp. of America north
of Mexico). —Scullen, 1972. Smithsn. Contrib. Zool. 110: 1-121, 173 figs. (Mexican and
Central American spp.).

Taxonomy: Cresson, 1865. Ent. Soc. Phila., Proc. 5: 122-132 (spp. of N. Amer. north of
Mexico). —Banks, 1912. Ent. Soc. Amer., Ann. 5: 11-30 (spp. of east. N. Amer.). —Banks,
1947. Psyche 54: 1-35 (West Coast spp.).

acanthophila Cockerell. Southwest. U. S.; Mexico (Baja California to Nuevo Leon, south to
Hidalgo).
Cerceris acanthophila Cockerell, 1897. Entomologist 30: 135. d.
Cerceris cockerelli Viereck, 1903 (1902). Acad. Nat. Sci. Phila., Proc. 54: 731. 6.
Cerceris (Apiratrix(!)) huachuca Banks, 1947. Psyche 54: 29. 6.
aequalis aequalis Provancher. Principally Calif. and Oreg., but scattered records in Wash.,
Wyo., Utah, Ariz., N. Mex., Tex.
Cerceris aequalis Provancher, 1888. Addit. Corr. Faune Ent. Canada Hym., p. 417. 2.
Cerceris vicinoides Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 140. “3” = 9.
Cerceris psamathe Banks, 1912. Ent. Soc. Amer., Ann. 5: 21. °.
aequalis bolingeri Scullen. Oreg., Nev.
Cerceris aequalis bolingeri Scullen, 1965. U. S. Natl. Mus., Proc. 116: 453, figs. 64, 146a, b,

@ ee
aequalis idahoensis Scullen. B. C. to Oreg., Idaho, Wyo., Colo. Predator: Philanthus zebratus
Cr.
Cerceris aequalis idahoensis Scullen, 1965. U. S. Natl. Mus., Proc. 116: 454, figs. 65, 147a,
b,c. @.

alceste Mickel. Nebr. (Mitchell).
Cerceris alceste Mickel, 1918 (1917). Nebr. Univ. Studies 17: 333. °.
argia Mickel. La., Tex. north to Kans. and Colo., west to Ariz.; Mexico (Chihuahua, Nuevo

Leon, Jalisco).
Cerceris argia Mickel, 1916. Amer. Ent. Soc., Trans. 42: 412. °.

Taxonomy: Scullen, 1968. Ent. News 79: 158, figs. 5, 6. d.
arizonella Banks. Ariz. (Tempe).
Cerceris arizonella Banks, 1947. Psyche 54: 32. 6.
astarte Banks. N. H. to N. C. west to Wis., Ill. and northeast. Tex.
Cerceris astarte Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 424. 9.
atramontensis Banks. Que. to N. C. west to N. Dak. and centr. Tex. Ecology: Nests in woods
in stony soil. Prey: Conotrachelus anaglypticus Say, C. naso LeC., C. nenuphar Hbst.,
C. posticatus Boh., Hyperodes sparsus Say.
Cerceris atramontensis Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 425. 9.
Cerceris arbuscula Mickel, 1916. Amer. Ent. Soc., Trans. 42: 410. °.

Biology: Krombein, 1956. Brooklyn Ent. Soc., Bul. 51: 43 (nest, prey). —Evans, 1971. Kans.
Ent. Soc., Jour. 44: 512-513 (nest, prey).

1730 Hymenoptera in America North of Mexico

azteca Saussure. Ariz.; Mexico to Argentina.
Cerceris azteca Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 97. 2.
Cerceris (Apiratrix(!)) seminigra Banks, 1947. Psyche 54: 33. d. Preoce.
banksi Scullen. Mass. to N. C., Ill., Mo., Kans.
Cerceris banksi Scullen, 1965. U. S. Natl. Mus., Proc. 116: 461, figs. 70, 152a, b,c. 2.
bicornuta Guerin. Transcont. in U. S., Mass., Mich., S. Dak. and Oreg. south into Mexico
(Coahuila, Chihuahua, Sonora, Baja California, Durango). Ecology: Nests in flat stony
soil or sand, makes its own burrow or extends a pre-existing burrow. Parasite:
Senotainia kansensis Tns.?; Hedychrum violaceum Br.? Prey: Eupagoderes sp.,
Sphenophorus aequalis Gyll., S. criosus (Oliv.), S. cultellatus (Horn), S. maidis (Chitt.),
S. marinus (Chitt.), S. parvulus (Gyll.), S. pertinax (Oliv.), S. venatus (Say), S. vestitus
(Chitt.), S. zeae (Walsh).
Cerceris bicornuta Guerin, 1844. Iconogr. Regne Anim., Ins., v. 7, p. 448. @.
Cerceris dufourti Guerin, 1844. Iconogr. Regne Anim., Ins., v. 7, p. 444. 3.
Cerceris venator Cresson, 1865. Ent. Soc. Phila., Proc. 5: 116. ¢.
Cerceris curvicornis Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 124, pl. 8, fig. 3. ¢.
Cerceris fidelis Viereck and Cockerell, 1904. N. Y. Ent. Soe., Jour. 12: 132. @.
Cerceris venatrix Schulz, 1906. Spolia Hym., p. 195. Emend.

Biology: Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 337-341, figs. 35, 36 (nest, prey, parasite
?). —Cartwright, 1929. S. C. Agr. Expt. Sta., Bul. 257: 31 (prey). —Strandtmann, 1945. Ent.
Soe. Amer., Ann. 38: 311, fig. 8 (nest, prey). —Krombein, 1953. Ent. Soc. Wash., Proc. 55:
118-119 (nest, prey). —Krombein, 1960. Brooklyn Ent. Soc., Bul. 55: 75-76 (nest, prey,
parasite ?). —Evans, 1971. Kans. Ent. Soc., Jour. 44: 513 (prey).

blakei Cresson. N. J. to Fla., west to east. Tex., Ill., Kans. Ecology: Makes vertical burrow in

flat bare sand, makes 3 or more cells in a completed nest, stores 42-69 very small
beetles, mostly weevils, per cell. Parasite: Miltogrammini sp. Prey: Derolomus basalis
LeC. the most common prey, Limnobaris confusa (Boh.), Anthonomus sexguttatus
Dietz, Hyperodes sp.; Graphops floridana Bl.; Blapstinus interruptus (Say).

Cerceris elegans Smith, 1856. Cat. Hym. Brit. Mus., v. 5, p. 467. d. Preoce.

Cerceris blakei Cresson, 1865. Ent. Soc. Phila., Proc. 5: 121. @.

Cerceris elegantissima Schletterer, 1887. Zool. Jahrb., Ztschr. f. System. 2: 490. N. name.

Biology: Krombein, 1963. Brooklyn Ent. Soc., Bul. 58: 72-79 (nest, prey capture and

transport, parasite).
boharti Scullen. Ariz., N. Mex.; Mexico (Coahuila).

‘Cerceris boharti Scullen, 1965. U. S. Natl. Mus., Proc. 116: 466, figs. 73, 154a, b,c. ¢.
bolingeriana Krombein, n. name. Ariz. (Herford); Mexico (Durango).

Cerceris bolingeri Scullen, 1972. Smithsn. Contrib. Zool. 110: 72, figs. 68, 146a, b. 2. Preoce.
bridwelli Scullen. South. Calif. and Ariz.; Mexico (Baja California, Sonora).

Cerceris bridwelli Seullen, 1965. U. S. Natl. Mus., Proc. 116: 361, figs. 5, 11la, b,c. 2, d.
butleri Scullen. South. Ariz., southwest. Tex.; Mexico (Sonora).

Cerceris butleri Scullen, 1965. U. S. Natl. Mus., Proce. 116: 363, figs. 6, 112a, b, c. °.

californica Cresson. B. C. and Idaho south to Calif. and Ariz., N. Mex., west. Tex.; Mexico

(Baja California to Nuevo Leon south to Queretaro). Parasite: Dasymutilla
coccineohirta (Bl.); Metopia argyrocephala (Meig.), Amobia floridensis (Tns.) ?,
Senotainia trilineata (Wulp) ? Prey: Acmaeodera acuta LeC., A. adenostomae Caz., A.
angelica Fall, A. coquilletti Fall, A. dohrni Horn, A. dolorosa Fall, A. fenyesi Fall, A.
gemina Horn, A. hepburni LeC., A. holsteni White, A. jocosa Fall, A. nexa Fall, A.
perlanosa Timb., A. plagiaticauda Horn, A. prorsa Fall, A. quadriseriata Fall, A.
sinuata Van Dyke, Agrilus angelicus Horn, A. blandus Horn, A. politus (Say),
Anthaxia aeneogaster Cast., Chrysobothris deleta LeC., C. lucana Horn, C. femorata
(Oliv.).

Cerceris californica Cresson, 1865. Ent. Soe. Phila., Proc. 5: 128. d.

Cerceris ferruginior Viereck and Cockerell, 1904. N. Y. Ent. Soe., Jour. 12: 134. 6.

Cerceris populorum Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 135. 3.

Cerceris garciana Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 135. 6.

Cerceris argyrotricha Rohwer, 1908. Canad. Ent. 40: 324. 9.

Superfamily SPHECOIDEA 1731

Cerceris cognata Mickel, 1916. Amer. Ent. Soc., Trans. 42: 408. °.
Cerceris denticularis Banks, 1917. Mus. Comp. Zool., Bul. 61: 113. 2, d.
Cerceris interjecta Banks, 1919. Canad. Ent. 51: 84. 6.

Cerceris arno Banks, 1947. Psyche 54: 19. 2.

Cerceris calodera Banks, 1947. Psyche 54: 22. 6.

Cerceris illota Banks, 1947. Psyche 54: 28. 6.

Cerceris isolde Banks, 1947. Psyche 54: 24. 6.

Biology: Linsley and MacSwain, 1956. Ent. Soc. Amer., Ann. 49: 71-84, 3 figs. (nest, prey
selection, parasites).
calochorti calochorti Rohwer. East slope of Rocky Mts., Alta. south to N. Mex. and Tex.
Predator: Philanthus zebratus Cr.
Cerceris calochorti Rohwer, 1908. Canad. Ent. 40: 322. 9°.
calochorti hidalgo Scullen. Ariz. (Pima Co.); Mexico (Higher elevations south to Mexico City).
Cerceris calochorti hidalgo Seullen, 1972. Smithsn. Contrib. Zool. 110: 75, figs. 71, 149a-e.
OPEOr
carrizonensis Banks. Tex., (Uvalde).
Cerceris carrizonensis Banks, 1915. Canad. Ent. 47: 403. do.
chilopsidis Viereck and Cockerell. Southwest. desert area.
Cerceris chilopsidis Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 136. °.
cleomae Rohwer. Colo. (Denver).
Cerceris cleomae Rohwer, 1908. Canad. Ent. 40: 325. 6.
clypeata clypeata Dahlbom. Maine and Ont. to N. Dak. south to north. Fla. and east Tex.
Ecology: Nests in gravelly soil. Parasite: Dasymutilla scaevola (Bl.) ? Prey: Chalepus
dorsalis Thunb., Lema trilineata (Oliv.); Curculio nasicus (Say), Pissodes strobi (Peck),
Tanymecus confusus (Say).
Cerceris clypeata Dahlbom, 1844. Hym. Europaea, v. 1, pp. 221, 500. d, 2.
Cerceris imitator Cresson, 1865. Ent. Soe. Phila., Proe. 5: 125. d. Preoce.
Cerceris imitatoria Schletterer, 1887. Zool. Jahrb., Ztschr. f. System. 2: 494. N. name.
Cerceris chryssipe Banks, 1912. Ent. Soc. Amer., Ann. 5: 18. 2, ¢.
Cerceris clymene Banks, 1912. Ent. Soc. Amer., Ann. 5: 20. d, 9.
Cerceris zobeide Brimley, 1929. Ent. News 40: 194. 6.
Cerceris zosma Brimley, 1929. Ent. News 40: 195. d.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 86, fig. 31 (larva).

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Surv., Bul. 2: 109-114 (nest,

prey). —Krombein, 1954. Brooklyn Ent. Soc., Bul. 49: 6-7 (nest, prey). —Scullen, 1965. U.
S. Natl. Mus., Proc. 116: 471 (prey).

clypeata dakotensis Banks. North cent. States.
Cerceris dakotensis Banks, 1915. Canad. Ent. 47: 402. 2, 6.

clypeata gnarina Banks. Cent. States.
Cerceris gnarina Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 237. 2.

clypeata prominens Banks. East. States.
Cerceris prominens Banks, 1912. Ent. Soc. Amer., Ann. 5: 19. °, ¢.
Cerceris alaope Banks, 1912. Ent. Soc. Amer., Ann. 5: 22. ¢, 2.

cochisi Scullen. South. Calif., Ariz., N. Mex., Tex.; Mexico (Baja California, Sonora, Zacatecas,

San Luis Potosi, Nayarit).

Cerceris cochisi Seullen, 1965. U. S. Natl. Mus., Proc. 116: 416, fig. 39. 2, ¢.
Cerceris cochise (!) Secullen, 1972. Smithsn. Contrib. Zool. 110: 42.

compacta Cresson. East. and cent. States southwest to south. Calif.; Mexico south to Veracruz

and Oaxaca. Prey: Colaspis brunnea (F.).

Cerceris compacta Cresson, 1865. Ent. Soc. Phila., Proc. 5: 127. 6, °.
Cerceris aureo-facialis Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 112. ¢.
Cerceris solidaginis Rohwer, 1908. Canad. Ent. 40: 323. 6.
Cerceris belfragei Banks, 1917. Mus. Compar. Zool., Bul. 61: 114. 2, 6.

Biology: Scullen, 1965. U. S. Natl. Mus., Proc. 116: 420 (prey).

1732 Hymenoptera in America North of Mexico

compar albinota Scullen. Southwest. Tex., south. Ariz.; Mexico (Chihuahua, Durango).
Cerceris compar albinota Scullen, 1972. Smithsn. Contrib. Zool. 110: 60, figs. 53, 137a-e. 2,
3.
compar compar Cresson. Maine to S. Dak., south to Fla. and N. Mex.; Mexico (Guanajuato).
Cerceris compar Cresson, 1865. Ent. Soc. Phila., Proc. 5: 126. d.
Cerceris jucunda Cresson, 1872. Amer. Ent. Soc., Trans. 4: 231. d.
Cerceris catawba Banks, 1912. Ent. Soc. Amer., Ann. 5: 25. 2, 6.
Cerceris jucunda var. carolina Banks, 1912. Ent. Soe. Amer., Ann. 5: 26. d.
compar geniculata Cameron. South. Ariz. to north. Guatemala.
Cerceris geniculata Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 113, pl. 7, fig. 7. 2.
Cerceris feralis Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 113. ¢.
compar orestes Banks. South. Ariz. to cent. Mexico.
Cerceris orestes Banks, 1947. Psyche 54: 13. @, 6.
completa Banks. Calif. and south. Oreg.
Cerceris completa Banks, 1919. Canad. Ent. 51: 83. d.
Cerceris grandis percna Scullen, 1965. U. S. Natl Mus., Proce. 116: 415, fig. 38. 2.
conifrons Mickel. Alta., Nebr., Wyo., Utah, Nev., Calif. south to Ariz. and Tex.; Mexico (south
to Zacatecas at higher elevations).
Cerceris conifrons Mickel, 1916. Amer. Ent. Soc., Trans. 42: 410. °.
convergens Viereck and Cockerell. B. C., Wash., Mont. and S. Dak. south to Calif. and Tex.;
Mexico south to Morelos, Veracruz and Baja California.
Cerceris convergens Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 136. °.
Cerceris rinconis Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 137. °.
Cerceris hesperina Banks, 1917. Mus. Compar. Zool., Bul. 61: 115. 2, d.
Cerceris pudorosa Mickel, 1918 (1917). Nebr. Univ. Studies 17: 338. 2, d.
Cerceris snowi Banks, 1919. Canad. Ent. 51: 84. 6.
crandalli Scullen. South. Ariz. to west. Tex.; Mexico (Baja California, Sinaloa, Chihuahua).
Cerceris crandalli Scullen, 1965. U. S. Natl. Mus., Proc. 116: 372, figs. 13, 116a, b, ce. 2, ¢.

Taxonomy: Scullen, 1968. Ent. News 79: 158, figs. 7, 8. 6.
crotonella Viereck and Cockerell. N. Dak., Idaho and north. Calif. south to Ariz. and Tex.;
Mexico (Chihuahua, Sonora). Prey: Carpophilus pallipennis (Say).
Cerceris crotonella Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 139. 9.

Biology: Evans, 1971. Kans. Ent. Soc., Jour. 44: 510-511 (prey).
dentifrons Cresson. Maine to N. C., Ont., Mich., Wis., Minn., Iowa, S. Dak.
Cerceris dentifrons Cresson, 1865. Ent. Soc. Phila., Proc. 5: 124. 9.
deserta Say. Northeast. and centr. States and Provinces, N. B. and Maine west to Alta. and
Mont. south to N. C. and Colo. Prey: Conotrachelus posticatus Boh.
Cerceris deserta Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, app., p. 343. d.
Cerceris fulvipes Cresson, 1865. Ent. Soe. Phila., Proc. 5: 126. 2. Preoce.
Cerceris fulvipediculata Sehletterer, 1887. Zool. Jahrb., Ztschr. f. System. 2: 492. 2. N.
name.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Survey, Bul. 2: 115-116 (nest,
prey transport).
dilatata chisosensis Scullen. South. Ariz., southwest. Tex.; Mexico (Chihuahua, Coahuila,

Durango, Zacatecas).
Cerceris dilatata chisosensis Seullen, 1965. U. S. Natl. Mus., Proc. 116: 409, fig. 35. 2, 6.

dilatata dilatata Spinola. Ariz. to Argentina.
Cerceris dilatata Spinola, 1841. Soc. Ent. France, Ann. 10: 118. d.
Cerceris maximiliani Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 94. °.
Cerceris contracta Taschenberg, 1875. Ztschr. Gesam. Naturw. 45: 396. °.
Cerceris Caridei Holmberg, 1903. Mus. Nac. Buenos Aires, An. 9 (3): 478. °.
Cerceris Vigilii Brethes, 1910. Mus. Nac. Buenos Aires, An. 20 (3): 270. 2.
Cerceris divisa Brethes, 1910. Mus. Nac. Buenos Aires, An. 20 (3): 270. d.
Cerceris olymponis Strand, 1910. Zool. Jahrb. 29: 140. @.
Cerceris semiatra Banks, 1947. Psyche 54: 25. 6.

Superfamily SPHECOIDEA 1733

echo atrata Scullen. Maine, Mass., N. Y., N. J., Va. N. C., Ga., Wis., Ill, Kans. Ecology: Makes
multicellular nest in flat, fine-grained sand, stores up <o 30 prey per cell. Prey: Olibrus

neglectus Casey, O. sp., Phalacrus sp.
Cerceris echo atrata Scullen, 1965. U.S. Natl. Mus., Proc. 116: 377, fig. 16. 2.

Biology: Evans, 1971. Kans. Ent. Soc., Jour. 44: 509, figs. 6-7 (nest, prey).
echo echo Mickel. Iowa, N. Dak. to Tex., west to Idaho, north. Calif. and Ariz.; Mexico
(Chihuahua, Coahuila, Durango, Zacatecas, Aguascalientes). Ecology: Nests in open sand.
Prey: Phalacrus sp.
Cerceris echo Mickel, 1916. Amer. Ent. Soc., Trans. 42: 412. 9.

Biology: Evans, 1971. Kans. Ent. Soc., Jour. 44: 509-510 (nest, prey).
femurrubrum Viereck and Cockerell. Colo. and west. Tex. to south. Calif.; Mexico (Baja

California and Coahuila south to Puebla). Ecology: Nests in moist sand. Prey:
Eurymetopon rufipes Esch., Steriphanus sp.

Cerceris femur-rubrum Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 135. 6.

Cerceris thione Banks, 1947. Psyche 54: 18. 6.

Cerceris athene Banks, 1947. Psyche 54: 20. °.

Cerceris femurrubra rossi Scullen, 1972. Smithsn. Contrib. Zool. 110: 56, fig. 50. 2, d.

Biology: Wasbauer, 1957. Pan-Pacifie Ent. 33: 131 (prey transport). —Aleock, 1974. Jour. Nat.
Hist. 8: 650-651 (nest, prey).
finitima Cresson. Sask., transcont. in U.S., N. Y. to Fla. west to N. Dak., Wyo., Utah and

Calif. south to Guatemala. Parasite: Dasymutilla scaevola (Bl.)? Prey: Chaetocnema
pulicaria Melsh.

Cerceris finitima Cresson, 1865. Ent. Soe. Phila., Proc. 5: 122. 9.

Cerceris finitima var. nigroris Banks, 1912. Ent. Soc. Amer., Ann. 5: 27. 6.

Cerceris (Apiratrix(!)) vierecki Banks, 1947. Psyche 54: 30. °.

Cerceris finitima citrina Scullen, 1965. U. S. Natl. Mus., Proc. 116: 380, fig. 18. 2, ¢.

Cerceris finitima morelos Scullen, 1972. Smithsn. Contrib. Zool. 110: 25, fig. 13. 2, d.

Biology: Strandtmann, 1945. Ent. Soc. Amer., Ann. 38: 312, fig. 10 (nest, prey).
flavofasciata flavofasciata Smith. N. J. to N. C., Ill, Minn. and S. Dak. to Okla. Ecology:

Nests in vertical sand banks. Parasite: Dasymutilla nigripes (F.). Prey: Chlamisus sp.
probably plicata (F.), Cryptocephalus mutabilis Melsh., C. notatus F., C. fulvipennis
Hald., C. guttulatus Oliv., C. sp. probably quadrimaculatus Say, Bassareus clathratus
Melsh., B. sp. probably sellatus Suffr.

Cerceris flavofasciata Smith, 1908. Nebr. Univ. Studies 8: 364. 2.

Cerceris natallenus Brimley, 1927. Ent. News 38: 238. °.

Biology: Krombein, 1959. Ent. Soc. Wash., Proc. 61: 197-198 (nest, prey, cocoon, parasite).
—Scullen, 1965. U. S. Natl. Mus., Proc. 116: 422 (prey).
flavofasciata floridensis Banks. N. C. to Fla., Ala. Ecology: Nests in vertical face of sand pit,
makes more than 2 cells per nest, stores 9-11 prey per cell. Prey: Cryptocephalus bivius
Newm., C. binominis Newm., C. guttulatus Suffr., Chlamisus sp. probably nodulosa
Blatch., Coscinoptera dominicana (F.).
Cerceris floridensis Banks, 1915. Canad. Ent. 47: 403. ¢.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 276, figs. 68-70 (larva).

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 20-22 (nest, prey, life cycle).
frontata Say. Ill. and Iowa west to Calif., south to Tex. and Ariz.; Mexico (Baja California,

Chihuahua, Coahuila, Durango). Ecology: Nests in flat, hard-packed sometimes stony
soil, makes 3 or more cells per nest, stores 6 prey per cell. Parasite: Parametopia sp.?;
Exoprosopa fasciata Macq.; Hedychrum violaceum Br.? Prey: Cleonus pulvereus (LeC.),
Eupagoderes sp., Lixus concavus Say, L. mucidus LeC., Thecesternus humeralis (Say).

Cerceris frontata Say, 1823. West. Quart. Rptr. 2: 80. 2.

Cerceis Texensis Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 89. @.

Cerceris occidentalis Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 100. 9.

Cerceris raui Rohwer, 1920. U. S. Natl. Mus., Proc. 57: 230. 2, do.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 277-278 (larva).

1734 Hymenoptera in America North of Mexico

Biology: Rau, 1928. Acad. Sci. St. Louis, Trans. 25: 325-337, figs. 30-34 (nest, prey, parasites
?). —Krombein, 1960. Brooklyn Ent. Soe., Bul. 55: 76-77 (nest, prey). —Scullen, 1965. U.S.
Natl. Mus., Proc. 116: 482, pl. 1 (prey, mating).

fumipennis Say. U.S. east of Rocky Mts., Wyo., Minn., Mich. and N. H. south to N. Mex. and

Fla. Ecology: Nests in hard-packed sand. Parasite: Vidia (Crabrovidia) sp.; Phrosinella
fumosa Allen, Senotainia trilineata (Wulp)? Prey: Acmaeodera pulchella (Herbst),
Actenodes acornis (Say), Agrilus abjectus Horn, A. anxius Gory, A. a. arcuatus (Say),
A. a. fulgens LeC., A. a. torquatus LeC., A. b. bilineatus (Weber), A. b. carpini Knull, A.
obsoletoguttatus Gory, A. politus Say, A. ruficollis (F.), A. spp., Brachus ovatus Weber,
Buprestis consularis Gory, B. fasciata F., B. intricata Casey, B. lineata F., B.
maculipennis Gory, B. rufipes (Oliv.), B. striata F., Chrysobothris adelpha Germ., C.
azurea LeC., C. blanchardi Horn, C. breviloba Fall, C. dentipes (Germ.), C. femorata
(Oliv.), C. floricola Gory, C. monticola Fall, C. purpureovittata Horn, C. sexsignata
(Say), C. trinervia (Kby.), C. verdigripennis Frost, Cinyra gracilipes (Melsh.), Dicerca
americana (Hbst.), D. caudata LeC., D. divaricata (Say), D. lurida (F.), D. obscura (F.),
D. punctulata (Schon.), D. spreta (Gory), D. tuberculata (Cast.), Eupristocerus cogitans
(Weber), Melanophila fulvoguttata (Harr.), Poecilonotus cyanipes (Say); Chlamisus sp.;
Conotrachelus sp.

Cerceris fumipennis Say, 1837. Boston Jour. Nat. Hist. 1: 381. 6.

Cerceris cincta Dahlbom, 1844. Hym. Europaea, v. 1, p. 204. 6.

Cerceris unicincta Taschenberg, 1875. Ztschr. Gesam. Naturw. Berlin 11: 397. °.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 84, figs. 21-26 (larva).

Biology: Peckham and Peckham, 1900. Wis. Nat. Hist. Soc., Bul. 1: 90 (nest, prey transport).
—Hartman, 1905. Tex. Univ. Bul. 65: 66 (prey). —Grossbeck, 1912. N. Y. Ent. Soc., Jour.
20: 1385, 299. —Cartwright, 1931. Ent. News 42: 269 (nest, prey). —Krombein, 1958. Ent.
Soe. Wash., Proc. 60: 110 (nest, prey). —Scullen, 1965. U. S. Natl. Mus., Proc. 116: 412-413
(prey). —Evans, 1971. Kans. Ent. Soc., Jour. 44: 501-508, figs. 1-5 (mating, nest, prey,
cocoon, parasites).

Morphology: Snodgrass, 1941. Smithsn. Inst., Mise. Collect. 99 (14): pl. 21, figs. A-E (male
genitalia).
gandarai Rohwer. Tex. (Brooks Co.); Mexico (Coahuila and Sinaloa south to Puebla and
Veracruz).
Cerceris gandarai Rohwer, 1912. U.S. Natl. Mus., Proc. 41: 470, fig. 9. 3.
Cerceris gandari(!) Seullen, 1972. Smithsn. Contrib. Zool. 110: 82. 2, d.
grandis Banks. South. Calif. and Ariz.; Mexico (Baja California, Sonora).
Cerceris grandis Banks, 1913. Amer. Mus. Nat. Hist., Bul. 32: 423. °.
graphica Smith. IIl., Nebr. to Tex. west to Utah, south. Calif. and south to northwest. South
America. Ecology: Makes vertical burrow and multicellular nest in soil ranging from
heavy to wind-blown sand, stores 3-10 prey per cell. Parasite: Senotainia trilineata
(Wulp) complex?; Nysson sp.? Prey: Eleodes opaca (Say), Metapoloba pruinosa (Horn),
Tenebrionidae sp.
Cerceris graphica Smith, 1873. Ann. and Mag. Nat. Hist. (4) 12: 410. °.
Cerceris hebes Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 124, pl. 8, fig. 4. d.
Cerceris macrosticta Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 133. 6.
Cerceris ampla Banks, 1912. Ent. Soc. Amer., Ann. 5: 16. 2, d.

Biology: Scullen, 1965. U. S. Natl. Mus., Proc. 116: 440 (prey). —Lin, 1967. Ent. Soc. Wash.,

Proc. 69: 312-317, 3 figs. (nest, prey transport). —Alcock, 1974. Jour. Nat. Hist. 8: 645-650,

7 figs. (nest, prey transport, parasites ?). —Alcock and Gamboa, 1975 (1974). Psyche 81:

528-533, 2 figs. (home ranges of males).

halone Banks. Man., N. Dak. east to N. H. south to Tex., Ark. and S. C. Parasite: Hedychrum

confusum Buyss. Prey: Curculio confusor Ham., C. nasicus (Say), C. proboscideus F., C.
rectus Say, C. sulcatulus (Casey).

Cerceris halone Banks, 1912. Ent. Soc. Amer., Ann. 5: 24. 2.

Cerceris architis Mickel, 1916. Amer. Ent. Soc., Trans. 42: 409. 9.

Cerceris alacris Mickel, 1918 (1917). Nebr. Univ. Studies 17: 334. 9.

Cerceris salome Banks, 1923. Canad. Ent. 55: 21. °.

Superfamily SPHECOIDEA 1735

Cerceris shermani Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 200. 9.

Biology: Abbott, 1928. Ent. News 39: 205-206 (nesting habits). —Krombein, 1958. U.S. Dept.
Agr., Monog. 2, Sup. 1, p. 197 (prey). —Byers, 1962. Kans. Ent. Soce., Jour. 35: 317-321
(nest, prey carriage, parasite (misdet. as H. violacewm Br.)). —Scullen, 1965. U.S. Natl.
Mus., Proce. 116: 484 (prey). —Evans, 1971. Kans. Ent. Soc., Jour. 44: 511-512, fig. 9 (nest,
prey).

hurdi Scullen. Ariz. (Patagonia) south through Mexico to Nicaragua.
Cerceris hurdi Seullen, 1972. Smithsn. Contrib. Zool. 110: 48, figs. 40, 128. 2, 6.
insolita albida Scullen. Ariz., N. Mex., west. Tex.; Mexico (Veracruz).
Cerceris insolita albida Seullen, 1965. U. S. Natl. Mus., Proc. 116: 447, fig. 59. 2, ¢.
insolita atrafemori Scullen. Ariz.; Mexico (Sinaloa, Nayarit).
Cerceris insolita atrafemori Scullen, 1965. U. S. Natl. Mus., Proc. 116: 448, figs. 6, 143a, b,
Q.
insolita insolita Cresson. N. J. to S. Dak. south to Fla. and Ariz.; Mexico (Chihuahua, Jalisco).
Ecology: Nests in flat, coarse-grained sand. Prey: Rhabdopterus praetextus (Say).
Cerceris insolita Cresson, 1865. Ent. Soc. Phila., Proc. 5: 129. 6.
Cerceris intractibilis Mickel, 1916. Amer. Ent. Soc., Trans. 42: 411. 2.

Biology: Krombein, 1964. Brooklyn Ent. Soc., Bul. 58: 119 (nest, prey).

irene Banks. Minn., Wyo., Colo., Kans., Okla., Tex., La.; Mexico (Tamaulipas).
Cerceris irene Banks, 1912. Ent. Soe. Amer., Ann. 5: 26. °.

kennicottii kennicottii Cresson. Md. to Fla. west to S. Dak., Colo., Tex.; Mexico (Jalisco,
Morelos, Veracruz).
Cerceris kennicottii Cresson, 1865. Ent. Soc. Phila., Proc. 5: 128. 3.
Cerceris eriogoni Viereck and Cockerell, 1904. N. Y. Ent. Soe., Jour. 12: 139. 3.

kennicottii zapoteca Saussure. South. Ariz. to west. Tex. south to Panama.

Cerceris Zapoteca Saussure, 1867. Reise d. novara, Zool., v. 2, Hym., p. 89. 6.

Cerceris montivaga Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 119, pl. 7, fig. 16. 2.

Cerceris kennicottii beali Seullen, 1965. U. S. Natl. Mus., Proce. 116: 386, fig. 22. 2, 3.
krombeini Scullen. South. Ariz.

Cerceris krombeini Scullen, 1965. U. S. Natl. Mus., Proc. 116: 388, figs. 23, 122a, b,c. 2, d.
macswaini Scullen. South. Calif. and Ariz.

Cerceris macswaini Scullen, 1965. U.S. Natl. Mus., Proc. 116: 485, figs. 84, 164a, b,c. 2, 6.
mandibularis Patton. N. Y. to Ga., W. Va. to Iowa south to Tenn. and La.

Cerceris mandibularis Patton, 1881. Boston Soc. Nat. Hist., Proce. 20: 403. 2, 6.
melanthe Banks. Que. to N. C. west to IIl., southwest to south. Ariz.

Cerceris melanthe Banks, 1947. Psyche 54: 21. 9.

Cerceris nitida Banks, 19138. Amer. Mus. Nat. Hist., Bul. 32: 424. 2, 3. Preoce.
mimica Cresson. Fla. to Ariz., Kans.; Mexico-(Nuevo Leon and Chihuahua south to Oaxaca).

Cerceris mimica Cresson, 1872. Amer. Ent. Soc., Trans. 4: 228. 2, ¢.

Cerceris esau Schletterer, 1887. Zool. Jahrb., Ztschr. f. System. 2: 458. 9.

Cerceris minima(!) Schletterer, 1887. Zool. Jahrb., Ztschr. f. System. 2: 497.

Cerceris englehardti Banks, 1947. Psyche 54: 12. 6.
minax Mickel. West. U. S.

Cerceris minax Mickel, 1918 (1917). Nebr. Univ. Studies 17: 339. 2, 6.
morata Cresson. Nebr. to Tex. west to Utah and Ariz., south in Mexico to Oaxaca. Ecology:

Makes multicellular nest in heavy soil, stores 59-67 prey per cell. Prey: Rhyssomatus
parvulus Casey, Tachiinae probably Sibinia sp.
Cerceris morata Cresson, 1872. Amer. Ent. Soc., Trans. 4: 230. 2.
Cerceris nasica Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 132. 9.

Biology: Alcock, 1974. Jour. Nat. Hist. 8: 651 (nest, prey).
neahminax Scullen. N. Mex., Ariz., Utah.
Cerceris neahminax Scullen, 1965. U.S. Natl. Mus., Proc. 116: 390, figs. 24, 128a, b, e. @.
nebrascensis Smith. S. Dak., Nebr., Wyo., Colo.
Cerceris nebrascensis Smith, 1908. Nebr. Univ. Studies 8: 368. 2.

1736 Hymenoptera in America North of Mexico

nigrescens Smith. Que. to B. C. south to N. C., Tex. and Calif. Ecology: Nests in small

aggregations in hard-packed, stony or sandy loam, stores up to 25 prey per cell.
Parasite: Senotainia trilineata (Wulp)? Prey: Calomycterus setarius Roelofs, Sitona
scissifrons (Say), S. hispidula (F.), Hyperodes delumbis Gyll., Gymnetron antirrhini
Payk., G. sp.

Cerceris nigrescens Smith, 1856. Cat. Hym. Brit. Mus. 4: 466. 2.

Cerceris arelate Banks, 1912. Ent. Soc. Amer., Ann. 5: 18. @.

Cerceris nigritulus Banks, 1915. Canad. Ent. 47: 402. 6.

Cerceris munda Mickel, 1918 (1917). Nebr. Univ. Studies 17: 337. 2, 6.

Cerceris abbreviata Banks, 1919. Canad. Ent. 51: 84. 6.

Cerceris crawfordi Brimley, 1928. Elisha Mitchell Sci. Soc., Jour. 43: 199. 6.

Biology: Peckham and Peckham, 1898. Wis. Geol. Nat. Hist. Surv., Bul. 2: 116-117 (nest,
prey). —Krombein, 1936. Ent. News 47: 94-95 (nest, prey, parasite ?). —Krombein, 1938.
Ent. News 49: 1-3 (nest, prey). —Evans, 1971. Kans. Ent. Soc., Jour. 44: 512 (nest, prey).

occipitomaculata Packard. Minn. and S. Dak. to Tex., Colo., Ariz.; Mexico (Chihuahua).
Cerceris occipitomaculata Packard, 1866. Ent. Soc. Phila., Proc. 6: 62. 3.
Cerceris fasciola Cresson, 1872. Amer. Ent. Soc., Trans. 4: 230. 6.
Cerceris novomexicana Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 137. °.
parkeri Scullen. South. Tex. and Ariz., south in Mexico to Chiapas.
Cerceris parkeri Seullen, 1972. Smithsn. Contrib. Zool. 110: 35, figs. 22, 112. 2, ¢.
poculum Scullen. South. Ariz., west. Tex.
Cerceris poculum Scullen, 1965. U. S. Natl. Mus., Proc. 116: 391, figs. 25, 124a, b, c. 2.
posticata Banks. N. Mex. (Jemez Mts.).
Cerceris posticata Banks, 1916. Ent. News 27: 64. 6.
rhois Rohwer. N. Mex. (White Mts.).
Cerceris rhois Rohwer, 1908. Canad. Ent. 40: 325. 6d.
robertsonii bifida Scullen. Va., N. C., Ga., Ind.
Cerceris robertsonti bifidus Scullen, 1965. U. S. Natl. Mus., Proce. 116: 428, figs. 45, 137a, b,
Cage
robertsonii emmiltosa Scullen. Fla., Ga. Ecology: Nests in slightly sloping, sparsely vegetated
sand, makes more than 6 cells per nest, stores 6-14 prey per cell. Prey: Colaspis favosa
Say, Paria sp.
Cerceris robertsonii emmiltosus Scullen, 1964. Ent. News 75: 144. 9.

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 277 (larva).

Biology: Krombein, 1964. Amer. Mus. Novitates 2201: 22-23 (nest, prey).
robertsonii miltosa Scullen. Fla.
Cerceris robertsonii miltosus Scullen, 1965. U. S. Natl. Mus., Proc. 116: 429, fig. 47. 2.
robertsonii robertsonii Fox. Que., Ont. to N. C. west to S. Dak. and Kans., Tex. Ecology:
Nests in small aggregations in fine, firm sand to coarse sandy gravel, stores 8-12 prey
per cell. Parasite: Metopia argyrocephala (Meig.), Miltogrammini sp. Prey:
Cryptocephalus notatus F., Pachybrachis dilatatus Suffr., Rhabdopterus picipes (Oliv.),
Tymnes tricolor F.
Cerceris robertsonii Fox, 1893. N. Y. Ent. Soe., Jour. 1: 55. 2, 6.
Cerceris austrina Fox, 1893. Psyche 6: 556. 2, 6.
Cerceris pleuralis Smith, 1908. Nebr. Univ. Studies 8: 366. 6, 9.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 86, figs. 27-30 (larva; described both as
nigrescens Sm. and robertsonii). —Evans, 1959. Amer. Ent. Soc., Trans. 85: 156-157
(larva).

Biology: Krombein, 1953 (1952). Wasmann Jour. Biol. 10: 288-294 (nest, prey, egg, life cycle,
parasite). —Krombein, 1953. Ent. Soe. Wash., Proc. 55: 119-121 (nest, prey). —Evans, 1959.
Amer. Ent. Soc., Trans. 85: 157 (prey). —Evans, 1971. Kans. Ent. Soc., Jour. 44: 511, fig. 8
(nest, prey).

rozeni Scullen. Fla. (Highlands Co.).
Cerceris rozeni Scullen, 1971. N. Y. Ent. Soc., Jour. 79: 130, 4 figs. 2, 6.

Superfamily SPHECOIDEA 1737

rubrata Bohart and Menke. Fla.
Cerceris rufa Seullen, 1965. U. S. Natl. Mus., Proc. 116: 449, fig. 61. 2, ¢. Preoce.

Cerceris rubrata Bohart and Menke, 1976. Sphecid wasps of world, p. 586. N. name.
rufinoda Cresson. Md., Va., N. C., Ohio, N. Dak. to Tex. west to B. C., Calif.; Mexico (Durango,
San Luis Potosi). Prey: Miccotrogus picirostris (F.), Smicronyx squalidus Casey.

Cerceris rufinoda Cresson, 1865. Ent. Soc. Phila., Proce. 5: 121. 6.
Cerceris rufinoda var. crucis Viereck and Cockerell, 1904. N. Y. Ent. Soce., Jour. 12: 139. 9.

Biology: Strandtmann, 1945. Ent. Soc. Amer., Ann. 38: 311, fig. 9 (nest, prey). —Scullen, 1951.

U.S. Dept. Agr., Monog. 2: 1010 (prey).

rufopicta Smith. Fla. to N. C.
Cerceris rufo-picta Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 467. 6.

sandiegensis Scullen. South. Calif. to southwest. N. Mex.
Cerceris sandiegensis Scullen, 1965. U.S. Natl. Mus., Proc. 116: 432, fig. 49. 2, d.

serripes (Fabricius). North America.
Vespa serripes Fabricius, 1781. Spec. Ins., v. 1, p. 46. 2.

Taxonomy: Scullen, 1965. U. S. Natl. Mus., Proc. 116: 503 (notes on type).

sexta Say. Nebr., Wyo. and Utah south to N. Mex. and Ariz.
Cerceris sexta Say, 1837. Boston Jour. Nat. Hist. 1: 382. 6.
Cerceris biungulata Cresson, 1865. Ent. Soe. Phila., Proc. 5: 118. 9.
Cerceris orphne Banks, 1947. Psyche 54: 15. ¢.

sextoides Banks. B. C. to south. Calif., Idaho, Utah. Prey: Sitona californica Fahr.,

Trigonoscuta pilosa Motsch.

Cerceris sextoides Banks, 1947. Psyche 54: 10. 2, 6.
Cerceris eurymele Banks, 1947. Psyche 54: 11. 2, 6.

Taxonomy: Scullen, 1969. Ent. News 80: 283-284 (separation of sextoides and tepaneca
Sauss.).

Biology: Scullen, 1965. U. S. Natl. Mus., Proc. 116: 510 (prey).

squamulifera Mickel. N. C., Ark., Nebr. to Tex.
Cerceris squamulifera Mickel, 1916. Amer. Ent. Soc., Trans. 42: 411. 2.

Taxonomy: Scullen, 1968. Ent. News 79: 154-156, 4 figs. 6.
stigmosalis Banks. Alta., B. C., Minn. to Mont. south to Tex.; Mexico (Sonora, Durango, San
Luis Potosi, Mexico).
Cerceris stigmosalis Banks, 1916. Ent. News 27: 64. 6.
Cerceris fugatrix Mickel, 1918 (1917). Nebr. Univ. Studies 17: 335. d.
Cerceris sayi Banks, 1923. Canad. Ent. 55: 21. @, ¢.
Cerceris stevensi Banks, 1923. Canad. Ent. 55: 22. 2.
texana Scullen. Tex.
Cerceris texana Scullen, 1965. U.S. Natl. Mus., Proc. 116: 511, figs. 97, 177a, b, e. 2.
tolteca Saussure. Ga., Fla., La., Okla., Ark., Tex., Ariz.; Mexico, Honduras.
Cerceris Tolteca Saussure, 1867. Reise d. Novara, Zool., v. 2, Hym., p. 94. 2, 6.
Cerceris cosmiocephala Cameron, 1904. Invertebrata Pacifica 1: 67. 3.
townsendi Viereck and Cockerell. N. Mex. (Las Cruces).
Cerceris townsendi Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 140. 2.
truncata Cameron. South. Tex, N. Mex., Ariz., south to Costa Rica. Ecology: Nests in large
aggregation in hard-packed soil, stores up to 10 prey per cell. Prey: Algarobius prosopis
(LeC.), Mimosestes protractus (Horn), M. amicus (Horn), Neltumius arizonensis
(Sehffr.).
Cerceris truncata Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 121, pl. 7, figs. 19, 20. 2,
é.

Biology: Werner, 1960. Psyche 67: 43-44 (nest, male activity, prey).
vanduzeei Banks. B. C. to Calif., Idaho, Utah.
Cerceris vanduzeei Banks, 1917. Mus. Compar. Zool., Bul. 61: 114. @.
Cerceris complanata Mickel, 1918 (1917). Nebr. Univ. Studies 17: 340. ¢, 2.
Cerceris vanduzeei eburnea Scullen, 1965. U. S. Natl. Mus., Proc. 116: 399, fig. 30. 2.

1738 Hymenoptera in America North of Mexico

varians Mickel. Calif., Nev.; Mexico (Michoacan).
Cerceris varians Mickel, 1918 (1917). Nebr. Univ. Studies 17: 336. ?, d.

veracruz veracruz Scullen. Tex. south in Mexico to Campeche. Another subsp. occurs in El

Salvador.
Cerceris veracruz veracruz Seullen, 1972. Smithsn. Contrib. Zool. 110: 53, figs. 45, 132a-d.
OPaor
verticalis Smith. N. C. to Fla. west to Kans. and Tex.; Mexico (Coahuila, Nuevo Leon,
Tamaulipas).

Cerceris verticalis Smith, 1856. Cat. Hym. Brit. Mus., v. 4, p. 466. 9.

Cerceris gnara Cresson, 1872. Amer. Ent. Soc., Trans. 4: 229. 6.

Cerceris firma Cresson, 1872. Amer. Ent. Soc., Trans. 4: 229. 2.
vicina Cresson. Nebr., Wyo., Colo., Kans., Tex.

Cerceris vicina Cresson, 1865. Ent. Soe. Phila., Proe. 5: 120. °.

Cerceris platyrhina Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 133. 2.
wyomingensis Scullen. N. Dak., Wyo.

Cerceris wyomingensis Scullen, 1965. U. S. Natl. Mus., Proc. 116: 519, figs. 102, 182a, b, e.

OF

zelica Banks. N. Y., Minn. and S. Dak. to N. C. and Tex.

Cerceris zelica Banks, 1912. Ent. Soe. Amer., Ann. 5: 23. 6, °.

Genus EUCERCERIS Cresson

Eucerceris Cresson, 1865. Ent. Soc. Phila., Proc. 5: 104.
Type-species: Eucerceris fulvipes Cresson. Desig. by Pate, 1937.

Revision: Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 1-80, figs. 1-158e. —Scullen,
1968. U. S. Natl. Mus., Bul. 268: 1-97, figs. 1-98h.

Taxonomy: Scullen, 1948. Pan-Pacifie Ent. 24: 155-180 (revised key).

angulata Rohwer. N. Mex., Ariz.; Mexico (Sonora, Baja California).
Eucerceris angulata Rohwer, 1912. Amer. Mus. Nat. Hist., Bul. 31: 326. 2.

Taxonomy: Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 57. 6.

arenaria Scullen. Southwest. Tex. to Calif., Nev., Colo.
Eucerceris arenaria Seullen, 1948 Pan-Pacific Ent. 24: 168. 2, do.

Biology: Alcock, 1975. Kans. Ent. Soc., Jour. 48: 533-536, figs. 1, 3 (male mating strategy).
bitruncata Scullen. Tex., N. Mex., Ariz. Ecology: Nests in flat, stony soil, makes a multicellular
nest, stores 39-41 small prey per cell; first progeny may transform to adults before

mother completes nest. Prey: Minyomerus languidus Horn.
Eucerceris bitruncata Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 35. 9.
Eucerceris triciliata Seullen, 1948. Pan-Pacific Ent. 24: 172. 3d. N. syn. (R. M. Bohart).

Taxonomy: Evans, 1964. Amer. Ent. Soc., Trans. 90: 278-279, figs. 71-73 (larva).

Biology: Krombein, 1960. Brooklyn Ent. Soc., Bul. 55: 77-79 (nest, prey, cocoon).
canaliculata (Say). Ark., S. Dak. and Mont. south to Tex. and Ariz., Calif.; Mexico (Baja

California and Durango south to Hidalgo).

Philanthus canaliculatus Say, 1823. West Quart. Rptr. 2: 79. d.

Cerceris bidentata Say, 1823. West. Quart. Rptr. 2: 80. ?. Preoce.

Cerceris cameroni Schulz, 1906. Spolia Hym., p. 194. N. name.

Eucerceris canaliculata var. atronitida Scullen, 1939. Oreg. State Monog., Studies in Ent.
IE HOS Cpe

Eucerceris biconica Seullen, 1948. Pan-Pacific Ent. 24: 178. 2.

Biology: Alcock, 1975. Kans. Ent. Soc., Jour. 48: 536-538, figs. 2, 4 (male mating strategy).
conata Scullen. Nebr., Tex.
Eucerceris conata Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 34. @.
Eucerceris hespera Scullen, 1948. Pan-Pacifie Ent. 24: 171. 6.

Superfamily SPHECOIDEA 1739

elegans elegans Cresson. Nev., Calif.; Mexico (Baja California).
Eucerceris elegans Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proe., p. xxiii. d. Preoce. in
Cerceris.
Cerceris nevadensis Dalla Torre, 1890. Wien. Ent. Ztg. 9: 200. N. name.
elegans ferruginosa Scullen. South. Calif.
Eucerceris ferruginosa Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 45. 9.
Eucerceris mojavensis Scullen, 1968. U. S. Natl. Mus., Bul. 268: 44, figs. 26, 80a-c. ¢.
elegans monoensis Scullen. East. centr. Calif.
Eucerceris elegans monoensis Scullen, 1968. U. S. Natl. Mus., Bul. 268: 28, figs. 14, 69a-f.
eG
flavocincta Cresson. Man. to B. C. south to N. Mex. and Calif. Ecology: Nests in hard stony
soil, provides up to 7 cells per nest, stores 6-7 prey per cell. Parasite: Hedychrum
nigropilosum Mocs.? Prey: Dyslobus lecontei Casey, D. segnis (LeC.), D. sp., Panscopus
aequalis (Horn), Peritaxia nigricollis Horn.
Eucerceris flavocinctus Cresson, 1865. Ent. Soc. Phila., Proce. 5: 109. 9.
Eucerceris cingulatus Cresson, 1865. Ent. Soc. Phila., Proc. 5: 110. 6.
Eucerceris striareata Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 85. 2.
Eucerceris chapmanae Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 86. 3.

Taxonomy: Evans, 1957. Amer. Ent. Soc., Trans. 83: 88, figs. 32-35 (larva).

Biology: Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 12-14 (nest, prey, cocoon, male
behavior). —Bohart and Powell, 1956. Pan-Pacific Ent. 32 143-144 (nest, prey, parasite 7).
—Evans, 1970. Mus. Compar. Zool., Bul. 140: 502 (nest, prey).

fulvipes Cresson. Alta., Mont., S. Dak. to Wyo., south to Tex., Ariz. and south. Calif. Ecology:

Nests in hard stony soil, makes more than 5 cells per nest, stores 12-18 prey per cell.
Parasite: Taxigramma heteroneura (Meig.)?; Hedychrum parvum Aar.? Prey:
Brachyrhinus ovatus L., Ceutorhynchus punctiger Gyll., Hyperodes sp. Predator:
Philanthus zebratus Cr.

Eucerceris fulvipes Cresson, 1865. Ent. Soc. Phila., Proc. 5: 111. 2, 6. Preoce. in Cerceris.

Cerceris cressoni Schletterer, 1887. Zool. Jahrb., Ztschr. f. System. 2: 489. N. name.

Eucerceris flavipes(!) Ashmead, 1899. Canad. Ent. 31: 295. °.

Eucerceris simulatrix Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12: 87. d.

Biology: Evans, 1970. Mus. Compar. Zool., Bul. 140: 502 (nest, prey transport, parasites ?).
insignis Provancher. Nev., Calif.; Mexico (Baja California).
Eucerceris insignis Provancher, 1889. Addit. Corr. Faune Ent. Canada Hym., p. 418. 6.
Preoce. in Cerceris.
Cerceris provancheri Dalla Torre, 1890. Wien. Ent. Ztg. 9: 204. N. name.
lacunosa lacunosa Scullen. South. Ariz. to southwest. Tex.; Mexico (Chihuahua, Durango).
Eucerceris lacunosa Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 19. d.
Eucerceris arizonensis Seullen, 1939. Oreg. State Monog., Studies in Ent. 1: 20. 9.
lacunosa sabinasae Scullen. South. Ariz.; Mexico (Coahuila).
Eucerceris lacunosa sabinasae Scullen, 1968. U.S. Natl. Mus., Bul. 268: 36, fig. 20. ¢, 9.
melanovittata Scullen. South. Ariz., N. Mex., west. Tex.; Mexico (Nuevo Leon).
Eucerceris melanovittata Seullen, 1948. Pan-Pacific Ent. 24: 164. ¢.
mellea Scullen. Southwest. Tex., N. Mex.; Mexico (Chihuahua).
Eucerceris mellea Seullen, 1948. Pan-Pacifie Ent. 24: 165. 2, ¢.
montana Cresson. Mont., Utah to Kans., south to Ariz. and west. Tex., south in Mexico to
Jalisco and San Luis Potosi.
Eucerceris montanus Cresson, 1882. Amer. Ent. Soc., Trans. 10: Proc., p. viii. 2, d.
Cerceris sonorensis Cameron, 1891. Biol. Cent.-Amer., Hym., v. 2, p. 129, pl. 8, fig. 10. ¢.
morula albarenae Scullen. South. N. Mex., southwest. Tex.
Eucerceris morula albarenae Scullen, 1968. U. S. Natl. Mus., Bul. 268: 46, fig. 29. 2, ¢.
morula morula Scullen. South. N. Mex., southwest. Tex.; Mexico (San Luis Potosi,
Aguascalientes, Zacatecas, Queretaro, Hidalgo).
Eucerceris morula morula Seullen, 1968. U. S. Natl. Mus., Bul. 268: 49, figs. 30, 82a-f. 2,
3.

1740 Hymenoptera in America North of Mexico

pimarum Cockerell and Rohwer. S. Dak. through Colo., Utah, Nev. to Calif. south to west.
Tex. and Ariz.; Mexico (Chihuahua, Coahuila, Durango).
Eucerceris pimarum Cockerell and Rohwer, 1908. In Rohwer, Canad. Ent. 40: 326. °.
Eucerceris apicata Banks, 1915. Canad. Ent. 47: 404. 6.
rubripes Cresson. Rocky Mts. to east. Nebr. and Kans., north to Canada and south to Tex.
Prey: Peritaxia sp.
Eucerceris rubripes Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxiii. d.
Eucerceris unicornis Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 359. °.
Aphilanthops marginipennis Cameron, 1890. Biol. Cent.-Amer., Hym., v. 2, p. 105. d.

Biology: Scullen, 1968. U. S. Natl. Mus., Bul. 268: 56 (prey). —Aleock, 1975. Kans. Ent. Soc.,
Jour. 48: 538 (male mating strategy).
ruficeps Scullen. Cent. Calif., Nev. Ecology: Nests in hard-packed sand in abandoned burrows
of the bee Sphecodogastra aberrans (Cwfd.), makes up to 5 or more cells per nest,
stores 6-22 prey per cell. Parasite: Miltogrammini spp. Prey: Dysticheus rotwndicollis
Van Dyke, Sitona californicus Fahr.
Eucerceris ruficeps Scullen, 1948. Pan-Pacifie Ent. 24: 175. 2.

Biology: Linsley and MacSwain, 1954. Pan-Pacifie Ent. 30: 11-14 (nest, prey, parasites).
similis Cresson. Oreg., Calif., Nev., south. Idaho, west. Wyo., Colo.
Eucerceris similis Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxiv. 2, 6.
Eucerceris barri Seullen, 1968. U.S. Natl. Mus., Bul. 268: 19, figs. 7, 65a-g. 2, ¢.
sinuata Scullen. South centr. Tex.; Mexico (Nuevo Leon, Coahuila).
Eucerceris sinuata Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 47. °.
superba bicolor Cresson. Alta., Mont., N. Dak., S. Dak., Wyo. Prey: Ophryastes sulcirostris
(Say), O. sp. in sulcirostris-porosus complex.
Eucerceris bicolor Cresson, 1881. Amer. Ent. Soc., Trans. 9: Proe., p. xxxviii. 2. Preoce. in

Cerceris.
Cerceris dichroa Dalla Torre, 1890. Wien. Ent. Ztg. 9: 199. 2. N. name.

Biology: Scullen, 1939. Oreg. State Monog., Studies in Ent. 1: 40 (prey). —Scullen, 1968. U.S.
Natl. Mus., Bul. 268: 66 (prey).
superba superba Cresson. Rocky Mts. east to N. Dak., S. Dak., Nebr., Kans.
Eucerceris superbus Cresson, 1865. Ent. Soc. Phila., Proc. 5: 108. d.
Eucerceris fulviceps Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxiii. °.
Eucerceris fulviceps var. rhodops Viereck and Cockerell, 1904. N. Y. Ent. Soc., Jour. 12:
Sieh, Be ”
tricolor Cockerell. Ariz., N. Mex., west Tex.; Mexico (Chihuahua, Coahuila, Hidalgo, Zacatecas).
Eucerceris vittatifrons var. tricolor Cockerell, 1897. Entomologist 30: 136. ¢.

Biology: Alcock, 1975. Kans. Ent. Soc., Jour. 48: 538 (male mating strategy).
vittatifrons Cresson. Wash., Calif., Nev., Utah, Colo., Ariz., N. Mex., Oreg., Idaho.
Eucerceris vittatifrons Cresson, 1879. Amer. Ent. Soc., Trans. 7: Proc., p. xxiv. d.
zonata (Say). New England States, N. Y., Ont. west to N. Dak., north. centr. States west to
Wyo. and Colo., Ark., Tex. Ecology: Makes multicellular nest of as many as 9 cells in
fine-grained friable sand, stores 3-7 prey per cell. Prey: Cleonus plumbeus LeC.
Philanthus zonata Say, 1823. West. Quart. Rptr. 2: 80. d.
Eucerceris laticeps Cresson, 1865. Ent. Soc. Phila., Proc. 5: 107. 6, @.

Taxonomy: Evans, 1971. Kans. Ent. Soc., Jour. 44: 517-518, figs. 15-21 (larva).

Biology: Evans, 1971. Kans. Ent. Soce., Jour. 44: 514-517, figs. 11, 18, 14 (nest, prey, egg,
cocoon).

1741

Superfamily APOIDEA

By PAUL D. HuRD, JR.

This superfamily contains the bees which, like many other aculeates, visit flowers for nectar.
However, unlike nearly all other aculeates, the bees and most wasps of the Masaridae provide or
feed their larvae with a mixture of pollen and nectar or, in some bees, the mixture is converted
into glandular substances which are then fed to their larvae as well as to certain adults. It is
presumed by many specialists that bees as a group evolved from flower-visiting wasps, most
likely the sphecoid wasps (but possibly also other groups of aculeate wasps), by developing a de-
pendence for food upon pollen and nectar (and sometimes other substances, such as plant oils)
and thereby have abandoned the habit of provisioning their nests with insect or spider prey. It
is not known when this dependency arose, but it could not have occurred before the advent of
Angiosperms which did not begin to flourish until the latter half of the Cretaceous period and
which then became the dominant flora of the earth by the Tertiary. In general, the evolution of
the entomophilous flower has resulted in the replacement of a shallow, flat or bowl-shaped
flower by a corolla-tube. The progressive increase in the depth of the corolla-tube conceivably
has resulted from coevolutionary interactions between the flowers and entomophilous insects,
especially bees, with progressively more elongated mouthparts. Although the earliest known
fossils of bees (Tertiary) are insufficient to establish ancestral relationships with other aculeates
or to demonstrate that the evolution of mouthparts proceeded from the short to the
long-tongued condition, it is probably significant that this sequence is being corroborated by stu-
dies on the systematics, morphology, biology, behavior and biogeography of the contemporary
bee fauna of the world. This fauna is at present considered to consist of eight families which in
most current classifications are usually arranged phylogenetically as follows: Colletidae, Ox-
aeidae, Andrenidae, Halictidae, Melittidae, Megachilidae (including the Fideliinae),
Anthophoridae and Apidae. Even though there is some and sometimes much variation in the
length of the glossa within each of these families, the first five families listed above contain the
so-called short-tongued bees while the remaining families consist of the so-called long-tongued
bees. Although the short-tongued bees of only three families (Colletidae, Melittidae and Halic-
tidae) are present on all continents, only the Colletidae are exceptionally diverse and
well-represented on the southern continents, especially in Australia where other presumably an-
cient groups of plants and animals survive today. The glossa of the Colletidae, in addition to
being normally short and bilobed, is also usually truncate and bifid and therefore is more
wasp-like in structure than is the glossa of any other family of bees. Thus it is believed that the
initial evolution of bees resulted in the development of short-tongued bees which radiated
throughout the earth and this event was subsequently followed by the coevolution of the corol-
la-tube and the long-tongued bees which also have spread throughout the world. Consequently,
the Colletidae are regarded by most specialists as representing the most primitive group of liv-
ing bees. Most, but not all, authors believe that the Apoidea represent a monophyletic assem-

1742 Hymenoptera in America North of Mexico

blage which evolved from a sphecoid ancestor. Brothers (1975) has concluded that the sphecoid
wasps and bees belong to a single superfamily, the Sphecoidea, which according to him consists
of two informal groups, the Spheciformes (8 families) and the Apiformes (9 families). However,
Lanham (1960) has commented upon the similarities and differences of the Sphecoidea and
Apoidea and points out that the presence or absence of the strigil on the hind leg long known to
European hymenopterists may be of as much phylogenetic significance as other characters of
presumably phylogenetic importance. The value of the strigil in these matters was employed by
Boerner (1919) who divides the aculeate Hymenoptera into the Haplocnemata (ants, scolioid
wasps, and bees) and the Diplocnemata (sphecoids, pompilids, and vespids) thereby indicating
that bees are more closely related to the scolioids than to the sphecoid wasps. The phylogeny of
the Aculeata deserves a more thorough study and reevaluation before we can dismiss the possi-
bility that the Apoidea is a polyphyletic assemblage having been derived from both scolioid and
sphecoid ancestors.

Most species of bees construct their nests in the ground usually excavating their own tunnels
and cells, although many others appropriate preexisting burrows or other cavities in the ground
and sometimes modify these to accomplish their needs. Still many others make their nests above
ground. Among these are species that gnaw out their nesting tunnels in wood substrates of vari-
ous kinds including hollow stemmed plants, while others appropriate a wide variety of preexist-
ing cavities, such as abandoned beetle burrows, hollow trees, old mason wasp nests, old bird
nests, empty snail shells and old insect galls, while still others make their nests of wax and other
materials such as mud, resin, pebbles, pieces of leaves or petals, plant down, etc., and place their
nests either in the open attaching them to branches and so forth or place their nests under
eaves, bridges, rocks, cow chips and so on. As a consequence of these habits, many species of the
families Megachilidae and Apidae readily accept artificial nesting devices (hives, trap-nests, etc.)
which not only has made possible detailed studies of their biologies, but also has made possible
the manipulation and management of several species, including the common honeybee, for use in
the pollination of agricultural crops or for the production of honey and other useful products of
value and benefit to mankind.

In nesting behavior the vast majority of bees are solitary including all members of the fami-
lies Colletidae, Oxaeidae and Melittidae. Except for a few communal species, all members of the
family Andrenidae also exhibit solitary nesting behavior. Similarly most species of the family
Megachilidae are solitary in habit, although some nest communally (parasocial behavior) while
some are quasisocial and possibly a few are even semisocial in their nesting behavior. Among the
remaining three families (Halictidae, Anthophoridae and Apidae), the Halictidae and
Anthophoridae, each represented by many solitary species, contain some parasocial and eusocial
(primitively social) species as well as some subsocial species in the anthophorid tribe Ceratinini.
While most species of the family Apidae live in perennial, highly eusocial colonies, others exhibit
solitary, parasocial or primitively social nesting behavior.

Bees are exceptionally important pollinators of both native vegetation and many agricultural
crops. Unquestionably bee-plant (bee-flower) relationships reflect various strategies on the part
of both sets of participants and two of the most commonly recognized behavioral modes of pollen
collection by bees are oligolecty and polylecty (see reviews by Grant, 1950; Linsley, 1958; and
Baker and Hurd, 1968). The intrafloral relationships of bees not only involves their own survival
and evolution, but also through coevolutionary interactions with flowers insures the main-
tenance and evolution of much of the earth’s flora that depends upon entomophilous pollination
for reproduction. These interrelationships are of unus»al significance to us because we in turn
depend upon the earth’s flora for our own livelihood and welfare.

In America north of Mexico there are about 3,500 species of bees. This area represents nearly
one-sixth of the earth’s land surface and, disregarding ecological diversity, this means that the
world fauna of bees approximates 21,000 species. Somewhat more than 2,700 of our species are
pollen-collecting bees while slightly more than 700 species (or about 21%) are cleptoparasitic spe-
cies. Only about 800 species of bees occur east of the Mississippi River and thus the apifauna of
the larger and more ecologically diverse western portion of America north of Mexico is more
than three times richer in species. Since it is well established that the apifaunas of arid regions
are consistently much richer in species than any other climatic regions, it is not surprising that
most of the North American species of bees are to be found in the southwestern United States
and adjacent northern Mexico. All eight recognized families of bees are present in America
north of Mexico and are represented by the following numbers of species: Colletidae (153), Ox-

Superfamily APOIDEA 1743

aeidae (4), Andrenidae (1,199), Halictidae (506), Melittidae (30), Megachilidae (607),
Anthophoridae (919), and Apidae (47). Among the largest genera in our fauna are: Andrena
(511), Perdita (502), Nomada (286), Dialictus (189), Osmia (133), Megachile (114), Triepeolus
(102), Melissodes (99), and Colletes (96). In all, our fauna consists of 121 genera of which 92 con-
tain the pollen-collecting species. Several of our species are Holarctic in distribution and eight
species, including the European honeybee, are of introduced origin.

The biology of the Apoidea including their behavior, intrafloral ecology, nesting habits, life
histories, communication and the like has always fascinated and attracted much interest and
study by the laity, the beekeeper and the specialist. An immense literature on these and related
subjects has already accumulated and continues to develop so that it is becoming almost an im-
possibility to accomplish an in-depth bibliography of the Apoidea. For example, there are now
more than 100,000 references to the European honeybee alone. Fortunately, the task is made
easier by several literature information retrieval systems and especially by the International
Bee Research Association, headquartered in England, which publishes the key abstracting jour-
nal on bees, Apicultural Abstracts, two key research journals in English, Bee World and the
Journal of Apicultural Research, as well as comprehensive bibliographies (J. B. R. A. Bibliogra-
phies) on selected subjects pertaining to bees.

Revision: Dalla Torre, 1896. Cat. Hym., v. 10: viii and 643 pp. (classification, catalog of world
spp.; lists 6,165 spp. in 186 genera distributed among 14 subfamilies and all placed in one
family, the Apidae). —Boerner, 1919. Biol. Zentrbl. 39: 145-186, 6 figs. (classification).
—Friese, 1923. Die Europaischen Bienen (Apidae), vii and 456 pp., W. de Gruyter and Co.,
Berlin and Leipzig (classification, life histories). —Grutte, 1935. Arch. Naturgesch. (n, f.) 4:
449-534 (classification of parasitic spp.). —Sandhouse, 1943. U.S. Natl. Mus., Proc. 92:
519-619 (type-species of genera and subgenera). —Michener, 1944. Amer. Mus. Nat. Hist.,
Bul. 82: 151-326, text figs. 1-246, diagrams 1-13 (morphology, phylogeny and classification).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 1-538, 134 figs., 16 tables (east. U. S.
spp. of Colletidae, Andrenidae, Halictidae and Melittidae). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 1-557, 134 figs., 18 tables (east. U. S. spp. of Megachilidae,
Anthophoridae, Xylocopidae, Apidae). —Michener, 1965. Amer. Mus. Nat. Hist., Bul. 130:
1-362, 789 text figs., 15 pls., 4 tables (classification). —Weber, 1965. Colo. Univ. Studies,
Series in Bibliography 1: 1-124, 1 frontis. (bibliography of T. D. A. Cockerell). —Stephen,
Bohart and Torchio, 1969. Oreg. State Univ. Agr. Expt. Sta., pp. 1-140, 320 figs.
(classification, morphology, phylogeny and biology of northwest. U. S. spp.). —Michener,
1974. The social behavior of the bees, chapter 3: 25-29. Cambridge, Mass. The Belknap
Press of Harvard Univ. Press (classification). |

Taxonomy: Cockerell, 1898. Sci. Lab. Denison Univ., Bul. 11: 41-73 (N. Mex. spp.).
—Cockerell, 1898. N. Mex. Univ., Bul. 1: 43-73 (N. Mex. spp.). —Ashmead, 1899. Amer.
Ent. Soc., Trans. 26: 49-100 (classification). —Fowler, 1902. Calif. Agr. Expt. Sta., Rpts.
1899-1901, pt. 2, pp. 316-330 (long-tongued Calif. spp.). —Cockerell, 1903. Psyche 10: 74-78
(Calif. spp.). —Robertson, 1903. Amer. Ent. Soc., Trans. 29: 163-189 (synopsis of
Megachilidae and Bombinae). —Cockerell and Robbins, 1910. Colo. Univ. Studies 7:
179-195, 8 pls. (Rocky Mts. spp.). —Graenicher, 1911. Wis. Nat. Hist. Soc., Bul. 1: 221-249
(north. Wis. spp.). —Lutz and Cockerell, 1920. Amer. Mus. Nat. Hist., Bul. 42: 491-641
(notes on distribution, bibliography and floral records of N. Amer. spp. of Anthophoridae
and Apidae). —Cockerell, 1924. Ent. Soc. Wash., Proc. 26: 77-85 (tax. characters).
—Cockerell, 1928. Colo. Univ. Studies 16: 99-126 (Colo. spp.). —Michener, 1941. Sixth
Pacific Sci. Congr., Proc. 4: 297-303 (distributional history of N. Amer. fauna). —Michener,
1947. Amer. Midland Nat. 38: 443-455 (south. Miss. spp.). —Stevens, 1948. N. Dak. Agr.
Expt. Sta. Bimonthly Bul. 10: 187-194 (N. Dak. spp.). —Stevens, 1948. N. Dak. Agr. Expt.
Sta., Bimonthly Bul. 11: 49-54, 2 figs. (N. Dak. spp.). —Stevens, 1949. N. Dak. Agr. Expt.
Sta., Bimonthly Bul. 11: 130-135, 210-225, 4 figs. (N. Dak. spp.). —Richards, 1949. Linn. Soe.
London, Proe. 161: 40-41 (evolution of cuckoo spp.). —Stevens, 1949. N. Dak. Agr. Expt.
Sta. Bimonthly Bul. 12: 14-22 (N. Dak. spp.). —Stevens, 1950. N. Dak. Agr. Expt. Sta.,
Bimonthly Bul. 12: 90-98, 3 figs. (N. Dak. spp.). —Stevens, 1950. N. Dak. Agr. Expt. Sta.,
Bimonthly Bul. 13: 72-80, 4 figs. (N. Dak. spp.). —Bohart and Knowlton, 1950. Utah State
Agr. Expt. Sta. mimeo series, 371: 1-5 (Utah spp.). —Buckell, 1951. Ent. Soe. Brit.
Columbia, Proc. 47: 7-24 (B. C. spp.). —Stevens, 1951. N. Dak. Agr. Expt. Sta., Bimonthly

1744 Hymenoptera in America North of Mexico

Bul. 13: 199-205, 2 figs. (N. Dak. spp.). —Stevens, 1951. N. Dak. Agr. Expt. Sta., Bimonthly
Bul. 14: 27-31, 59-64, 2 figs. (N. Dak. spp.). —Stevens, 1952. N. Dak. Agr. Expt. Sta.,
Bimonthly Bul. 14: 105-112, 2 figs. (collecting, mounting, labeling, identifying, study, life
histories). —Michener, 1953. Kans. Univ. Sci. Bul. 35: 987-1102, figs. 1-287 (larvae).
—Michener, 1954. Pan-Pacific Ent. 30: 63-70, fig. 1, table 1 (pupae). —Michener, 1954.
Amer. Mus. Nat. Hist., Bul. 104: 1-176, figs. 1-55, tables 1-3 (classification). —Michener,
1955. A century of progress in the natural sciences, pp. 575-579, Calif. Acad. Sci., San
Francisco (Apoidea). —LaBerge, 1956. Kans. Univ. Sci. Bul. 38: 501-531 (types in Snow
Entomological Museum). —Krunic, 1959. Zbornik Matice Srpske (Novi Sad) 17: 102-111
(transitional forms between solitary and social spp.). —Lanham, 1960. Ent. News 71: 85-86
(diagnostic characters). —Moure, 1960. Studia Ent. 3: 97-160 (Fabricius types of
Neotropical spp.). —Hurd, 1966. Ent. Soc. Amer., Bul. 12: 110-111 (distributional patterns
in west. N. Amer.). —Nielsen and Bohart, 1967. Ent. Soc. Amer., Ann. 60: 414-419, 18 figs.
(larval sex characters). —Covell, 1972. Ent. Soc. Wash., Proc. 74: 10-18 (Lovell types).
—Kerr and da Silveira, 1972. Evolution 26: 197-202 (karyotypic evolution and tax.
implications). —Bohart and Knowlton, 1973. Utah Acad. Sci. Arts and Letters, Proc. 50:
1-9 (spp. of Curlew Valley of Utah and Idaho). —Brothers, 1975. Kans. Univ. Sci. Bul. 50:
586-587, 640-641 (phylogeny). —Moldenke, 1977 (1976). Wasmann Jour. Biol. 34: 147-178, 1
fig., 6 tables (evolutionary history and diversity of faunas of Chile and Pacific North
America).

Biology: Loew, 1884. Jahrb. Bot. Garten 3: 69-118 (floral relationships, oligotrophy and
polytrophy). —Robertson, 1888. Bot. Gazette 13: 33-34 (effect of wind on behavior).
—Verhoeff, 1892. Zool. Anz. 15: 41-43 (relationships between host and parasitic bee
larvae). —Bulman, 1892. Sci. Gossip 329: 98-99 (floral constancy). —Robertson, 1899. Bot.
Gazette 28: 215 (oligotrophy). —Friese, 1899. Zool. Jahrb., Abt. Syst. 3: 847-870 (parasitic
bees and their hosts). —Bulman, 1902. Zoologist 6: 220-222 (floral constancy).
—Graenicher, 1905. Wis. Nat. Hist. Soe., Bul. 3: 153-167 (life history and habits of parasitic
bees). —Latter, 1906. Nature 74: 200 (how inquiline bees find their hosts). —Lovell, 1913.
Ent. News 24: 104-112 (origin of oligotrophy). —Robertson, 1914. Ent. News 25: 67-73
(origin of oligotrophy). —Lovell, 1914. Ent. News 25: 314-321 (origin of oligotrophism).
—Gutbier, 1915. Soc. Ent. Ross., Horae 41: 1-57, 2 pls. (classification and evolution of
nests). —Robertson, 1918. Ent. News 29: 340-342 (proterandry and flight behavior).
—Betts, 1920. Bee World 2: 10-11 (floral constancy). —Clements and Long, 1923. Carnegie
Inst. Wash., Pub. 336: 1-274 (experimental pollination). —Lutz, 1924. N. Y. Acad. Sci., Ann.
29: 181-283 (u. v. floral patterns and flower visiting habits). —Robertson, 1924. Ecology 5:
393-407 (phenology of entomophilous flowers). —Robertson, 1925. Ecology 6: 412-436
(heterotrophy). —Robertson, 1926. Psyche 33: 116-120 (phenology of inquiline and
nest-making bees). —Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 157-277, 8 pls. (life
histories). —Hicks, 1926. Colo. Univ. Studies 15: 217-310 (nesting habits and parasites of
certain Boulder County, Colo. spp.). —Robertson, 1926. Ecology 7: 378-380 (list of
oligolectie spp.). —Robertson, 1928. List of visitors of 453 flowers, 221 pp., Carlinville,
Illinois. —Robertson, 1929. Psyche 36: 112-118 (phenology of oligolectic spp.). —Robertson,
1929. Flowers and insects, 221 pp., Lancaster, Pa., Science Press. —Bromley, 1930. N. Y.
Ent. Soce., Jour. 38: 159-175 (bee-killing robber flies). —Graenicher, 1930. Ent. Soc. Amer.,
Ann. 23: 285-310 (bee-fauna and vegetation of Miami, Fla.). —Robertson, 1930. Ent. News
41: 154-157, 331-336 (proterandry and flight behavior). —Atwood, 1933. Canad. Jour.
Research 9: 443-457 (apple blossom visiting spp. in N. S.). —Cockerell, 1933. Amer. Nat.
67: 1-3 (excessive abundance). —Pearson, 1933. Ecol. Monog. 3: 374-441 (ecological
relationships of spp. in Chicago region). —Rau, 1933. Jungle bees and wasps of Barro
Colorado Island, 324 pp., 112 figs., Kirkwood, Mo. (life histories). —Hicks, 1934. Colo. Univ.
Studies 21: 265-271 (parasites). —Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 219-224
(behavior of certain solitary and social spp.). —Betts, 1935. Bee World 16: 111-113 (floral
constancy). —Cockerell, 1935. Science 81: 458-459 (origin of higher flowering plants and
their insect visitors). —Graenicher, 1935. Ent. Soc. Amer., Ann. 38: 285-310 (bee-fauna and
vegetation of Wis., visitors). —Malyshev, 1935. Eos 11: 201-309, pls. III-XV (nesting
habits of solitary spp.). —Linsley and MacSwain, 1942. South. Calif. Acad. Sci., Bul. 40:
126-137 (nest drepredation by Ptinws californicus Pic). —Linsley, 1942. Calif. Univ. Pubs.

Superfamily APOIDEA 1745

Ent. 7: 189-206, pls. 6-7, 1 text fig. (bionomics of Hornia, a nest parasite). —Linsley and
MacSwain, 1944 (1943) Ent. Soe. Amer., Ann. 36: 589-601 (predation by Trichodes ornatus
Say). —Linsley, 1944. Pan-Pacific Ent. 20: 67-68 (bee prey records of Callinicus calcaneus
Loew). —Linsley, 1944. Brooklyn Ent. Soe., Bul. 39: 54-55 (sapygid parasites). —Popov,
1945. Zhur. Obsch. Biol. 6: 183-203 (parasitism in bees). —Linsley, 1946. Econ. Ent., Jour.
39: 18-29 (alfalfa pollinating spp. in Calif.). — Mitchell, 1946. Research and Farming,
Raleigh, N. C. 4: 1-2, 11 (DDT as threat to bees). —Peck and Bolton, 1946. Sci,Agr. 26:
338-418 (alfalfa pollinating spp. in Sask.). —Bohart, 1947. Farm and Home Sci. Utah Agr.
Expt. Sta. 8: 13-14 (alfalfa pollinating spp. in Utah). —Linsley and MacSwain, 1947. Econ.
Ent., Jour. 40: 349-357 (factors influencing effectiveness of alfalfa pollinating spp. in
Calif.). —Richards, 1949. Linn. Soe. London, Proc. 161: 40-41 (evolution of cuckoo bees and
wasps). —Linsley, MacSwain and Smith, 1950. Econ. Ent., Jour. 43: 59-62 (DDT
susceptibility). —Grant, 1950. Bot. Rev. 16: 379-398 (flower constancy). —Linsley and
MacSwain, 1951. South. Calif. Acad. Sci. Bul. 50: 92-95 (parasitism by Tricrania stansburyi
Hald.). —Larking, 1952. Agron. Jour. 44: 216-218 (alfalfa pollinating spp.). —Linsley and
MacSwain, 1952. Wasmann Jour. Biol. 10: 91-102 (parasitism by Nemognatha spp.).
—Linsley, MacSwain and Smith, 1952. Ecology 33: 558-567 (outline for study of life
histories of solitary and semisocial spp.). —Pengelly, 1953. 84th Ann. Rpt. Ent. Soc. Ont.,
pp. 101-118 (alfalfa pollinating spp. in Ont.). —Michener, 1953. Century of Progress in the
natural sciences, pp. 575-578, Calif. Acad. Sci., San Francisco (Apoidea). —Michener, Cross,
Daly, Rettenmeyer and Wille, 1955. Ins. Sociaux 2: 237-246 (techniques for studying
behavior). —Stephen, 1955. Econ. Ent., Jour. 48: 543-548 (alfalfa pollinating spp. in Man.).
—Bohart and Nye, 1956. Gleanings in Bee Culture 84: 265-268, 317, 331-333, 337, 400-405,
468-472, 508, 602-606, 639 (place of bees in the world of insects). —Kerr and Laidlaw, 1956.
Advances in Genetics 8: 109-153 (genetics of bees). —Manning, 1956. Behaviour 9: 114-139
(honey-guides). —Manning, 1956. Royal Physiol. Soc., Proc. 25: 67-71 (floral constancy).
—Leppik, 1957. Evolution 11: 466-481 (coevolution of entomophilous plants and
anthophilous insects). —Michener and Lange, 1957. Kans. Ent. Soc., Jour. 30: 71-80
(ethology of colletid spp.). —Hobbs, 1957. Canad. Ent. 89: 230-235 (alfalfa and red clover as
sources of nectar and pollen). —Linsley and MacSwain, 1957. Calif. Univ. Pubs. Ent. 11:
395-430 (stylopization). —Bohart, 1958. Internat]. Congr. Ent., Proc. 10: 929-937 (alfalfa
pollinating spp.). —Koerber and Medler, 1958. Wis. Acad. Sci. Arts and Letters 47: 58-63
(trap-nest survey of solitary spp. in Wis.). —Linsley, 1958. Hilgardia 27: 543-599, 3 figs., 8
tables (ecology of solitary Apoidea). —Linsley and MacSwain, 1958. Evolution, 12: 219-223
(significance of floral constancy). —Medler, 1958. Ent. News 69: 21-24 (parasitism by
Leucospis affinis Say of trap-nesting spp.). —Michener, 1958. Xth Internatl. Congr. Ent.,
Proc. 2: 441-448 (evolution of social behavior). —Michener, Lange, Bigarella and Salamuni,
1958. Ecology 39: 207-217 (factors influencing distribution of nests in earth banks).
—Michener and Lange, 1958. Science 127: 1046-1047 (primitive social behavior). —Hobbs,
1958 (1956). Tenth Internat]. Congr. Ent. 4: 939-942 (factors affecting value of bees as
pollinators of alfalfa and red clover.). —Evans and Lin, 1959. Wasmann Jour. Biol. 17:
115-132 (predation by Philanthus spp.). —Linsley and Hurd, 1959. Ent. News 70: 63-68
(ethological observations on spp. in Ariz. and N. Mex.). —Linsley and MacSwain, 1959.
Kans. Ent. Soe., Jour. 32: 8 (sound production in nocturnal spp.). —Linsley and MacSwain,
1959. Calif. Univ. Pubs. Ent. 16: 1-46 (ethology of Ranunculus visiting spp.). —Powell and
Chemsak, 1959. Kans. Ent. Soc., Jour. 32: 115-120 (predation by Philanthus spp.).
—Linsley, 1960. N. Y. Ent. Soc., Jour. 68: 13-20 (matinal bees at flowers of Cucurbita,
Ipomoea and Datura). —Evans and Linsley, 1960. South. Calif. Acad. Sci., Bul. 59: 30-37, 1
pl. (sleeping aggregations). —Linsley, 1960. Calif. Univ. Pubs. Ent. 16: 357-392, pls. 48-55
(ethology of bee- and wasp-killing robber flies in Ariz. and N. Mex.). —Hobbs, Nummi and
Virostek, 1961. Canad. Ent. 93: 409-419 (food gathering behavior of honey-, bumble-, and
leaf-cutter bees in Alta.). —Bohart, 1962. 1st Internat]. Symp. Pollination, August 1960.
Swedish Seed Assoc. Copenhagen Publ. Comm., Proc. 7: 181-188 (introduction of foreign
pollinating spp.). —Linsley, 1962. Ent. Soc. Amer., Ann. 55: 148-164 (sleeping
aggregations). —Linsley, 1962. Sartyrck ur Meddelande nr 7 fran Sveriges
Froodlareforbund, pp. 189-197 (ethological adaptations of solitary spp. for pollination of
desert plants). —Michener, 1962. Rev. Biol. Tropical 10: 167-175, 2 figs. (pollen collection
from flowers with tubular anthers). —Linsley and Cazier, 1963. Pan-Pacifie Ent. 39: 1-18, 6

1746 Hymenoptera in America North of Mexico

figs., 2 tables (spp. which take pollen from Solanwm flowers). —Linsley, MacSwain and
Raven, 1963. Calif. Univ. Pubs. Ent. 33: 1-58, 6 pls., 6 text figs. (comparative behavior of
Camissonia and Oenothera spp. of the Colorado Desert and the Great Basin). —Linsley,
MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 59-98, 3 pls. (comparative behavior
of Camissonia and Oenothera spp. of the Mojave Desert). —Michener, 1963. Science 141:
434-435 (division of labor among primitively social spp.). —Wille, 1963. Rev. Biol. Tropical
11: 205-210 (behavioral adaptations of pollen-collecting spp. at flowers of Cassia).
—Michener, 1964. Ins. Sociaux 11: 317-342 (reproductive efficiency in relation to colony
size). —Michener, 1964. Amer. Zool. 4: 227-239 (evolution of nests). —Armitage, 1965.
Kans. Ent. Soc., Jour. 38: 89-100, 4 figs., 4 tables (predation by Philanthus spp.).
—Matthews and Fischer, 1965. North Central Branch, Ent. Soc. Amer., Proc. 19: 79-81, 1
fig. (modified trap-nest). —Fye, 1965. Econ. Ent., Jour. 58: 803-804, 4 figs. (trap-nesting
methods). —Fye, 1965. Canad. Ent. 97: 863-877, 6 figs., 4 tables (ethology of spp. taken in
trap-nests in northwest Ont.). —Parker and Bohart, 1966. Pan-Pacific Ent. 42: 91-98
(host-parasite relationships as determined by use of trap-nests). —Butler, Werner and
Levin, 1966. Kans. Ent. Soc., Jour. 39: 434-436 (safflower visiting spp.). —Levin and
Butler, 1966. Econ Ent., Jour. 59: 654-657, 3 tables (safflower pollinating spp.).
—Krombein, 1967. Trap-nesting wasps and bees, vi and 570 pp., 29 pls., Smithsn. Press
(life histories, nest architecture, nest associates). —Kerfoot, 1967. Amer. Nat. 101: 65-70
(correlation between ocellar size and foraging activities). —Parker and Bohart, 1968.
Pan-Pacific Ent. 44: 1-6 (host-parasite relationships as determined by use of trap-nests).
—Baker and Hurd, 1968. Ann. Rev. Ent. 13: 385-414 (intrafloral ecology). —Batra and
Torchio, 1968. Mycologia 60: 189-190 (N. Amer. records of Ascocphaera apis L.). —May
and Stockhammer, 1968. Kans. Ent. Soc., Jour. 41: 339-341, 1 fig. (mass colonization by use
of artificial substrate). —Kerr, 1969. Ecol. Biol. 3: 119-175 (evolution of social bees).
—Michener, 1969. Ann. Rev. Ent. 14: 299-342, 3 tables (comparative behavior of social
bees). —Bohart, 1970. Ent. Soc. Amer., Bul. 16: 8-9 (management of native spp. for
commercial crop production). —Schlissing, 1970. Ecology 51: 1061-1067 (foraging behavior
in flowers of Ipomoea and Aniseia). —Gerber and Klostermeyer, 1970. Science 167: 82-84
(sex control). —Darchen, 1970. Gaz. Apicult. 754: 48-51 (division of labor in social spp.).
—Mickel, 1970. Minn. Univ. Agr. Expt. Sta. Tech. Bul. 27: 1-77 (references to literature
pertaining to Mutillidae parasitic on Apoidea). —Bohart, 1970. Utah State Univ. 41st
Faculty Honor Lecture, 33 pp. (evolution of parasitism). —Macior, 1970. Amer. Jour. Bot.
57: 716-728 (pollinating spp. of Pedicularis in Colo.). —Hurd, Linsley and Whitaker, 1971.
Evolution 25: 218-234, 4 figs., 3 tables (squash and gourd bees and origin of cultivated
Cucurbita). —Bohart, 1971. Tall timbers Conf. Ecol. Animal Contrib. Habitat
Management, Proc., pp. 253-266, 9 figs. (management of habitats for native spp.).
—Michener and Brothers, 1971. Kans. Ent. Soc., Jour. 44: 236-239, 4 figs. (observation nest
for burrowing spp.). —Batra, 1972. Kans. Ent. Soc., Jour. 45: 208-218 (nest-building
secretions). —Osgood, 1972. Maine life Sci. Agr. Expt. Sta. Tech. Bul. 59: 1-8 (nesting sites
of native spp. associated with low-bush blueberries in Maine). —Bohart, 1972. Ann. Rev.
Ent. 17: 287-312, 4 tables (management of native spp. for crop pollination). —Michener,
1972. Kans. Ent. Soc., Jour. 45: 373-376 (direct food transferring behavior). —Cruden, 1972.
Evolution 26: 363-389 (pollination biology of Nemophila menziesii with comments on
evolution of oligolectic bees). —Cruden, 1973. Amer. Jour. Botany 60: 802-809 (pollination
of Mirabilis). —Thorp, 1973. Pan-Pacific Ent. 49: 89 (robber fly predation). —Batra, Batra
and Bohart, 1973. Mycopath. Mycol. Appl. 49: 13-44 (mycoflora of domesticated and wild
bees). —MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 1-80, 3 pls., 10 figs.,
18 tables (comparative behavior of Clarkia visiting spp.). —Linsley, MacSwain, Raven and
Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 1-68, 6 pls., 15 figs., 10 tables (comparative
behavior of Camissonia and Oenothera visiting spp. in cismontane Calif. and Baja
California). — Vogel, 1973. Umschau 73 (22): 701-702 (collection of fatty oil from plants and
its incorporation into larval food.). —Macior, 1973. Amer. Jour. Bot. 60: 863-871 (pollinating
spp. of Pedicularis on Mt. Rainier). —Torchio, 1973. Kans. Ent. Soc., Jour. 46: 446-453, 1
fig., 4 tables (relative toxicity of insecticides to honeybee, alkali bee and alfalfa leafcutting
bee). —Frankie, 1973. Ent. Soc. Amer., Ann. 66: 690-691, 1 fig. (field technique for marking
bees). —Macior, 1974. Melanderia 15: 1-59 (pollinating spp. of Front Range of Colorado
Rocky Mts.). —Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 1-20, figs. 1-6, tables

Superfamily APOIDEA 1747

1-2 (foraging behavior of spp. visiting flowers of Kallstroemia grandiflora). —Jones and
Buchman, 1974. Anim. Behaviour 22: 481-485, 2 tables (u. v. floral patterns as orientation
guides). —Estes and Thorp, 1974. Torrey Bot. Club, Bul. 101: 272-276 (pollinating spp.
visiting flowers of Ludwigia peploides). —Michener, 1974. The social behavior of the bees,
xli and 404 pp., The Belknap Press of Harvard Univ. Press. —Michener and Brothers,
1974. Natl. Acad. Sci. U. S. A., Proc. 71: 671-674 (queen inhibitory behavior). —Macior,
1974. Missouri Bot. Garden, Ann. 61: 760-769, 21 figs. (coadaptation between flowers and
pollinating spp.). —Moldenke and Neff, 1974. The bees of California: A catalogue with
special relevance to pollination and ecological research, Origin and Structure of
Ecosystems (IBP) Technical Rpts. 74-1 to 74-6, Calif. Univ. Santa Cruz. —Austin and
Oliver, 1974. Arnold Arboreteum Harvard Univ., Jour. 55: 291-299 (pollinators of
Sisyrinchium solstitiale, a Fla. endemic). —Bottema, 1975. Paleohistoria 17: 17-35, 9 figs.,
7 tables (contamination of pollen spectra by burrowing bees in prehistoric settlements).
—Cazier and Linsley, 1975. Pan-Pacifie Ent. 51: 248-253, 6 figs., 2 tables (bee and wasp
visitors to the flowers of Kallstroemia grandiflora after two years drought). —Thorp,
Briggs, Estes and Erickson, 1975. Science 189: 476-478, 1 fig. (nectar flourescense and
foraging efficiency of bees). —Estes and Thorp, 1975. Amer. Jour. Bot. 62: 148-159
(pollination of Pyrrhopappus carolinianus). —Thorp and Estes, 1975. Kans. Ent. Soc.,
Jour. 48: 175-184, 6 figs. (intrafloral behavior of bees on flowers of Cassia fasciculata).
—Heinrich, 1975. Ann. Rev. Ecol. Syst. 6: 139-170 (energetics of pollination). —Heinrich,
1975. Evolution 29: 325-334, 4 figs., 2 tables (bee flowers). —Barrows, Bell and Michener, |
1975. Natl. Acad. Sci. USA, Proc. 72: 2824-2828 (odor differences and their social function).
—Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: iii and 74 pp., 18 figs., 15 tables
(principal Larrea visiting spp. of southwest. U. S.). —Kevan, 1975. Biol. Conservation 7:
301-309, 1 fig., 4 tables (effect of fenitrothion on pollinators of lowbush blueberries).
—Macior, 1975. Amer. Jour. Bot. 62: 1009-1016, 19 figs., 17 tables (pollination of
Delphinium tricorne). —Macior, 1975. Amer. Jour. Bot. 62: 1065-1072, 21 figs., 7 tables
(pollination ecology of Pedicularis in the Yukon Territory). —Evans, 1975. Ent. Soc.
Amer., Ann. 68: 888-892, 6 figs., 3 tables (predation by Philanthus albopilosus Cress.).
—Linsley, 1976. Pan-Pacifie Ent. 52: 177-178 (defensive behavior of males about plants not
visited by their females). —Bouseman, 1976. Pan-Pacific Ent. 52: 178-179 (predation by
Apiomerus crassipes). —Iwata, 1976. Evolution of instinct, comparative ethology of
Hymenoptera, 535 pp., Washington, D. C., Smithsonian Institution (behavior). —Erickson,
Enns and Werner, 1976. Ent. Soc. Amer., Ann. 69: 959-970, 4 tables (bee-associated
Meloidae). —Jander, 1976. Physiol. Ent. 1: 179-194, 8 figs., 2 tables (grooming and pollen
manipulation). —McGregor, 1976. U. S. Dept. Agr., Agr. Handbook 496: 1-411, 196 figs.
(pollination of cultivated crop plants). —Frankie, Opler and Bawa, 1976. Jour. Ecol. 64:
1049-1057, 1 fig., 4 tables (foraging behavior). —Rust and Clement, 1977. Kans. Ent. Soc.,
Jour. 50: 37-48, 5 figs., 4 tables (role in pollination of Collinsia sparsiflora).

Morphology: Braue, 1913. Jenaische Ztschr. Naturwiss. 50: 1-96 (pollen-collecting apparatus).
—Stoeckhert, 1924. Arch. Naturgesch. (A) 90 (2): 109-131 (gynandromorphism). —Kuhn,
1927. Ztschr. vergleich. Physiol. 5: 762-800 (color vision). —Pessotzkaya, 1929. Soe. Nat.
Leningrad. Trav. 59: 21-46 (gland apparatus in instinctive behavior). —Beck, 1933. Utah
Acad. Sci., Proc. 10: 89-137, 8 pls. (male genitalia). —Michener, 1943. Pan-Pacifie Ent. 19:
96-100 (homologies between male and female appendages). —Michener, 1944. Ent. Soe.
Amer., Ann. 37: 336-351 (appendages of eighth and ninth abdominal segments).
—Michener, 1944. Amer. Mus. Nat. Hist., Bul. 82: 158-225 (comparative external
morphology). —Auclair and Jamieson, 1948. Science 108: 357-358 (amino acids in pollen
collected by bees). —Wille, 1956. Kans. Univ. Sci. Bul. 38: 439-499 (thoracic musculature).
—Wille, 1958. Ent. Soc. Amer., Ann. 51: 538-546, 24 figs. (dorsal vessel). — Altenkirch,
1962. Zool. Beitrag (n. f.) 7: 161-238 (abdomen). —Michener, 1965. Amer. Mus. Nat. Hist.,
Bul. 130: 27-32, figs. 2-17 (morphological terminology). —Cruz-Landim, 1967. Arq. Zool. S.
Paulo 15: 177-290 (glands). —Rothenbuhler, Kulincevic and Kerr, 1968. Ann. Rev. Genetics
2: 413-438 (genetics). —Graf, 1968. Bol. Univ. Federal Parana, Zool. 3: 65-78 (salivary
gland). —Stephen, Bohart and Torchio, 1969. Oreg. State Univ. Agr. Expt. Sta., pp. 3-31,
figs. 1-112 (external morphology). —Tanabe, Tamaki and Nakamo, 1970. Japanese Jour.
Genetics 45: 425-428 (variation in esterase isozymes). —Tulloch, 1970. Lipids 5: 247-258

1748 Hymenoptera in America North of Mexico

(composition of beeswax). —Lello, 1971. Kans. Ent. Soc., Jour. 44: 5-20, 21 figs. (adnexal
glands of sting apparatus). —Lello 1971. Ciencia e Cultura 23: 253-258 (adnexal glands of
sting apparatus). — Wille, 1971. Rev. Biol. Tropical 18: 33-51 (musculature of salivary
syringe and neck region). —Kerr, 1972. Kans. Ent. Soc., Jour. 45: 111-122, 20 figs.
(chromosome numbers). —Almeida Correia, 1973. Faculdade Cienc. Porto Univ., An. 56:
67-175 (morphological and morphometric study of mouthparts of principal genera).

— Almeida Correia, 1973. Inst. Zool. “Dr. Augusto Nobre” Facul. Cienc. Porto 118: 1-117, 8
pls. (mouthparts). —Cruz-Landim, 1973. Studia Ent. 16: 209-215, 2 figs., 1 table (thoracic
salivary glands). —Iuga, 1973. Mus. Hist. Nat. “Grigore Antipa”, Trav. 13: 203-226, 26 figs.
(apical abdominal appendages). —Michener, 1974. The social behavior of the bees, Chapter
1: 3-19 (development, structure and function). —Snyder, 1975. Evolution 28: 687-689
(allozymie variability). —Pasteels and Pasteels, 1975. Arch. Biol., Bruxelles 86: 453-466, 13
figs. (stereoscan studies of pollen collecting scopae of Fideliidae). —Lello, 1976. Kans. Ent.
Soe., Jour. 49: 85-99, 22 figs., 3 tables (adnexal glands of sting apparatus). —Pasteels and
Pasteels, 1976. Arch. Biol., Bruxelles 87: 79-102, 25 figs. (stereoscan studies of pollen
collecting scopae of Colletidae and Oxaeidae).

Family COLLETIDAE

This fainily differs from all other bees in that the females line their cells with a cel-
lophane-like substance. The glossa is short, usually truncate or bifid, and this wasp-like feature
is the primary basis for considering the Colletidae to be the most primitive family of living bees.
Although the Colletidae are found throughout much of the world, the family is especially abun-
dant and diverse in the southern hemisphere being exceptionally well represented in Australia
by a large fauna consisting chiefly of the subfamilies Colletinae (Paracolletini), Euryglossinae
and Hylaeinae.

In North America the family is represented by four subfamilies of which the nominate tribe
of the Colletinae (Colletes) and the Hylaeinae (Hylaews) comprise the largest assemblage of the
family in America north of Mexico. Only a single species of the Paracolletini (Eulonchopria
punctatissima Michener) is known to range north of Mexico into the United States (Arizona).
The other two subfamilies present in North America are the Xeromelissinae and Diphaglossinae
which occur only in the New World and are chiefly centered in the Neotropical Region.
Representatives of these subfamilies have been found as far north as southern Mexico
(Xeromelissinae) and the southern United States (Diphaglossinae).

While most of the species in America north of Mexico depend upon a comparatively wide
variety of flowers for nectar and pollen, the females of a number of species of Colletes are either
oligoleges or exhibit narrow polylectic tendencies.

SUBFAMILY COLLETINAE
TRIBE PARACOLLETINI

Genus EULONCHOPRIA Brethes

Eulonchopria Brethes, 1909. Mus. Nac. Buenos Aires, An. 19: 247.
Type-species: Eulonchopria psaenythioides Brethes. Monotypic.
punctatissima Michener. Southern Ariz.; Mexico (Guerrero). Pollen: Unknown, but visits

flowers of Acacia and Baccharis.
Eulonchopria punctatissima Michener, 1963. Ent. Soc. Amer., Ann. 56: 847, fig. 1-2, 7-9. °,
3.

TRIBE COLLETINI

Genus COLLETES Latreille

Colletes Latreille, 1802. Hist. Nat. Fourmis, p. 423.

Type-species: Apis succincta Linnaeus. Monotypic.
Evodia Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 207.

Type-species: Apis succincta Linnaeus. Monotypic. (=Apis calendarwm Panzer).
Colletes subg. Rhinocolletes Cockerell, 1910. Entomologist 43: 242.

Superfamily APOIDEA 1749

Type-species: Colletes nasutus Smith. Monotypic.
Colletes subg. Ptilopoda Friese, 1921. Stettin. Ent. Ztg. 82: 83.
Type-species: Colletes spiloptera Cockerell. Monotypic. (=Colletes maculipennis
Friese).
Colletes subg. Denticolletes Noskiewicz, 1936. Prace Nauk. Wydawnictwo Towarzystwa
Nauk. Lwowie (2) 3: 486.
Type-species: Colletes graeffei Alfken. Monotypic.
Colletes subg. Puncticolletes Noskiewicz, 1936. Prace Nauk. Wydawnictwo Towarzystwa
Nauk. Lwowie (2) 3: 490. Proposed without a type-species designation and therefore
invalid under article 13b of International Rules of Zoological Nomenclature.

Several of the subgenera cited in the above synonymy are doubtless valid, but as the sub-
generic groupings of Colletes have not been properly worked out they are listed as synonyms.

Some species are polytropic, others oligotropic. Several of the principally autumnal species
have small vernal broods.

Revision: Stephen, 1954. Kans. Univ. Sci. Bul. 36: 149-527, 87 figs., 8 maps (U. S. spp.).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 24-59, figs. 7-10 (eastern U. S. spp.).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 20-22, figs. 1-3 (Larrea visiting
spp.).
SPECIES GROUP PRODUCTUS

arizonensis Stephen. Ariz. mts.
Colletes arizonensis Stephen, 1954. Kans. Univ. Sci. Bul. 36: 204. 3, @.
cercidii Timberlake. Calif., Ariz. Pollen: Unknown, but visits flowers of Cercidiwm
torreyanum.
Colletes cercidii Timberlake, 1951. Wasmann Jour. Biol. 9: 198. 6, °.
gilensis Cockerell. Colo., Tex., N. Mex., Ariz.; Mexico. Pollen: Unknown, but visits flowers of
Melilotus alba, Petalostemon candidus, P. flavescens, P. oligophyllus, Potentilla
thurberi, Solidago canadensis.
Colletes gilensis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 41. ¢.
perileucus Cockerell. Calif., Ariz., Tex., Mexico. Pollen: Unknown, but visits flowers of
Melilotus alba, Prosopis glandulosa.
Colletes perileucus Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 535. 9.
productus Robertson. Mass., west to Wis., south to Ga. and Ala. Pollen: Unknown, but visits
flowers of Apocynum cannabinum, Batodendron, Rhus copallina, R. glabra, Xolisma
ligustrina.
Colletes producta Robertson, 1891. Amer. Ent. Soc., Trans. 18: 62. ¢.
rudis Timberlake. Calif. (near Indio). Pollen: Unknown, but visits flowers of Encelia farinosa.
Colletes rudis Timberlake, 1951. Wasmann Jour. Biol. 9: 197. d.
skinneri Viereck. Ariz., N. Mex.
Colletes skinneri Viereck, 1903. Amer. Ent. Soc., Trans. 29: 58. 2, 6.
vandykei Timberlake. Ariz. (Santa Rita Mts.).
Colletes vandykei Timberlake, 1951. Wasmann Jour. Biol. 9: 200. d.

SPECIES GROUP COMPACTUS

compactus compactus Cresson. N.S. to Ga., west to Wis., Mo., Colo., and Ariz. Parasite:
Epeolus pusillus Cress., Epeolus autumnalis Robt.? Pollen: Compositae, especially
flowers of Astereae; Heliantheae; Helenieae.
Colletes compactus Cresson, 1868. Boston Soc. Nat. Hist., Proc. 12: 166. 2, ¢.

Biology: Rau and Rau, 1916. Anim. Behavior, Jour. 6: 367. —Rau, 1922. Acad. Sci. St. Louis,
Trans. 24 (7): 37. —Rozen and Favreau, 1968. N. Y. Ent. Soc., Jour. 76: 106-111.

compactus hesperius Swenk. B. C., and Idaho south to Calif., Ariz.; Mexico.
Colletes hesperius Swenk, 1906. Ent. News 12: 257. °, 6.

1750 | Hymenoptera in America North of Mexico

SPECIES GROUP LATITARSIS

latitarsis Robertson. Mich. to Fla., west to Mont., Colo., Tex., Ariz. Parasite: Epeolus
bifasciatus Cresson? Pollen: Unknown, but visits flowers of Asclepias incarnata, A.
syriaca, Campanula americana, Ceanothus americanus, Cicuta maculata, Lycopus
americanus, Medicago sativa, Melilotus alba, Passiflora lutea, Physalis lanceolata, P.
subglabrata, P. virginiana, Polygonum hydropiperoides, Pycnanthemum flexuosum,
Solidago, Symphoricarpos occidentalis, Trifolium.
Colletes latitarsis Robertson, 1891. Amer. Ent. Soc., Trans. 18: 60. 2, 3.
punctipennis maurus Stephen. Tex. (Brownsville). The typical punctipennis Cresson occurs
only in Mexico.
Colletes punctipennis maurus Stephen, 1954. Kans. Univ. Sci. Bul. 36: 232, fig. 11. d.

SPECIES GROUP SIMULANS

angelicus Cockerell. Calif. Ecology: Possibly produces two generations annually. Pollen:
Unknown, but visits predominantly fall blooming composites including but not restricted
to Baccharis emoryi, Gutierrezia sarothrae, Haplopappus venetus, H. vernonioides,
Hemizonia paniculata. Males and females have also been taken at the flowers of an
unidentified Eriogonum.

Colletes angelicus Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 32. ¢.
birkmanni Swenk. Kans., Okla., Tex., N. Mex. Pollen: Unknown, but visits flowers of Rhus
microphylla.

Colletes birkmanni Swenk, 1906. Ent. News 17: 259. °, 6.
bryanti Timberlake. Ariz., N. Mex.
Colletes bryanti Timberlake, 1951. Wasmann Jour. Biol. 9: 208. 6, 9.
delodontus Viereck. N. Mex. Pollen: Unknown, but visits flowers of Rorripa nasturtium.
Colletes delodontus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 59. °.
eulophi Robertson. South. half of U. S., west to Colo. and Ariz. Parasite: Epeolus minimus
(Robt.). Pollen: Unknown, but visits flowers of Aster, Ceanothus fendleri, C.
americanus, Chrysanthemum leucanthemum, Clematis, Eupatorium perfoliatum,
Melilotus, Rubus, Solidago nemoralis, Sophia obtusa.
Colletes eulophi Robertson, 1891. Amer. Ent. Soc., Trans. 18: 61. 2, d.
Colletes illinoiensis Robertson, 1891. Amer. Ent. Soc., Trans. 18: 62. 2.
fulgidus fulgidus Swenk. B. C., Mont., Alta., S. Dak. south to Tex. and Calif. Pollen: Unknown,
but visits flowers of Aster, Baccharis emoryi, Calochortus luteus, Chaenactis sterioides,
Chrysothamnus, Clarkia, Coreopsis lanceolata, Cryptantha intermedia, Encelia
farinosa, Erigeron, Eriogonum fasciculatum, E. latifolium nudum, Eriophyllum
confertifolium, Grindelia, Gutierrezia californica, G. sarothrae, Haplopappus
arborescens, H. parishii, H. vernonioides, Medicago sativa, Melilotus alba, Rhus
laurina, Solidago multiradiata, Sphenosciadium capitellatum.
Colletes fulgidus Swenk, 1904. In Viereck, Canad. Ent. 36: 95. 2, 6.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1018, figs. 27, 30-33, 35 (larva).

fulgidus longiplumosus Stephen. Coastal north. Calif. and San Joaquin Valley. Parasite:
Epeolus sp. A (Rozen). Pollen: Unknown, but visits flowers of Armeria californica,
Clarkia rubicunda, Convolvulus, Eriophyllum staechadifolium, Grindelia latifolia,
Haploppapus, Layia, Solidago occidentalis.
Colletes fulgidus longiplumosus Stephen, 1954. Kans. Univ. Sci. Bul. 36: 248. 5, 2.
kincaidii Cockerell. N. S. south to N. C. west to B. C., Calif., Ariz. Pollen: Unknown, but visits
flowers of Chamaenerion angustifolium, Eriogonum mariifolium, E. subscaposum,
Heracleum, Horkelia bernardina, Medicago sativa, Melilotus alba, Mentha arvensis,
Penstemon, Potentilla, Solidago californica, S. confinis, Sphenosciadium capitellatum,
Symphoricarpos.
Colletes kincaidii Cockerell, 1898. Acad. Nat. Sci. Phila., Proc. 50: 52. 9, ¢.
Colletes sieverti Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 424. ¢.
Colletes eulophi albertensis Cockerell, 1938. Canad. Ent. 70: 70. 2.
Colletes stricklandi Cockerell, 1938. Canad. Ent. 70: 70. °.

Superfamily APOIDEA 1751

louisae Cockerell. Tex. west to south. Calif.; Mexico. Pollen: Polylectic, produces two
generations annually, the spring brood visits various Compositae (e.g., Baileya, Encelia,
Geraea, Lasthenia, Malacothrix) and desert shrubs including Agave, Cercidiwm and
Larrea. The autumnal brood obtains pollen exclusively from Compositae (e.g.,
Chrysothamnus, Haplopappus, Lepidospartum). Predator: Philanthus gibbosus (Fabr.).
Colletes louisae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 46. 6.
Colletes tucsonensis Cockerell, 1906. Canad. Ent. 38: 163. 6.

rufocinctus Cockerell. Minn. west to Alta., south to S. Dak., Colo., Ariz. Pollen: Polylege of
Compositae including Aster paniculatus, Grindelia squarrosa, Helianthus petiolaris,
Solidago canadensis, S. rigida, Taraxacum vulgare.
Colletes rufocinctus Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 298. ¢.
Colletes truncatus Timberlake, 1943. Amer. Mus. Nat. Hist., Bul. 81: 396. 9.
simulans armatus Patton. N.S. south to N. C., west to Nebr. and Alta. Pollen: Oligolege of
autumnal flowering Compositae including Aster ericoides, Baccharis, Bidens aristosa,
Eupatorium, Solidago canadensis, S. graminifolia, S. nemoralis, S. ulmifolia, but also
visits Cicuta maculata, Polygonum hydropiperoides for nectar. Predator: Philanthus
albopilosus Cress.
Colletes armata Patton, 1879. Boston Soe. Nat. Hist., Proc. 20: 143. 2, 6.
Colletes scitula Patton, 1879. Boston Soe. Nat. Hist., Proc. 20: 144. ¢.
Colletes spinosa Robertson, 1891. Amer. Ent. Soc., Trans. 18: 60. 2, 6.

Biology: Evans, 1975. Ent. Soc. Amer., Ann. 68: 891 (predator).

simulans miamiensis Mitchell. Fla. (Miami). Intergrades between Colletes s. simulans and C.
s. miamiensis occur in Miss. and Tex. Pollen: Presumably oligolege of autumnal
flowering Compositae, but has been collected only on one occasion at flowers of Bidens
pilosa.

Colletes simulans miamiensis Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 236. 6.
simulans nevadensis Swenk. B. C., Wash., Oreg., Calif., Nev., Utah. Pollen: Oligolege of
autumnal flowering Compositae including Baccharis emoryi, Chrysothamnus
vicidiflorus, Gutierrezia californica, G. confinis, G. sarothrae, Haplopappus acradenius,
H. bernardina, H. pinifolius, H. vernonioides, Lepidospartum squamatum, Solidago
lucida, but also visits Eriogonum and Melilotus alba for nectar.
Colletes nevadensis Swenk, 1908. Nebr. Univ., Dept. Ent. Contrib. 1: 52. ¢.

simulans simulans Cresson. Nebr. west to B. C., Colo., Ariz. Pollen: Oligolege of autumnal
flowering Compositae including Aster, Bigelovia wrightii, Grindelia, Senecio.
Colletes simulans Cresson, 1868. Boston Soc. Nat. Hist., Proc. 12: 168. d.
Colletes bigeloviae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 40. 9.
Colletes brevispinosus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 62. 2.
Colletes tegularis Swenk, 1905. Canad. Ent. 37: 304. °.
Colletes coloradensis Cockerell, 1933. Ent. Soc. Amer., Ann. 26: 41. ¢.
slevini Cockerell. B. C. to Wyo., south to Calif. and Ariz.; Mexico (Baja California). Pollen:
Polylectic, visiting a wide variety of flowers including Adenostoma fasciculatum, A.
sparsifolium, Amorpha fruticosa, Aster, Baccharis pilularis, Bigelovia, Ceanothus
integerrimus, C. palmeri, Chrysanthemum, Chrysothamnus pumilis, Cirsium,
Eriogonum elatum, E. elongatum, E. fasciculatum, E. latifolium nudum, E.
plumatella, E. subscaposum, E. wrightii, Euphorbia albomarginata, Fremontia
californica, Gutierrezia californica, G. lucida, Haplopappus palmeri, H. pinifolius, H.
vernonioides, Heteromeles arbutifolia, Lepidium virginicum, Malvastrum thurberi,
Melilotus, Nolina, Phacelia frigida, Rhamnus californica, Rhus trilobata, Ribes roezlii,
Solidago californica, Sphenosciadium capitellatum, Stephanomeria exigua.
Colletes slevini Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 185. @.
Colletes eriogoni Cockerell, 1939. Pan-Pacific Ent. 15: 188. d, 2.

trigonatus Cockerell. Colo.

Colletes trigonatus Cockerell, 1933. Ent. Soc. Amer., Ann. 26: 42. °.
utilis Cockerell. N. Mex.

Colletes utilis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 39. 9.

1752 Hymenoptera in America North of Mexico

wootoni Cockerell. Tex., N. Mex., Ariz. Pollen: Polylectic, visits flowers of Amorpha fruticosa,
Dasylirion wheeleri, Helenium autumnale, H. laciniatum, Isocoma, Larrea tridentata,
Melilotus alba, Nolium microcarpa, Phacelia, Petalostemon flavescens, Sapindus
drummondi, Sphaeralcea laxa, Solidago occidentalis, Wislizenia refracta.
Colletes wootoni Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 42. 6.
Colletes apacheorum Timberlake, 1943. Amer. Mus. Nat. Hist., Bul. 81: 401. °.

SPECIES GROUP AESTIVALIS

aestivalis Patton. Mass. south to N. C. west to Tenn., Ill. Parasite: Epeolus interruptus Robt.?
Pollen: Unknown, but visits flowers of Apocynum cannabinum, Heracleum lanatum,
Heuchera americana, H. hispida, Krigia amplexicaulis, Polytaenia nuttallii, Rhus
glabra, Ribes gracile, Rubus villosus, Taenidia integerrima, Zizia aurea.
Colletes aestivalis Patton, 1879. Boston Soc. Nat. Hist., Proc. 20: 142. 2.
Colletes heucherae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 61. 2, d.
andrewsi Cockerell. Nebr., Colo., Wis., S. Dak., Man. Pollen: Apparently an oligolege of
Heuchera.
Colletes Andrewsi Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 311. 9.

SPECIES GROUP ROBERTSONII

kansensis Stephen. Kans. Pollen: Unknown, but visits flowers of Monarda punctata
occidentalis.
Colletes kansensis Stephen, 1954. Kans. Univ. Sci. Bul. 36: 297, fig. 29. d.
larreae Timberlake. South. Calif., Ariz. Pollen: Oligolege of Larrea tridentata; males also have
been collected at flowers of Acamptopappus sphaerocephalus, Barbarea orthoceras.
Colletes larreae Timberlake, 1951. Wasmann Jour. Biol. 9: 193. d, 2.
metzi Timberlake. Colo. (Boulder).
Colletes metzi Timberlake, 1951. Wasmann Jour. Biol. 9: 192. 6.
robertsonii Dalla Torre. Ill. and Miss., west to Man., Colo., and N. Mex. Pollen: Unknown, but
visits flowers of Amorpha canescens, Monarda punctata occidentalis, Petalostemon
candidus, P. purpureum, P. violaceus.
Colletes punctata Robertson, 1891. Amer. Ent. Soc., Trans. 18: 92. ¢. Preoce.
Colletes robertsonii Dalla Torre, 1896. Cat. Hym., v. 10, p. 44. N. name.
Colletes robustus Swenk, 1904. Ent. News 15: 251. @, 6.
timberlakei Stephen. Colo., Wyo. Pollen: Unknown, but visits flowers of Petalostemon.
Colletes timberlakei Stephen, 1954. Kans. Univ. Sci. Bul. 36: 290, fig. 31. ¢.
turgiventris Timberlake. Calif. Pollen: Unknown, but visits flowers of Phacelia including P.
hispida.
Colletes turgiventris Timberlake, 1951. Wasmann Jour. Biol. 9: 196. d, °.

SPECIES GRouP NUDUS

brimleyi Mitchell. N. J., N. C., Ga., Fla. Pollen: Unknown, but visits flowers of Ilex cassine.
Colletes brimleyi Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 231, figs. 14, 15. @, 6.
nudus Robertson. Ont. south to N. C. and La., west to Wis., Colo. Pollen: Unknown, but visits
flowers of Asclepias syriaca, Ceanothus, Daucus carota, Hydrangea arborescens,
Melilotus alba, Monarda mollis, Parthenocissus, Pycnanthemum flexuosum, Rhus
copallina, R. glabra, Symphoricarpos orbiculatus, S. vulgaris, Verbena urticifolia.
Colletes nudus Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 43. 2, 6.
Colletes Vierecki Swenk, 1905. Canad. Ent. 37: 301. °.
Colletes hydrophilus Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 313. d.

SPECIES GROUP AMERICANUS

aberrans Cockerell. Mich., west to Alta., Colo., and N. Mex. Pollen: Unknown, but visits
flowers of Melilotus, Petalostemon flavescens, P. villoswm.
Colletes aberrans Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 44. @.
albescens Cresson. Wis. and IIl., west to Man., Utah, and N. Mex. Pollen: Unknown, but visits
flowers of Amorpha canescens, Petalostemon purpureum.
Colletes albescens Cresson, 1868. Boston Soc. Nat. Hist., Proc. 12: 168. 2.

Superfamily APOIDEA 1753

americanus Cresson. Que., Man. and United States east of Rocky Mts. Parasite: Epeolus
pusillus Cress.? Pollen: Possibly oligolege of autumnal flowering Compositae, but has
been collected from these and wide range of other plants including Aster, Boltonia
asteroides, Chrysopsis microcephala, Eupatorium altissimum, Gnaphalium
polycephalum, Helianthus divaricatus, Lactuca floridana, Lespedeza virginica, Lycopus
americanus, Polygonum hydropiperoides, P. scandens, Rudbeckia trilobata, Sium
cicutaefolium, Solidago. Predator: Philanthus solivagus Say.
Colletes americana Cresson, 1868. Boston Soc. Nat. Hist., Proce. 12: 167. 2, 6.
annae annae Cockerell. N. Mex., Ariz. Pollen: Unknown, but visits flowers of Baccharis,
Solidago canadensis arizonica.
Colletes annae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 48. 3.
Colletes crucis Cockerell, 1902. Ent. News 13: 304. ¢.
Colletes cockerelli Timberlake, 1951. Wasmann Jour. Biol. 9: 221. °.
annae disseptus Timberlake. South Calif.; Mexico (Baja Calif.). Pollen: Possibly oligolege of
autumnal flowering Compositae including Conium maculatum, Lepidospartum
squamatum, Solidago.
Colletes disseptus Timberlake, 1951. Wasmann Jour. Biol. 9: 223. 2, ¢.
bradleyi Mitchell. N. J. (Chatsworth and Chesilhurst).
Colletes bradleyi Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 239. @.
gypsicolens Cockerell. B. C. to Calif., southeast to Colo. and N. Mex. Pollen: Possibly oligolege
of autumnal flowering Compositae including Aster, Chrysothamnus nauseosus,
Haplopappus vernonioides.
Colletes gypsicolens Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 47. ¢.
howardi Swenk. N. C. (Southern Pines). Parasite: Epeolus howardi Mitchell? Pollen: Unknown,
but visits Kuhnistera pinnata.
Colletes howardi Swenk, 1925. Amer. Mus. Novitates 186: 5. 2, 6.
laticinctus Timberlake. Nebr. west to Idaho, south to N. Mex. and Ariz. Pollen: Unknown, but
visits Gutierrezia sarothrae, Pectis papposa, Polygonum.
Colletes laticinctus Timberlake, 1951. Wasmann Jour. Biol. 9: 220. °.
mandibularis Smith. B. C., N.S., U. S., east of the Rocky Mts. Pollen: Unknown, but visits
flowers of Amorpha canadensis, Amphiachyris, Berlandiera subcaulis, Cassia
fasciculata, Ceanothus, Cirsium, Erigeron quercifolius, E. ramosus, Flaveria linearis,
Heterotheca subaxillaris, Isopappus divaricatus, Medicago sativa, Melilotus alba, M.
indica, Prionopsis ciliata, Pycnanthemum flexuosum.
Colletes mandibularis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 5. 6.
Colletes similis Robertson, 1904. Canad. Ent. 36: 276. 2. Preocce.
Colletes simulator Michener, 1951. In Muesebeck et al., U. S. Dept. Agr., Agr. Monog. 2:
1048. N. name.
micheneri Stephen. Nebr. (Halsey).
Colletes micheneri Stephen, 1954. Kans. Univ. Sci. Bul. 36: 357. d, 2.
mitchelli Stephen. U. S. east of Great Plains, except New England. Pollen: Unknown, but
visits flowers of Aster, Erigeron quercifolius, Haplopappus, Melilotus alba, Penstemon,
Solidago. It has also been collected at honey dew of Phylloxera on Quercus alba.
Colletes mitchelli Stephen, 1954. Kans. Univ. Sci. Bul. 36: 325, fig. 38. ¢, 2.
ochraceus Swenk. Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Eriogonum, Haplopappus vernonioides, Wislizenia refracta.
Colletes ochraceus Swenk, 1906. Canad. Ent. 38: 42. °.
saritensis Stephen. Tex. (near Sarita). Pollen: Unknown, but visits flowers of Dalea grisea.
Colletes saritensis Stephen, 1954. Kans. Univ. Sci. Bul. 36: 341, fig. 48. d, 9°.
solidaginis Swenk. Mass. to Ga., west to Minn. and Nebr. Pollen: Unknown, but visits flowers
of Solidago.
Colletes solidaginis Swenk, 1906. Canad. Ent. 38: 40. °, ¢.
susannae Swenk. Wis. and IIl., west to Colo. and Alta. Pollen: Unknown, but visits flowers of
Helianthus, Kuhnistera purpurea, Petalostemon purpureum, P. violaceus.
Colletes susannae Swenk, 1925. Amer. Mus. Novitates 186: 1. 9, 6.

1754 Hymenoptera in America North of Mexico

tectiventris Timberlake. Calif. Pollen: Possibly an oligolege of autumnal flowering Compositae,
visiting flowers of Eriogonum, Gutierrezia lucida and Haplopappus acradenius.
Colletes tectiventris Timberlake, 1951. Wasmann Jour. Biol. 9: 218. 9°, ¢.
thysanellae Mitchell. Fla., Ga., N. C., Va. Pollen: Unknown, but visits flowers of Aster,
Baccharis, Eupatorium, Solidago, and possibly Thysanella.
Colletes thysanellae Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 238, figs. 18-20. °,
é.

wilmattae Cockerell. Wis., Minn., Iowa, Man., Nebr., Tex., N. Mex. Pollen: Possibly an oligolege
of Petalostemon having been collected only at the flowers of Petalostemon candidum,
P. flavescens, P. oligophyllum.
Colletes wilmattae Cockerell, 1904. Canad. Ent. 36: 14. 2.

SPECIES GROUP HYALINUS

banksi Swenk. Mich. and N. Y. south to Fla. Pollen: Unknown, but visits flowers of
Batodendron, Ilex glabra.
Colletes banksi Swenk, 1908. Nebr. Univ., Dept. Ent. Contrib. 1: 19. 2.
Colletes ilicis Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 234, figs. 12, 13. 2, d.
distinctus Cresson. N. C., Ga., Fla. Pollen: Unknown, but visits flowers of Hypericum, Ilex
cassine, Melilotus alba.
Colletes distinctus Cresson, 1868. Boston Soe. Nat. Hist., Proc. 12: 167. 3.
Colletes nitidus Smith, 1879. Deser. New Species Hym. Coll. Brit. Mus., p. 1. 2, d.
Colletes carolinus Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 233. °.
hyalinus gaudialis Cockerell. South. Calif. Pollen: Unknown, but visits flowers of Asclepias
eriocarpa, Baccharis emoryi, Brassica, Eriogonum latifolium, Gutierrezia sarothrae,
Haplopappus vernonioides, Heliotropium curassavicum, Hemizonia corymbosa,
Melilotus alba, M. indica, Oenothera, Solidago.
Colletes gandialis Cockerell, 1905. South. Calif. Acad. Sci. Bul. 4: 32.
Colletes gaudialis Cockerell, 1905. South. Calif. Acad. Sci. Bul. 4: 106. Emend.
Colletes arenicola Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 18: 626. °.
hyalinus hyalinus Provancher. N.S. and Maine, west to Alta., Yukon, Colo., and N. Mex.
Pollen: Unknown, but visits flowers of Achillea, Cirsium, Epilobium, Frasera, Kalmia,
Melilotus, Rhus, Solidago. Predator: Philanthus albopilosus Cress.
Colletes hyalinus Provancher, 1888. Addit. Corr. Fauna Ent. Canada, Hym. p. 303. 2, 6.
Colletes spurcus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 58. 2, 6.
Colletes fraserae Swenk, 1908. Nebr. Univ., Dept. Ent. Contrib. 1: 41. 6.
Colletes cauponarius Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 297. 9.
Colletes nitidicaudus Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 299. ¢.

Biology: Evans, 1975. Ent. Soc. Amer., Ann. 68: 891 (predator).
hyalinus oregonensis Timberlake. Coastal Oreg. and Calif. Pollen: Unknown, but visits
flowers of Achillea borealis arenicola, Armeria californica, Baccharis pilularis, Cakile
edentula, Erigeron glaucus, Eriogonum, Polygonum paronychium, Potentilla, Silybum
marianum.
Colletes hyalinus oregonensis Timberlake, 1951. Wasmann Jour. Biol. 9: 211. °, ¢.
lutzi interior Timberlake. Northeast. Calif., Nev., Oreg., Wash., Idaho. Pollen: Unknown, but
visits flowers of Melilotus alba.
Colletes monticola interior Timberlake, 1951. Wasmann Jour. Biol. 9: 214. 2, ¢d.
lutzi lutzi Timberlake. Idaho, Wyo., Utah, Colo., N. Mex. Pollen: Unknown, but visits flowers
of Chrysothamnus, Cleome serrulata, Melilotus.
Colletes lutzi Timberlake, 1943. Amer. Mus. Nat. Hist., Bul. 81: 390. 2, d.
lutzi monticola Timberlake. Calif. (Sierra Nevada Mts.). Pollen: Unknown, but visits flowers
of Allium, Aster adscendens, Castanopsis, Chrysothamnus, Hemizonia corymbosa,
Solidago elongata.
Colletes monticola monticola Timberlake, 1951. Wasmann Jour. Biol. 9: 212. 2, 6.
Colletes inyoensis Timberlake, 1951. Wasmann Jour. Biol. 9: 215. 2.

Superfamily APOIDEA 1755

lutzi pinorum Timberlake. South Calif. (San Bernardino and San Jacinto mts.). Pollen:
Unknown, but visits flowers of Aster, Cryptantha, Eriogonum molestum var.
davidsoniu, Gnaphalium thermale, Hemizonia wheeleri, Layia platyglossa.
Colletes monticola pinorum Timberlake, 1951. Wasmann Jour. Biol. 9: 215. 2, d.
phaceliae Cockerell. B. C. to Man., south to Iowa, Kans., N. Mex., Ariz., Calif. Pollen:
Unknown, but visits flowers of Aster, Cleome serrulata, Epilobium spicatum, Grindelia,
Helenium lacinatum, Malvastrum coccineum, M. cockerelli, Melilotus alba,
Petalostemon, Phacelia, Psoralea tenuiflora, Ribes, Senecio douglasii, Solidago,
Sphenosciadium capitellatum, Tetradymia spinosa.
Colletes phaceliae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 315. @.
Colletes salicicola geranii Cockerell, 1906. Canad. Ent. 38: 163. 3.

SPECIES GROUP INAEQUALIS

inaequalis Say. N.S. to Ga., west to Sask., Kans., Tex., Idaho and Wash. Parasite: Tricrania

sanguinipennis Say. Pollen: Polylectic, visits flowers of Acer, Aesculus glabra sargentii,
Anemone virginiana, Arctostaphylos, Cercis canadensis, Claytonia palustris, Dendrium
buxifolium, Dentaria, Lomatium foeniculaceum, Prunus, Pyrus ioensis, Rhamnus
utilis, Rhus aromatica, R. canadensis, Ribes, Rubus, Salix, Spiraea vanhouttii, S.
thunbergi, Taraxacum, Viburnum acerifolium.

Colletes inaequalis Say, 1837. Boston Jour. Nat. Hist. 1: 391. 2, 6.

Colletes propinqua Cresson, 1868. Boston Soe. Nat. Hist., Proc. 12: 165. 2.

Colletes canadensis Cresson, 1868. Boston Soe. Nat. Hist., Proc. 12: 166. 3.

Colletes inaequalis ferrugineus Swenk, 1908. Nebr. Univ., Dept. Ent. Contrib. 1: 32. 2, 3.

Biology: Smith, 1898. Ent. News 9: 157 (as compactus). —Smith, 1900. N. Y. Ent. Soc., Jour.
8: 208 (as compactus). —Smith, 1901. N. Y. Ent. Soc., Jour. 9: 30 (as compactus). —Smith,
1901. N. Y. Ent. Soc., Jour. 9: 134. —Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7): 38.
—Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 219. —Stephen, 1954. Kans. Univ. Sci. Bul. 36:
155.

thoracicus Smith. Mass. to Fla., west to Okla. and Tex. Parasite: Tricrania sanguinipennis
Say. Pollen: Polylectic, visits flowers of Amelanchier, Aronia, Brassica, Ilex, Malus,
Melilotus, Padus, Prunus angustifolia, Salix, Vaccinium.
Colletes thoracica Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 5. 2, 6.
Colletes rufithorax Swenk, 1906. Canad. Ent. 38: 42. ?, 6.
Colletes pulcher Swenk, 1906. Canad. Ent. 38: 43. 6.

Biology: Parker and Boving, 1924. U.S. Natl. Mus., Proc. 64: 1-40. —Stephen, 1954. Kans.
Univ. Sci. Bul. 36: 156.
validus Cresson. N. B. to N. C., west to Mich. Ecology: A vernal species of wet woods and
swamplands. Pollen: Unknown, but visits flowers of Chamaedaphne calyculata,
Leucothoe racemosa, Prunus, Ribes oxyacanthoides, R. rubrum, Vaccinium, V.

corymbosum.
Colletes validus Cresson, 1868. Boston Soc. Nat. Hist., Proe. 12: 165. 2, d.

SPECIES GROUP IMPUNCTATUS

impunctatus lacustris Swenk. N. B. and N. H. west to B. C., Yukon, N. W. T., Alaska. Pollen:
Unknown, but possibly visits flowers of Gaylussacia. Typical impunctatus Nylander is
Palaearctic.
Colletes lacustris Swenk, 1906. Ent. News 17: 257. 2, 6.
Colletes vicinalis Graenicher, 1911. Pub. Mus. City Milwaukee, Bul. 1: 228, On

SPECIES GROUP WILLISTONI

brevicornis Robertson. United States east of Rocky Mts. Pollen: Unknown, but visits flowers
of Asclepias, Batodendron, Callirhoe involucrata, C. leiocarpa, Campanula, Crataegus,
Melilotus alba, Opuntia, Pastinaca sativa, Psoralea onobrychis, Rubus, Specularia.
Colletes brevicornis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 315. d.
Colletes opuntiae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 312. d, °.
Colletes brachyceros Graenicher, 1935. Ent. Soc. Amer., Ann. 28: 301. Preoce.

Taxonomy: Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 51. 2.

1756 Hymenoptera in America North of Mexico

willistoni Robertson. N.S. to Utah, south to Tex. and Fla. Pollen: Unknown, but visits flowers
of Ceanothus americanus, Melilotus alba, Monarda punctata occidentalis, Physalis
elliottii, P. virginiana, Psoralea tenuiflora, Pycnanthemum flexuosum, Rhus glabra,
Toxicodendron. Predator: Philanthus albopilosus Cress.
Colletes Willistoni Robertson, 1891. Amer. Ent. Soc., Trans. 18: 60. 2.

Taxonomy: Robertson, 1895. Amer. Ent. Soc., Trans. 22: 116. d.
Biology: Evans, 1975. Ent. Soc. Amer., Ann. 68: 891 (predator).
SPECIES GROUP CONSORS

californicus Provancher. South. and Centr. Calif. Pollen: Possibly oligolege of Phacelia,
visiting flowers of Baccharis, Cryptantha intermedia, Layia platyglossa, Nemophila
aurita, N. menziesii, Phacelia affinis, P. distans, P. tanacetifolia.
Colletes californica Provancher, 1895. Nat. Canad. 22: 189. 2, 6.
chamaesarachae Cockerell. Ariz., N. Mex. Pollen: Unknown, but visits flowers of
Chamaesaracha coronopus.
Colletes chamaesarachae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 49. 9.

consors consors Cresson. Mont., Idaho, Wyo., Colo., Calif. (Mono Basin). Pollen: Possibly
oligolege of Hydrophyllaceae, visiting flowers of Hydrophyllwm fendleri, Penstemon
virens, Phacelia leucophylla, Taraxacum.
Colletes consors Cresson, 1868. Boston Soe. Nat. Hist., Proc. 12: 168. ¢.
Colletes zonatus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 59. ¢.
Colletes myroni Cockerell, 1908. Entomologist 41: 293. 2.
consors mesocopus Swenk. N. S. to N. W. T. south to Alta., Mass., Mich., Wis. Pollen:
Unknown, but visits flowers of Geranium, Kalmia, Rubus.
Colletes mesocopus Swenk, 1907. Canad. Ent. 39: 364. 9, 6.

consors pascoensis Cockerell. B. C. and Idaho, south to Calif. and Utah. Pollen: Unknown, but
visits flowers of Nemophila aurita, Phacelia, Physalis, Sedum obtusatwm.
Colletes pascoensis Cockerell, 1898. Acad. Nat. Sci. Phila., Proce. 50: 51. 6, @.
linsleyi Timberlake. Calif. (Blythe). Pollen: Unknown, but visits flowers of the introduced
Tamariz.
Colletes linsleyi Timberlake, 1951. Wasmann Jour. Biol. 9: 205. 2.
nigrifrons Titus. North Que. to N. W. T. and Yukon, south into mts. of northeast Calif., Ariz.,
and N. Mex. Pollen: Unknown, but visits flowers of Arenaria nuttallii, Drymocallis
fissa, Eriogonum lobbii, E. latifolium nudum, E. mariifolium, Linum lewisii,
Mertensia sibirica, Oxytropis, Physalis, Polemonium, Potentilla anseriana, P. fruticosa,
P. glandulosa, P. hippiana, Ranunculus, Spiraea densiflora.
Colletes nigrifrons Titus, 1900. Canad. Ent. 32: 304. 2.
Colletes polemonii Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 425. 6.
Colletes florissantia Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 425. 6.

nitescens Timberlake. Calif. Pollen: Unknown, but visits flowers of Salix.
Colletes nitescens Timberlake, 1951. Wasmann Jour. Biol. 9: 187. 6, @.
paniscus mertensiae Timberlake. Calif. (Sierra Nevada and White mts.). Pollen: Unknown, but
visits flowers of Mertensia ciliata var. stomatechoides.
Colletes paniscus mertensiae Timberlake, 1951. Wasmann Jour. Biol. 9: 182. 6, °.
paniscus paniscus Viereck. Mont. and Wyo. south into mts. of Ariz., and N. Mex. Pollen:
Unknown, but visits flowers of Houstonia, Iris missouriensis, Mertensia franciscana,
Salix.
Colletes paniscus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 60. 6.
Colletes oromontis Viereck, 1903. Amer. Ent. Soc., Trans. 29: 61. @.
Colletes pleuralis Swenk, 1906. Ent. News 17: 259. 2, 6.
Colletes grisescens Cockerell, 1930. Amer. Mus. Novitates 397: 4. 6.
paniscus sculleni Timberlake. Wash. to Mont., south to Utah, Wyo., and Calif. Pollen:
Unknown, but visits flowers of Mertensia.
Colletes paniscus sculleni Timberlake, 1951. Wasmann Jour. Biol. 9: 183. 6, @.

Superfamily APOIDEA —= 1757

scopiventer Swenk. Tex., Ariz. and South. Calif. Pollen: Unknown, but visits flowers of
Baccharis emoryi, Brodiaea, Chamaesaracha conioides, Solanum rostratum.
Colletes scopiventer Swenk, 1908. Nebr. Univ., Dept. Ent. Contrib. 1: 46. 2 (d misdet.).
sphaeralceae Timberlake. Calif., Ariz., Nev., Utah. Pollen: Apparently oligolege of
Sphaeralacea, visiting flowers of S. alata, S. ambigua, S. orcutti, S. rosacea.
Colletes sphaeralceae Timberlake, 1951. Wasmann Jour. Biol. 9: 189. d, 2.
swenki Stephen. Tex., Kans. Pollen: Unknown, but visits flowers of Acacia greggii,
Chamaesaracha conioides, Coreopsis douglasti, Marrubium vulgare, Prosopis
glandulosa, Quincula lobata, Rhus microphylla.
Colletes swenki Stephen, 1954. Kans. Univ. Sci. Bul. 36: 450, fig. 68. 2, d.
texanus crawfordi Swenk. North. Tex. Pollen: Unknown, but visits flowers of Physalis.
Colletes crawfordi Swenk, 1906. Ent. News 17: 257. °.
texanus texanus Cresson. South. Tex.
Colletes texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 249. °.
wickhami Timberlake. Kans., Tex., N. Mex., Ariz., South. Calif. Pollen: Unknown, but visits
flowers of Chrysothamnus nauseosus, Helenium lacinatum, Phacelia popei, Quincula
lobata.
Colletes wickhami Timberlake, 1943. Amer. Mus. Nat. Hist., Bul. 81: 394. d.
xerophilus cismontanus Timberlake. Calif. (Hemet). Pollen: Unknown, but visits flowers of
Salix goodingii.
Colletes cismontanus Timberlake, 1951. Wasmann Jour. Biol. 9: 186. d, 2.
xerophilus sonoranus Timberlake. N. Mex. (Pecos). Pollen: Unknown, but visits flowers of
Salix.
Colletes xerophilus sonoranus Timberlake, 1951. Wasmann Jour. Biol. 9: 186. 2.
xerophilus xerophilus Timberlake. South. Calif. deserts. Pollen: Unknown, but visits flowers
of Salix.
Colletes xerophilus xerophilus Timberlake, 1951. Wasmann Jour. Biol. 9: 184. d, 9.

SPECIES GROUP INTERMIXTUS

bulbotibialis Stephen. N. Mex. (Belen).
Colletes bulbotibialis Stephen, 1954. Kans. Univ. Sci. Bul. 36: 458, fig. 71. 6.
intermixtus Swenk. Tex., N. Mex., Colo., Ariz., south. Calif. Pollen: Unknown, but visits
flowers of Cotoneaster integerrimum, Eriogonum fasciculatum, Gnaphalium beneolens,
Gutierrezia sarothrae, Haplopappus palmeri, Lippia, Physalis ixocarpa, Schinus molle,
Solanum douglasii.
Colletes intermixtus Swenk, 1905. Canad. Ent. 37: 302. °.
Colletes lippiarum Cockerell, 1909. Canad. Ent. 41: 394. °.

SPECIES GROUP CILIATUS

beamerorum Stephen. Tex.; Mexico (Chihuahua). Pollen: Unknown, but visits flowers of
Monarda punctata cory.
Colletes beamerorum Stephen, 1954. Kans. Univ. Sci. Bul. 36: 465, fig. 73. d.
ciliatoides Stephen. Utah, Idaho. Parasite: Epeolus pusillus Cress. Pollen: Unknown, but

visits flowers of Cuscuta.
Colletes ciliatoides Stephen, 1954. Kans. Univ. Sci. Bul. 36: 463, fig. 74. ¢.

Biology: Torchio, 1965. Kans. Ent. Soc., Jour. 38: 182-187.
ciliatus Patton. Va., Ill., lowa, Nebr., Kans., Colo. Pollen: Unknown, but visits flowers of
Cuscuta, Eryngium yuccifolium.
Colletes ciliata Patton, 1979. U. S. Geol. Geog. Survey Bul. 5: 369. °.
Colletes speciosa Robertson, 1891. Amer. Ent. Soc., Trans. 18: 62. °.
Colletes brevihirtus Timberlake, 1943. Amer. Mus. Nat. Hist., Bul. 81: 395. d.

1758 Hymenoptera in America North of Mexico

SPECIES GROUP DALEAE

algarobiae Cockerell. Tex., N. Mex., Utah, Ariz., south Calif.; north. Mexico; deserts. Pollen:
Apparently oligolege of Prosopis including P. glandulosa, P. pubescens, but also visits
flowers of Cercidium torreyanum, Dalea, Eriogonum, Melilotus, Rhus.
Colletes algarobiae Cockerell, 1900. Entomologist 33: 244. d, °.
clypeonitens Swenk. South Calif., Ariz., Nev., Utah, west. Tex. Parasite: Epeolus mesillae
(Ckll.). Pollen: Oligolege of Larrea tridentata, but also visits flowers of Cercidium
floridum, Dalea polyadenia.
Colletes clypeonitens Swenk, 1906. Canad. Ent. 38: 39. °.
covilleae Timberlake. South. Calif., Ariz.; Mexico (Baja Calif.). Pollen: Apparently oligolege of
Larrea tridentata.
Colletes covilleae Timberlake, 1951. Wasmann Jour. Biol. 9: 231. 6, °.
daleae Cockerell. Utah, Tex. to Calif.; Mexico (Baja Calif.). Pollen: Unknown, but visits flowers
of Dalea scoparia, Eriogonum fasciculatum, E. gracilentum, Prosopis, Rhus lawrina.
Colletes daleae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 43. 6.
deserticola Timberlake. N. Mex., Ariz., Calif.; Mexico (Baja Calif.). Parasite: Epeolus pusillus
Cress. Pollen: Apparently oligolege of Prosopis, including P. glandulosa, but also visits
flowers of Melilotus indica.
Colletes deserticola Timberlake, 1951. Wasmann Jour. Biol. 9: 232. 6, °.
petalostemonis Swenk. N. Dak. and Nebr. to Alta., Ariz., and Calif. Pollen: Unknown, but
visits flowers of Dalea polyadenia, Melilotus alba, Petalostemon flavescens.
Colletes petalostemonis Swenk, 1906. Canad. Ent. 38: 40. 2, ¢.
prosopidis Cockerell. Tex., N. Mex., south. Calif.; north. Mexico; deserts. Pollen: Apparently
oligolege of Prosopis, including P. glandulosa var. torreyanum, but also visits flowers of
Melilotus indica.
Colletes prosopidis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 46. 6.
salicicola Cockerell. Tex. to south. Calif., deserts. Pollen: Polylege of a wide variety of plants,
including Acacia, Cercidium, Dalea, Eriogonum, Hyptis, Larrea, Prosopis, but also
visits flowers of Asclepias, Phacelia and Salix for nectar.
Colletes salicicola Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 50. 9.
solitarius Timberlake. Calif. (Inyo Co.).
Colletes solitarius Timberlake, 1951. Wasmann Jour. Biol. 9: 216. °.
stepheni Timberlake. Calif. (Riverside Co.). Pollen: Oligolege of Larrea tridentata, but also
visits flowers of Cercidiwm floridum and Geraea canescens for nectar.
Colletes stepheni Timberlake, 1958. Pan-Pacifie Ent. 34: 148. d, 9.

Biology: Hurd and Powell, 1958. Pan-Pacifie Ent. 34: 147-153, 1 fig. (nesting habits, male
behavior).

SPECIES GROUP ARIDUS

aridus Stephen. Tex., N. Mex. Pollen: Unknown, but visits flowers of Phacelia popei.
Colletes aridus Stephen, 1954. Kans. Univ. Sci. Bul. 36: 494, fig. 75. 5, °.

SPECIES GROUP TITUSENSIS

titusensis Mitchell. Fla. (Titusville, Tampa).
Colletes titusensis Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 237, figs. 16, 17. 3.

SPECIES GROUP LONGIFACIES

longifacies Stephen. Fla. Pollen: Unknown, but visits flowers of Liatris laevigata.
Colletes longifacies Stephen, 1954. Kans. Univ. Sci. Bul. 36: 500. 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 43.
SPECIES GROUP UNASSIGNED

platyenema Snelling. Ariz. (Huachuca Mts.). Pollen: Unknown, but visits flowers of Acacia
angustissima.
Colletes platyenema Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267: 2, figs. 5-8. ¢,
On

Superfamily APOIDEA 1759

SUBFAMILY DIPHAGLOSSINAE

This is a New World group of large to very large, robust bees which occur in the tropics, sub-
tropics as well as the warm temperate regions of both North and South America.

Taxonomy: Michener, 1966. Kans. Univ. Sci. Bul. 46: 717-751, 48 figs.
TRIBE CAUPOLICANINI

Genus PTILOGLOSSA Smith

Ptiloglossa Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 7.
Type-species: Ptiloglossa ducalis Smith. Monotypic.
Ptiloglossa subg. Ptiloglossodes Moure, 1945. Arq. do Mus. Paranaense 4: 153.
Type-species: Megacilissa tarsata Friese. Orig. desig.
arizonensis Timberlake. Ariz. Pollen: Polylege of Cassia bauwhinioides, Larrea tridentata,
Solanum elaeagnifolium, S. rostratum; females also visit flowers of Cucurbita
foetidissima for nectar.
Ptiloglossa arizonensis Timberlake, 1946. Pan-Pacific Ent. 22: 157. 9.

Biology: Linsley, 1962. Pan-Pacifiec Ent. 38: 75-82. —Linsley and Cazier, 1970. Kans. Ent.
Soc., Jour. 43: 251-261, 3 tables. —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 19
(floral relationships).

jonesi Timberlake. Ariz. Pollen: Polylege of Datura meteloides, Larrea tridentata, Solanwm

elaeagnifolium, and S. rostratum.
Ptiloglossa jonesi Timberlake, 1946. Pan-Pacifie Ent. 22: 158. 2.

Biology: Linsley and Cazier, 1963. Pan-Pacific Ent. 39: 1-18. —Linsley and Cazier, 1970.
Kans. Ent. Soc., Jour. 43: 251-261, 3 tables. —Hurd and Linsley, 1975. Smithsn. Contrib.
Zool. 193: 20 (floral relationships).

Genus CAUPOLICANA Spinola

Although a few species of this genus occur in tropical America, the largest numbers of species
chiefly inhabit the warm temperate and subtropical regions of both continents. Most of the spe-
cies have been described from Chile and only two of the five recognized subgenera are known to
occur in southwestern United States.

Genus CAUPOLICANA Subgenus CAUPOLICANA Spinola

Caupolicana Spinola, 1851. In Gay, Hist. Fis. Pol. Chile, Zool., v. 6, p. 212.
Type-species: Caupolicana gayi Spinola. Desig. by Sandhouse, 1943.
Megacilissa Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 123.
Type-species: Caupolicana fulvicollis Spinola. Monotypic. (=Megacilissa superba
Smith).
Megalocilissa Schulz, 1906. Spolia Hym., p. 243. Emend.

Taxonomy: Michener, 1966. Kans. Univ. Sci. Bul. 46: 732-736.
electa (Cresson). N. C., to Ga., northwest Fla., Ala. Pollen: Unknown, but visits flowers of

Gerardia and Trichostema dichotemum.
Megacilissa electa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 221. 6.

Taxonomy: Michener, 1966. Kans. Univ. Sci. Bul. 46: 736-738.

Biology: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 24 (seasonal activity and flight
period).
ocellata Michener. Kans., N. Mex., Ariz., Tex.; Mexico (Chihuahua). Pollen: Apparently an
oligolege of Dalea including D. lanata, D. scoparia, but visits flowers of Gawra coccinea
and Petalostemon flavescens for nectar.
Caupolicana (Caupolicana) ocellata Michener, 1966. Kans. Univ. Sci. Bul. 46: 738, figs. 12,
23-27, 29. 3, 2.

Biology: Michener, 1966. Kans. Univ. Sci. Bul. 46: 739-740 (nest, floral relationships).

1760 Hymenoptera in America North of Mexico

yarrowi (Cresson). Ariz., Utah (Zion Park), N. Mex., Tex.; Mexico. Pollen: Polylege of Cassia
bauhinioides, Datura meteloides, Hofmanseggia jamesu, Larrea tridentata, Mentzelia
pumila, Solanum elaeagnifolium, S. rostratum, but visits other flowers including
Aloysia wrightii, Eysenhardtia polystacha, and Melilotus alba for nectar.
Megacilissa yarrowi Cresson, 1875. Rpt. Geog. Geol. Explor. and Survey west of 100th
Meridian, v. 5, p. 723. °.

Taxonomy: Michener, 1966. Kans. Univ. Sci. Bul. 46: 733-736, figs. 12, 13-17, 29. —Snelling,
1975. Los Angeles Co. Mus., Contrib. Sci. 267: 2 (geogr. records).

Biology: Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 403. —Linsley and Hurd,
1959. Ent. News 70: 67. —Linsley, 1960. N. Y. Ent. Soc., Jour. 68: 18. —Linsley, 1962. Ent.
Soc. Amer., Ann. 55: 158-159. —Linsley and Cazier, 1963. Pan-Pacific Ent. 39: 1-18.
—Michener, 1966. Kans. Univ. Sci. Bul. 46: 734-736. —Linsley and Cazier, 1970. Kans. Ent.
Soc., Jour. 43: 251-261, 3 tables. —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193:
18-19 (summary of habits and floral relationships).

Genus CAUPOLICANA Subgenus ZIKANAPIS Moure

Zikanapis Moure, 1945. Arq. do Mus. Paranaense, 4: 147.

Type-species: Ptiloglossa zikani Friese. Orig. desig.
Foersterapis Moure, 1964. Studia Ent. 7: 441.

Type-species: Zikanapis foersteri Moure and Seabra. Orig. desig.

Taxonomy: Michener, 1966. Kans. Univ. Sci. Bul. 46: 727-728, figs. 12, 30-34, 47.
elegans Timberlake. Ariz. (Portal); Mexico (Tehuacan and Puebla). Pollen: Unknown, but visits
flowers of Salvia.
Caupolicana elegans Timberlake, 1965. N. Y. Ent. Soc., Jour. 73: 46. 6.

Taxonomy: Michener, 1966. Kans. Univ. Sci. Bul. 46: 743-745, fig. 47.

NOMEN NUDUM IN CAUPOLICANA SPINOLA

Megacilissa monticola Ashmead, 1889. In Cockerell, Colo. Biol. Assn. 10th Rpt., [p. 2].
Probably published originally in Custer County Courant newspaper.

SUBFAMILY HYLAEINAE

The females of this subfamily, like those of the Australian subfamily Euryglossinae and most
of those in the chiefly Neotropical subfamily Xeromelissinae, lack scopal hairs and transport pol-
len in the crop. The subfamily Hylaeinae is represented in North America only by the genus
Hylaeus. The species of this genus make cells of a transparent cellophane-like material in preex-
isting burrows, hollow stems, holes or cavities.

Revision: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 60-85 (spp. of eastern U. S.).

Taxonomy: Lovell, 1910. Psyche 17: 177 (Maine). —Metz, 1911. Amer. Ent. Soc., Trans. 37: 85
(spp.). —Meade-Waldo, 1923. Gen. Ins., v. 181, p. 29 (cat.). —Cockerell and Sumner, 1931.
Amer. Mus. Novitates 490: 14 (Rocky Mt. females). —Michener, 1942. N. Y. Ent. Soc.,
Jour. 50: 273 (subgenera). —Snelling, 1966. South. Calif. Acad. Sci., Bul. 65: 164-175
(Nearctic subgenera). —Warncke, 1970. Rech. Agron. Gembloux, Bul. 5 (n.s.): 745-747
(status of generic names Hylaeus and Prosopis).

Genus HYLAEUS Fabricius

Genus HYLAEUS Subgenus PROSOPIS Fabricius

Prosopis Fabricius, 1804. Syst. Piezatorum, p. 293.
Type-species: Mellinus bipwnctatus Fabricius. Desig. by Morice and Durant, 1914.
(=Sphex signata Panzer).
Prosapis Ashmead, 1894. Psyche 7: 43. Emend.

Revision: Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 1-18 (spp. of western U. S.).

aenigmus (Viereck). N. Mex. Possibly a synonym of H. episcopalis (Cockerell).
Prosopis aenigmus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 64. 3.

Superfamily APOIDEA 1761

affinis (Smith). New England and adjacent Canada west to B. C., Wash., Idaho, south to Utah,

Kans., Miss., and Ga. Pollen: Unknown, but visits a wide variety of flowers including
Amorpha, Apocynum, Aruncus, Ascelpias, Blephilia, Boltonia, Campanula, Ceanothus,
Cephalanthus, Cicuta, Clematis, Cornus, Crataegus, Cryptotaenia, Daucus carotus,
Erigeron, Eryngium, Eulophus, Eupatorium, Euphorbia, Fragaria, Geranium, Geum,
Gnaphalium, Heracleum, Houstonia, Hydrangea, Hypericum, Krigia, Lactuca, Lepidium,
Lycopus, Malva, Melilotus, Nelumbo, Osmorrhiza, Oxypolis, Pastinaca, Petalostemon,
Philadelphicus, Polygonum, Polytaenia, Potentilla, Ptelea, Pycnanthemum,
Pyracantha, Rhus, Rubus, Salix, Salvia, Sanicula, Sium, Solidago, Stellaria,
Stenanthium Symphoricarpos, Taenidia, Thaspium, Valerianella, Veronica, Viburnum,
Zizia. Predator: Philanthus crabroniformis Sm., P. politus Say.

Prosopis affinis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 24. 2 (d misdet.).

Prosopis ziziae Robertson, 1896. Canad. Ent. 28: 136. 2, d.

Prosopis ziziae dunningi Cockerell, 1898. Entomologist 31: 188. d.

Hylaeus albertensis Cockerell, 1937. Canad. Ent. 69: 126. 6, °.

Taxonomy: Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 3.
confluens (Smith). N. J. to Fla. Pollen: Unknown, but visits flowers of Chrysanthemum,
Chrysopsis, Erigeron, Eryngium, Hypericum, Ilex, Itea, Melilotus, Oxydendrum,
Oxypolis filiformis, Polygonum, Rhus, Solidago.
Prosopis confluens Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 24. 2.
Prosapis triangularis Cockerell, 1896. Psyche 7 (sup.): 31. d.
episcopalis coquilletti (Cockerell). Oreg. and Idaho south to Mexico (Baja California and
Sonora) east to Texas. Pollen: Unknown, but visits flowers of Acamptopappus, Agave
deserti, Asclepias subulata, Aster abatus, Baileya pleniradiata, Chamaebatiaria
millefolium Croton californicus, Dalea californica, D. fremontii, D. saundersii, D.
schottii, Datura meteloides Eriodictyon californicum, Eriogonum fasciculatum, E.
inflatum, Gilia davyi, Haplopappus cooperi, Hyptis emoryi, Isomeris arborea, Larrea
tridentata, Lepidium fremontii, Lupinus formosus, Melilotus alba, Peucephyllum
schottii, Prosopis juliflora glandulosa, Sphaeralcea orcuttii, Stanleya pinnata, Tamarix
gallica.
Prosapis coquilletti Cockerell, 1896. Psyche 7 (sup.): 439. 3.

Taxonomy: Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 10 (as H. rugulosus
episcopalis). —Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267: 8 (tax. status).
episcopalis episcopalis (Cockerell). Alta., south to Calif. and N. Mex. Pollen: Unknown, but

visits a wide variety of flowers including Achillea lanulosa, A. millaefolium,
Adenostoma fasciculatum, Angelica tomentosa, Anthemis cotulla, Apocynum
adrosaemifolium, Asclepias fascicularis, A. mexicana, Baccharis douglasii, Calochortus
venustus, Ceanothus integerrimus, Chaetopappa aurea, Chamaebatia foliolosa, Clarkia
bilobata, Erigeron ramosus, Eriodictyon californicum, Eriogonum fasciculatum, E.
inflatum, E. latifolium nudum, E. virgatum, Eriophyllum confertiflorum, Eryngium
aristatulum, Geranium richardsonii, Gnaphalium californicum, Gutierrezia californica,
G. sarothrae, Haplopappus cooperi, Heteromeles arbutifolia, Linanthus montanus,
Lupinus, Oxypolis occidentalis Penstemon breviflorus, Perideridia gairdneri, Phacelia,
Prunus ilicifolia, Rhamnus californica, Scrophularia californica, Solidago, Stanleya
pinnata, Swertia parryi, Trichostema parishii.

Prosapis episcopalis Cockerell, 1896. Psyche 7 (sup.): 29. d.

Prosopis universitatis Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 225. 6.

Taxonomy: Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 9, 10 (as. H. rugulosus
rugulosus and H. r. episocpalis). —Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267:
8 (synonymy, tax. status).
episcopalis giffardiellus Cockerell. Calif. (Sacramento and San Joaquin Valleys). Pollen:
Unknown, but visits flowers of Melilotus alba, Raphanus sativus, Rosa californica,
Rubus, Salix, Trifolium repens.
Hylaeus giffardiellus Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 186. d.

Taxonomy: Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 11 (as H. rugulosus
giffardiellus). —Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267: 8 (tax. status).

1762 Hymenoptera in America North of Mexico

episcopalis metzi Snelling. South. Calif. Pollen: Unknown, but visits a wide variety of flowers

including Adenostoma fasciculatum, Amorpha fruticosa, Achillea millaefolium,
Apocynum androsaemifolium, Chrysopsis villosa, Cordylanthus nevinii, Encelia,
Eriogonum davidsonii, E. elongatum, E. fasciculatum, E. fasciculatum var. polifolium,
E. gracile, E. nudum, E. subscaposum, E. wrightii, Eriodictyon, Geranium richardsonii,
Gilia gilioides, Haplopappus cooperi, Lupinus cytisoides, Melilotus alba, Mentzelia
laevicaulis, Monardella linoides, Penstemon grinnellii, P. labrosus, P. palmeri,
Phacelia ramosissima, Potentilla glandulosa, Rhamnus californica, R. crocea, Salvia
carnosa, Solidago californica, S. confinis, Sphenosciadium capitellatum.

Hylaeus (Prosopis) rugulosus metzi Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98:
ora

Taxonomy: Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267: 8 (tax. status).

flammipes (Robertson). Fla. (Citrus Co.).
Prosapis flammipes Robertson, 1893. Amer. Ent. Soc., Trans. 20: 2738. 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 64.

gaigei (Cockerell). Mich. Possibly a synonym of H. modestus Say.
Prosopis gaigei Cockerell, 1916. Mich. Univ. Mus. Zool., Oceas. Papers 23: 2. 2.

Taxonomy: Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci. 180: 27.
illinoisensis (Robertson). Ill. and W. Va. to N. Y., Mass., Maine. Pollen: Unknown, but visits

flowers of Apocynum cannabinum, Aruncus, Aster, Cicuta, Cornus, Crataegus,
Eulophus, Gonolobus, Heracleum, Lycopus, Osmorrhiza, Pastinaca, Rhus, Salix,
Solidago, Taenidia, Thaspium, Viburnum.

Prosopis Illinoisensis Robertson, 1896. Canad. Ent. 28: 138. ¢.

Hylaeus (Prosopis) certus Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 241, figs. 4-7.
36, 2. Preoce.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 64.
insolitus Snelling. Ariz., Tex.; Mexico (Chihuahua). Pollen: Unknown, but visits flowers of

Cleome serrulata.
Hylaeus (Prosopis) insolitus Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 16. 6,
OR

modestus citrinifrons (Cockerell). B. C., Alta., Mont., Wyo., Colo., N. Mex., Calif., Nev. Pollen:

Unknown, but visits a wide variety of flowers including Adenostoma fasciculatum,
Angelica arguta, A. tomentosa, Aralia californica, Baccharis douglasii, Chaenactis
glabriuscula lanosa, Eriodictyon californicum, Erigonum nudum, Eriophyllum
staechadifolium, Geranium dissectum, G. richardsoni, Gnaphalium californicum,
Holodiscus discolor, Melilotus alba, Oxybaphis occidentalis, Perideridia gairdneri,
Phacelia humilis, Potentilla glandulosa, Ranwnculus, Rhamnus californica, Rubus,
Salvia sonomensis, Scrophularia californica, Solidago, Sphenosciadium capitellatum,
Trichostema parishii.

Prosapis citrinifrons Cockerell, 1896. Psyche 7 (sup.): 27. ¢.

Prosapis rugosulus Cockerell, 1896. Psyche 7 (sup.): 28. 3.

Prosapis rugosulus var. fallax Cockerell, 1896. Psyche 7 (sup.): 28. ¢.

Taxonomy: Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 98: 5. —Snelling, 1975. Los
Angeles Co. Mus., Contrib. Sci. 267: 7-8 (synonymy, tax. status).
modestus modestus Say. East. Canada and Atlantic States to Minn., La. Parasite:

Gasteruption kirbii Westw. Pollen: Unknown, but visits a wide variety of flowers
including Acer, Achillea, Amorpha, Angelica, Apocynum, Arabis, Aralia, Aruncus,
Azalea, Cacalia, Castanea pumila, Ceanothus, Chrysanthemum, Cicuta maculata,
Cornus, Crataegus, Daucus carota, Erigeron, Eulophus, Eupatorium, Gerardia, Geum,
Heracleum, Houstonia purpurea, Hydrangea, Ilex, Koellia, Lepidium, Malva, Monarda,
Osmorrhiza, Pastinaca, Ptelea, Pycnanthemum, Pyracantha, Rhus, Rubus, Sanicula,
Sium, Solidago, Spiraea, Symphoricarpos, Taenidia, Thaspium, Zizia. Predator:
Philanthus bilunatus Cr., P. politus Say.

Hylaeus modestus Say, 1837. Boston Jour. Nat. Hist. 4: 392. 2 (¢ misdet.).

Prosapis pennsylvanica Cockerell, 1896. Psyche 7: 439. ¢.

Superfamily APOIDEA 1763

Prosopis nucleolus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 64. °.
Prosopis sayi Robertson, 1904. Canad. Ent. 36: 274, 275. 2, d.
Prosopis minyra Lovell, 1909. Ent. News 20: 413. ¢.

Prosopis binghami Lovell, 1910. Psyche 17: 180. ¢.

Prosopis supracurta Swenk and Cockerell, 1910. Ent. News 21: 71. ¢.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 71. —Rau, 1930. Psyche 37: 173-175.
—Krombein, 1967. Trap-nesting wasps and bees, p. 262. —Medler, 1966. Ent. Soc. Wash.,
Proce. 68: 181.

nelumbonis (Robertson). II]. and Ohio to Fla., Ala., La. Pollen: Unknown, but visits flowers of
Castalia tuberosa, Nelumbo lutea.
Prosopis nelumbonis Robertson, 1890. Amer. Ent. Soc., Trans. 17: 316. 2.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 67.
ornatus Mitchell. N. C. to Fla. Pollen: Unknown, but visits flowers of Cephalanthus,
Hypericum, Melilotus, Parthenocissus.
Hylaeus (Prosopis) ornatus Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 242, figs.
SHill, Gi, Ge

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 67.
schwarzii (Cockerell). Mass. to Fla. Pollen: Unknown, but visits flowers of Hydrocotyle
umbellata, Medicago lupulina, Melilotus alba, Sagittaria, Salix.
Prosapis Schwarzii Cockerell, 1896. Ent. Monthly Mag. 32: 218. 9.
Prosopis fossata Metz, 1911. Amer. Ent. Soc., Trans. 37: 135. ¢ (2 misdet.).

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 68.
transvittatus (Cockerell). South. Ariz. (mts.); Mexico (Durango, D. F. and Veracruz).
Prosopis transvittata Cockerell, 1917. Ann. and Mag. Nat. Hist. (8) 20: 437. 2, d.

volusiensis Mitchell. Fla. (Dade and Volusia Cos.).
Hylaeus (Prosopis) volusiensis Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 245. 2, ¢.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 69.
Genus HYLAEUS Subgenus PARAPROSOPIS Popov

Prosopis subg. Paraprosopis Popov, 1939. Acad. des Sci. U. R. S. S., Compt. Rend. (Dok.)
(n. s.) 25: 169.
Type-species: Prosopis pictipes (Nylander). Monotypic and orig. desig.

Revision: Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci. 180: 27-58, figs. 7-12 (Nearctic
species).
SPECIES GROUP ASININUS

asininus (Cockerell and Casad). West. Tex. to South. Calif.; Mexico (Baja California). Parasite:
Anthrax irroratus irroratus Say. Pollen: Unknown, but visits flowers of Baccharis
emoryi, Chrysothamnus paniculatus, Gutierrezia lucida, Haplopappus acradenius,
Eriogonum fasciculatum, Lepidospartum, Prosopis, Solidago californica.
Prosopis asininus Cockerell and Casad, 1895. Amer. Ent. Soc., Trans. 22: 299. ¢.
Prosopis bipes Cockerell and Casad, 1895. Amer. Ent. Soc., Trans. 22: 300. °.
Prosapis asinina var. bigeloviae Cockerell, 1898. N. Mex. Univ., Bul. 1: 72. 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 261 (nest, parasite).
SPECIES GROUP CALVUS

calvus (Metz). South. Oreg. to Mexico (Baja California), Nev., Ariz. Pollen: Unknown, but visits
flowers of Baccharis, Ceanothus cordulatus, C. cuneatus, C. integerrimus, Collinsia
heterophylla, Eriodictyon californicum, Eriogonum, Fremontia californica, Holodiscus
discolor, Lepidium virginicum, Penstemon antirrhinoides, Photinia heteromeles,
Prunus ilicifolia, Rhamnus californicus, R. crocea, Salix.
Prosopis calvus Metz, 1911. Amer. Ent. Soc., Trans. 37: 143. 6.

1764 Hymenoptera in America North of Mexico

georgicus (Cockerell). N. Y., N. C., Ga., Fla., Tex. Pollen: Unknown, but visits flowers of
Crataegus, Hydrangea, Pyracantha, Salix.
Prosapis georgica Cockerell, 1898. Psyche 7 (sup.): 488. ¢.
Prosopis georgica var. leana Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 27. 6.
Hylaeus (Metziella) hydrangeae Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 244. 2.

timberlakei Snelling. Calif. (Sierra Nevada Mts.). Pollen: Unknown, but visits flowers of
Ceanothus integerrimus, Eriodictyon californicum, Eriogonum nudum, Haplopappus
arborescens, Lessingia leptoclada, Perideridia gairdner, Solidago californica.
Hylaeus (Paraprosopis) timberlake Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci.
180: 50. 3d, @.

SPECIES GROUP MEGALOTIS

megalotis (Swenk and Cockerell). Mont., Colo., Nebr., Utah, Ariz. Pollen: Unknown, but visits
flowers of Cleome serrulata, Chrysothamnus, Eriogonum aureus.
Prosopis megalotis Swenk and Cockerell, 1910. Ent. News 21: 69. 6, °.

SPECIES GROUP SONORENSIS

sonorensis Cockerell. Ariz., south. Calif.; Mexico (Baja California and Sonora). Pollen:
Unknown, but visits flowers of Encelia, Eriogonum inflatum, Eucnide urens, Prosopis
glandulosa, Scrophularia californica, Stanleya pinnata.
Hylaeus sonorensis Cockerell, 1924. Calif. Acad. Sci., Proe. (4) 12: 530. 9.
Hylaeus sonorensis var. melanorhinus Cockerell, 1924. Calif. Acad. Sci., Proe. (4) 12: 531.
OF

SPECIES GROUP WOOTONI

coloradensis (Cockerell). B. C., Wash., Idaho, Colo., N. Mex., Ariz., Utah, Nev., east. and south.
Calif. Pollen: Unknown, but visits flowers of Achillea lanulosa, A. millaefolium,
Adenostoma fasciculatum, Baccharis pilularis, Barbarea orthoceras, Ceanothus
cordulatus, Chrysothamnus nauseosus, C. nauseosus consimilis, Descurainia sophia,
Erigeron canadensis, E. pygmaeus, Eriodictyon californicum, Eriogonum elegans, E.
lobbii, E. mariifolium, E. latifolium nudum, E. subscaposum, E. umbellatum, E.
virgatum, Gnaphalium thermale, Haplopappus arborescens, Holodiscus boursieri,
Ivesia muirii, Lasthenia chrysostoma, Lepidospartum squamatum, Mentzelia
laevicaulis, Oxypolis occidentalis, Penstemon davidsonii, P. newberryi, Potentilla
glandulosa, Rhamnus californica, Rhus, Solidago californica, S. multiradiata,
Sphenosciadium capitellatum.
Prosapis coloradensis Cockerell, 1896. Psyche 7 (sup.): 30. 6.
cookii (Metz). N. Mex., Nev. (Churchill Co.), Ariz., south. Calif. Pollen: Unknown, but visits
flowers of Asclepias tuberosa, Chrysothamnus stenophyllus, Erigeron canadensis,
Eriogonum aureum, E. fasciculatum, E. inflatum, E. plumatella, Fallugia, Gutierrezia
lucida, Haplopappus gracilis, H. linearifolius, Lepidospartum, Nolina parryi,
Petalostemon candidum, Phacelia ramosissima, Prosopis, Rhus trilobata, Salvia dorii,
S. vitosa, Solidago californica, Sphaeralcea ambigua, Tetradymia canescens.
Prosopis cookii Metz, 1911. Amer. Ent. Soc., Trans. 37: 139. ¢ (2 misdet.).
floridanus (Robertson). Maine, Minn., Mich., Ill., Ky., N. C., Ga., Fla. Pollen: Unknown, but
visits flowers of Aster, Cornus paniculata, Erigeron quercifolius, Eulophus
americanus, Ilex, Polygonella polygama, Pyracantha, Solidago.
Prosopis floridanus Robertson, 1893. Amer. Ent. Soc., Trans. 20: 273. °.
Prosopis eulophi Robertson, 1905. Canad. Ent. 37: 236. 2, é.
Hylaeus (Paraprosopis) packardi Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 244. ¢.
lunicraterius Snelling. Idaho (Craters of the Moon National Monument), Calif. Pollen:
Unknown, but visits flowers of Chamaebataria millefolium, Chrysothamnus nauseosis,
C. viscidiflorus, Eriogonum nudum, E. ovalifolium, Phacelia leucophylla, Populus
tremuloides, Senecio serra.
Hylaeus (Paraprosopis) lunicraterius Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci.
180: 46. 3, 2.

Superfamily APOIDEA 1765

nevadensis (Cockerell). B. C., Wash., Idaho, Oreg., Calif., Nev. Pollen: Unknown, but visits

flowers of Achillea lanulosa, Adenostoma fasciculatum, Angelica tomentosa,
Apocynum androsaemifolium, Baccharis douglasii, Ceanothus cuneatus, C.
integerrimus, C. soredatus, Centaurea melitensis, Cicuta, Collinsia, Cordylanthus
rigidus, Corethrogyne filaginifolia, Eriodictyon californicum, E. crassifolium, E.
trichocalyx, Erigeron heterophyllus, Eriogonum latifolium nudum, E. saxatile, E.
wrightii, Eriophyllum confertiflorum, Geranium richardsonii, Gnaphalium
californicum, Haplopappus arborescens, H. parishii, Holodiscus discolor, Horkelia
bernardina, Monardella linoides, M. odoratissima, Penstemon labrosus, Perideridia
gairdneri, Phacelia humilis, Potentilla glandulosa, Prunus ilicifolia, Rhamnus
californica, R. crocea, Solidago californica, S. occidentalis, Tetradymia canescens.

Prosapis nevadensis Cockerell, 1896. Psyche 7 (sup.): 32. 3.

Prosopis hesperiphila Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 29. 6.

Hylaeus oregonensis Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 160. 2, d.

personatellus (Cockerell). Colo. (Corona and Ouray), Calif. (Sierra Nevada Mts.). Pollen:

Unknown, but visits flowers of Aster, Brodiaea gracilis, Chrysothamnus nauseosus,
Erigeron pygmaeus, Eriogonum elatum, E. lobbii, E. mariifolium, E. nudum, E.
ovalifolium, Ivesia muiri, Monardella odoratissima, Penstemon davidsonii, P.
newberryi, Perideridia gairdneri, Potentilla glandulosa, Solidago multixadiata.

Prosopis personatellus Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 16: 485. °.

polifolii (Cockerell). Calif. Parasite: Eurytoma stigmi Ashm. Pollen: Unknown, but visits

flowers of Achillea millaefolium, Adenostoma fasciculatum, Alyssum maritimum,
Anaphalis margaritacea, Baccharis douglasii, B. emoryi, B. glutinosa, B. pilularis, B.
viminea, Ceanothus crassifolius, C. greggii, C. integerrimus, C. leucodermis, C.
soredatus, Chrysothanmus nauseosus, Coreopsis lanceolata, Cryptantha intermedia,
Dendromecon rigida, Erigeron canadensis, E. stenophyllus, Eriodictyon trichocalyx
lanatum, Eriogonum elongatum, E. fasciculatum, E. gracile, E. latifolium, E.
latifolium nudum, E. vimineum, Eriophyllum confertiflorum, Eryngium aristatulum,
Erysimum perenne, Euphorbia albomarginata, Gnaphalium beneolens, G. californicum,
Gutierrezia californica, G. sarothrae, Haplopappus arborescens, H. linearifolius, H.
parishii, Helenium puberulum, Heteromeles arbutifolia, Heterotheca grandiflora,
Lasthenia, Malacothrix tenuifolia, Melilotus alba, Mentha, Opuntia, Penstemon
antirrhinoides, Perideridia, Phacelia distans, Prunus demissa, P. ilicifolia,
Pyracantha, Rhamnus californica, R. crocea, Rhus diversiloba, R. laurina, R. trilobata,
Salix lasiolepis, Salvia, Sambucus caerulea, Scrophularia californica, Sedum, Senecio
californica, Solidago californica, S. occidentalis, Sphenosciadium capitellatum, Swertia
parryi, Tetradymia canescens, Trichostema parishii.

Prosopis polifolii Cockerell, 1901. Canad. Ent. 33: 281. d.

Hylaeus polifolii catalinensis Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 150. d, @.

wootoni (Cockerell). B. C. to Calif., Ariz., N. Mex.; Mexico (Chihuahua). Pollen: Unknown, but

visits flowers of Achillea lanulosa, Apocynum androsaemifolium, Aster canescens,
Calochortus paludicola, C. splendens, Ceanothus cordulatus, C. palmeri,
Chrysothamnus nauseosus, C. pumilius, C. viridulus, C. viscidiflorus puberulus,
Eriodictyon trichocalyx, Eriogonum davidsoni, E. fasciculatum, E. nudum, E.
subscaposum, E. umbellatum, E. wrightii, E. wrightii membranaceum, Eriophyllum,
Erysimum asperum, Haplopappus arborescens, H. uniflorus gossypinus, H. parishii,
Horkelia fusca, Lepidium virginicum, Mentzelia albicaulis, Phacelia imbricata,
Potentilla glandulosa, Ranunculus californicus, Rhamnus californicus, Solidago
californica, S. confinis, Tetradymia canescens.

Prosapis wootoni Cockerell, 1896. Psyche 7 (sup.): 26. d.

Prosapis divergens Cockerell, 1896. Psyche 7 (sup.): 29. 6.

Prosopis clandestinus Viereck, 1903. Amer. Ent. Soc., Trans. 17: 65. ¢.

Prosopis excavata Swenk and Cockerell, 1910. Ent. News 21: 70. °.

Hylaeus perparvus Cockerell and Sumner, 1931. Amer. Mus. Novitates 490: 10. d.

SPECIES GROUP UNASSIGNED

seclusus Cockerell and Sumner. Colo.
Hylaeus seclusus Cockerell and Sumner, 1931. Amer. Mus. Novitates 490: 12. d.

1766 | Hymenoptera in America North of Mexico

Genus HYLAEUS Subgenus PROSOPELLA Snelling

Hylaeus subg. Prosopella Snelling, 1966. Biol. Soc. Wash., Proc. 79: 189.
Type-species: Hylaeus hurdi Snelling. Monotypic and orig. desig.

Revision: Snelling, 1966. Biol. Soc. Wash., Proc. 79: 139-144.
hurdi Snelling. Ariz.; Mexico (Chihuahua). Pollen: Unknown, but visits flowers of Baccharis,

Ceanothus including C. greggii, Melilotus alba, Penstemon campanulatus, Verbesina.
Hylaeus (Prosopella) hurdi Snelling, 1966. Biol. Soc. Wash., Proc. 79: 140, fig. 1. d, 9.

Genus HYLAEUS Subgenus HYLAEANA Michener

Hylaeus subg. Hylaeana Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 28.
Type-species: Hylaeus panamensis Michener. Monotypic and orig. desig.
panamensis Michener. Tex. west to South. Calif., south to Panama. Pollen: Unknown, but
visits flowers of Dicraurus, Hyptis emoryi, Verbesina encelioides.
Hylaeus (Hylaeana) panamensis Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 28. ¢.

Taxonomy: Snelling, 1968. Los Angeles Co. Mus., Contrib. Sci. 144: 4 (as H. aztecus).
—Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267: 8 (tax. status).

Genus HYLAEUS Subgenus HYLAEUS Fabricius

Hylaeus Fabricius, 1793. Ent. system., v. 2, p. 302.
Type-species: Apis annulata Linnaeus. Desig. by Latreille, 1810 (~Prosopis
annulata Fabricius).

Revision: Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci. 180: 1-27, figs. 1-6.
SPECIES GROUP BISINUATUS

bisinuatus Forster. Holarctic; Transcont. south. Canada and north. U.S., south to Ga. and

Calif. (Central Valley); introduced into Hawaii; Europe. Possibly introduced from
Europe into the area around Fargo N. D. prior to 1912. Pollen: Exhibits a pronounced
tendency toward oligolecty on flowers of the family Leguminosae, especially the
introduced species of Melilotus, but also visits the flowers of other, mostly introduced,
plants including those of the Amaranthaceae, Asclepiadaceae, Compositae,
Cruciferaceae, Polygonaceae, Salicaceae, Tamaricaceae, Umbelliferae.

Hylaeus bisinuatus Forster, 1871. Zool.-Bot. Gesell. Wien, Verhandl. 21: 935. 2, 6.

Prosopis leptocephala Morawitz, 1871. Soc. Ent. Ross., Horae 7: 324. @, ¢.

Prosopis (Hylaeus) discrepans Schenck, 1875. Deut. Ent. Ztschr. 19: 326. ¢.

Prosopis stevensi Crawford, 1913. Canad. Ent. 45: 155. d, 9.

Prosopis incompleta Alfken, 1937. Ist. Ent. Bologna, Bol. 9: 104. 9.

Taxonomy: Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci. 180: 26-27 (as H. stevensi).
—Snelling, 1975. Los Angeles Co. Mus., Contrib. Sci. 267: 8-9 (synonymy).

Biology: Barrows, 1975. Psyche 82: 74-75 (nest).
SPECIES GROUP CONSPICUUS

conspicuus (Metz). Wash., Oreg., Idaho, Calif., Nev., Wyo. Pollen: Unknown, but visits flowers
of Anthemis cotula, Asclepias, Baccharis, Calochortus luteus, Ceanothus soredatus,
Centaurea solstitialis, Eriodictyon californicum, Eriogonum fasciculatum, Eryngium
aristatulum, Foeniculum vulgare, Frankenia grandiflora, Gnaphalium californicum,
Grindelia, Hemizonia pungens, Lasthenia, Medicago sativa, Melilotus, Phacelia distans,
Prunus ilicifolia, Salix, Salsola kali, Solidago elongata, Trichostema. Predator:
Philanthus pulcher Dalla Torre.
Prosopis conspicua Metz, 1911. Amer. Ent. Soe., Trans. 37: 114. 6, 2.
maritimus Bridwell. Calif. (coastal sand dunes). Pollen: Unknown, but visits flowers of
Achillea borealis arenicola, Agoseris aparagoides, Eriogonum, Grindelia robusta
platyphylla, Hemizonia corymbosa var. barclayi, Phacelia distans, Polygonum
paronychia.
Hylaeus maritimus Bridwell, 1910. Hawaii. Ent. Soc., Proc. 4: 159. 2, ¢.

Superfamily APOIDEA 1767

SPECIES GROUP MESILLAE

fedorica (Cockerell). Mich. to Minn., south to Tex. and east to N. C. Pollen: Unknown, but
visits flowers of Crataegus, Daucus, Pyracantha.
Prosopis digitata var. fedorica Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 27. 6.
Prosopis grossicornis Swenk and Cockerell, 1910. Ent. News 21: 67. d.
granulatus (Metz). Nev., Calif., ?Colo.
Prosopis rudbeckiae var. granulatus Metz, 1911. Amer. Ent. Soc., Trans. 37: 114. d.
labiatifrons (Cockerell). Ga. Possibly a synonym of H. cressoni cressoni (Cockerell).
Prosapis labiatifrons Cockerell, 1896. Psyche 7 (sup.): 437. 3.
mesillae cressoni (Cockerell). Transcont. from south. Canada to north. Mexico. Ecology: Nests
in twigs of Symphoricarpos. Parasite: Rhydinofoenus floridanus bradleyi Townes, R.
visaliae Bradley. Pollen: Polylectic, visits a wide variety of flowers including Achillea
millaefolium, A. lanulosa, Adenostoma fasciculatum, Alyssum maritimum, Amorpha,
Anethum, Anthemis cotula, Apium graveolens, Apocynum, Aruncus, Asclepias
fascicularis, A. mexicana, A. parishii, Aster adscendens delectabilis, Atriplex
semibaccata, Baccharis douglasii, B. emoryi, B. glutinosa, B. pilularis, Blephilia,
Boltonia, Brassica oleracea, Cacalia, Capsella, Cardamine, Castanea, Castanopsis,
Ceanothus crassifolius, C. cuneatus, C. integerrimus, Cerastium, Chamaebatia foliolosa,
Chrysothamnus consimilis, C. nauseosus speciosus, Cicuta, Cirsium vulgare, Cleome,
Cleomella obtusifolia, Cornus, Crataegus, Cryptantha angustifolia, Cryptotaenia,
Datura meteloides, Daucus carota, Descurainia sophia, Encelia farinosa, Epilobium,
Eremocarpus setigerus, Erigeron canadensis, Eriodictyon californicum, Eriogonum
elongatum, E. gracile, E. fasciculatum, E. nodosum, E. nudum, E. saxatile, E.
subscaposum, E. vimineum, E. wrightii, Eriophyllum confertiflorum, Eryngium
aristatulum, Eschscholzia caespitosa, E. californica, Eulophus, Eupatorium, Euphorbia
albomarginata, Evonymus, Foeniculum vulgare, Fragaria, Galium, Geum, Gnaphalium
californicum, Gutierrezia californica, G sarothrae, Haplopappus arborescens, H.
palmeri, Heliotropium curvassavicum, Heracleum, Heteromeles arbutifolia, Holodiscus
discolor, Hydrangea, Krigia, Lasthenia chrysostoma, L. gracilis, Lepidium,
Lepidospartum, Lotus americanus, Lupinus paynei, Lycopus, Malus, Malva, Maytenus
boaria, Melilotus alba, Mentha pulegium, Monardella lanceolata, Nemophila menziesii,
Oenothera tanacetifolia, Osmorrhiza, Oxypolis, Parthenium, Pastinaca, Perideridia
gairdneri, Phacelia distans, P. ramosissima, Philibertia heterophylla, Polygonum,
Polytaenia, Potentilla glandulosa, Prosopis, Prunus ilicifolia, Pycnanthemum,
Pyracantha, Rhamnus californica, R. crocea, Rhus diversiloba, Rosa, Rubus, Salix
exigua, S. goodingii, S. laevigata, S. lasiolepis, S. nigra, Salvia.apiana, S. mellifera, S.
officinalis, S. stachyoides, Sanicula, Schinus molle, Scrophularia californica, Sedum,
Sisymbrium irio, Sium, Solidago californica, S. occidentalis, Stanleya pinnata, Swertia
parryi, Taenidia, Tamarix gallica, Tetradymia canescens, Thaspium, Trichostema
lanceolatum, T. parishii, Valerianella, Verbena lasiostachys, Veronica, Viburnum,
Zizia.
Prosopis pygmaea Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 272. d. Preoce.
Prosopis cressoni Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 131. N. name.
Prosopis pasadenae Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 30. d, @.
Prosopis telepora Lovell, 1911. Ent. News 22: 213. 9, ¢.
Hylaeus laciniatus Cockerell and Sumner, 1931. Amer. Mus. Novitates 490: 9. ¢.
Hylaeus repolitus Cockerell and Sumner, 1931. Amer. Mus. Novitates 490: 13. ?.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 36 (as pygmaeus and cressoni). —Hicks,
1926. Colo. Univ., Studies 15: 221.

Morphology: Blum and Bohart, 1972. Ent. Soc. Amer., Ann. 65: 274 (mandibular gland
pheromones).
mesillae mesillae (Cockerell). Nebr., Kans., Tex. west to Nev., Calif.; Mexico (Baja California,
Chihuahua, Coahuila, Nuevo Leon and Sonora). Populations in Colo. and Utah are
largely sympatric with those of the nominate subspecies and hybridization occurs in
these areas. Pollen: Unknown, but visits flowers of Croton californicus, Datura

1768 | Hymenoptera in America North of Mexico

meteloides, Encelia farinosa, Eriogonum fasciculatum, E. heermannii, Melilotus alba,
Prosopis, Sphaeralcea ambigua.

Prosopis subtilis Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 295. 2. Preoce.

Prosopis mesillae Cockerell, 1896. Canad. Ent. 28: 42. N. name.

Prosopis cressoni var. magniclavis Swenk and Cockerell, 1910. Ent. News 21: 68. d, 2.

rudbeckiae (Cockerell and Casad). Canada (Ontario), transcont. U. S. from Conn.,? N. J., ?Ala.

to Rocky Mts. and Pacific Coast. Pollen: Unknown, but in California visits flowers of
Achillea millaefolium, Alyssum maritimum, Angelica arguta, Anthemis cotula,
Asclepias, Baccharis douglasii, B. emoryi, B. glutinosa, B. viminea, Calochortus
venustus, Calyptridium umbellatum, Chrysothamnus, Cirsium, Ceanothus crassifolius,
Descurainia sophia, Eriodictyon californicum, Eriogonum elatum, E. fasciculatum, E.
latifolium nudum, E. vimineum, Foeniculum vulgare, Horkelia fusca, Lasthenia
chrysostoma, Mimulus, Phacelia distans, Potentilla glandulosa, Ranunculus
californicus, Rhus diversiloba, Salix goodingti, Sedum, Sisymbrium, Solidago,
Symphoricarpos Trichostema lanatum, Veronica americana.

Prosopis rudbeckiae Cockerell and Casad, 1895. Amer. Ent. Soc., Trans. 22: 380. ¢.

Prosapis bakeri Cockerell, 1896. Psyche 7 (sup.): 26. 3.

Prosapis digitatus Cockerell, 1896. Psyche 7 (sup.): 30. 3.

Prosgapis rudbeckiae ruidosensis Cockerell, 1896. Psyche 7 (sup.): 30. 3.

Prosapis rudbeckiae subdigitata Cockerell, 1896. Psyche 7 (sup.): 31. 3.

saniculae (Robertson). Canada (Ontario, Nova Scotia), and New England States west to Minn.,

south to Tenn., Ga. Pollen: Unknown, but visits flowers of Achillea, Amorpha fruticosa,
Apocynum cannabinum, Aruncus sylvester, Cicuta maculata, Crataegus crus-galla,
Cryptotaenia canadensis, Eulophus americanus, Heracleum lanatum, Polygonum
scandens, Sanicula canadensis, S. marilandica, Taenidia integerrima, Thaspium
aureum trifoliatum.

Prosopis saniculae Robertson, 1896. Canad. Ent. 28: 137. d, 2.

sejunctus Snelling. Ariz. (Yuma), Nev. (Searchlight), south. Calif. deserts; Mexico (Mexicali).
Pollen: Unknown, but visits flowers of Croton californicus, Cryptantha barbigera,
Heliotropium curassavicum, Hyptis emoryi, Isomeris arborea Melilotus, Phacelia,
Prosopis glandulosa var. torreyanum, Salix, Tamarix gallica.
Hylaeus (Hylaeus) sejunctus Snelling, 1970. Los Angeles Co. Mus., Contrib. Sci. 180: 18,
figs. d, 9.

SPECIES GROUP ELLIPTICUS

ellipticus (Kirby). Alaska to Nova Scotia and Newfoundland, New England States south to

Ga., west to Pacific Coast States. Parasite: Aetrorys analis Ashm., Chrysis parvula
Fabr., Coelopencyrtus hylaeoleter Burks, Encyrtus sp., Gasteruption assectator (L.).
Pollen: Unknown, but visits flowers of Acer apicatum, Arenaria, Aruncus, Ceanothus,
Chrysanthemum, Eriogonum mariifolium, Houstonia purpurea, Hydrangea, Malus,
Oxypolis, Penstemon heterodoxus, P. newberryi, Potentilla glandulosa, Rhus, Rosa,
Rubus, Solidago multiradiata, Sphenosciadium capitellatum, Taraxacum. Predator:
Philanthus pulcher Dalla Torre.

Prosopis elliptica Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 266. °.

Prosopis antennata Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 271. ¢.

Prosopis varifrons Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 270. 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 71.

Biology: Davidson, 1895. Psyche 7: 315-316. —Fye, 1965. Canad. Ent. 97: 864-865, fig. 1, table
1 (nest). —Medler, 1966. Ent. Soc. Wash., Proc. 68: 131.

SPECIES GROUP VERTICALIS

verticalis (Cresson). Southeast Canada and transcont. U. S. Pollen: Unknown, but visits
flowers of Allium haematochiton, Apocynum androsaemifolium, Arctostaphylos patula,
Asclepias, Baccharis, Ceanothus cordulatus, C. integerrimus, C. soredatus, Chamaebatia
foliolosa, Collinsia heterophylla, Cryptantha intermedia, Diplacus auriantiacus,
Eriodictyon californicum, E. trichocalyx, Eriogonum, Eriophyllum multicaule,

Superfamily APOIDEA 1769

Erysimum, Euphorbia crenulata, Heracleum lanatum, Horkelia parryi, Lomatium
dasycarpum, Lupinus, Malus, Microseris nutans, Nemophila aurita, Penstemon
grinnelli, P. palmeri, Phacelia distans, P. humilis, Potentilla glandulosa, Prunus
subcordata, Rhamnus californicus, R. crocea, R. rubra, Ribes, Rosa californica, Rubus
leucodermis, Salix, Scrophularia californica, S. laciniata, Solidago canadensis, S.
multiradiata, Tamarix. Predator: Philanthus politus Say.

Prosopis verticalis Cresson, 1869. Boston Soe. Nat. Hist., Proce. 12: 271. 6.

Prosapis tridentulus Cockerell, 1896. Psyche 7 (sup.): 27. d.

Prosopis tridens Cockerell, 1896. Psyche 7 (sup.): 28. 3.

Prosopis subtristis Swenk and Cockerell, 1910. Ent. News 21: 70. 2.

Prosopis melitina Lovell, 1911. Ent. News 22: 214.9.

Biology: Fye, 1965. Canad. Ent. 97: 867, table 1 (nest).
Genus HYLAEUS Subgenus CEPHALYLAEUS Michener

Hylaeus subg. Cephalylaeus Michener, 1942. N. Y.. Ent. Soc., Jour. 50: 278.
Type-species: Prosopis basalis Smith. Orig. desig.

Revision: Snelling, 1968. Los Angeles Co. Mus., Contrib. Sci. 144: 1-3.
basalis (Smith). Transcont. in U. S., south. Canada (montane areas, alpine in southern
distribution). Pollen: Unknown, but exhibits a decided preference for flowers of the
Rosaceae; visits a wide variety of flowers including Arctostaphylos patula, Calochortus
leichtlinni, Ceanothus, Chamaebatia foliolosa, Cymopterus terebinthus, Erigeron
coulteri, Eriogonum mariifolium, Horkelia fusca, Oxypolis occidentalis, Phyllodoce
breweri, Potentilla fruticosa, P. glandulosa, Rhamnus californica, Rubus,
Sphenosciadium capitellatum, Veronica americana. Predator: Philanthus pulcher
Dalla Torre.
Prosopis basalis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 22. 2, d.

Biology: Fye, 1965. Canad. Ent. 97: 864, table 1 (nest).

nunenmacheri Bridwell. Calif. (sea level to moderate elevations in the Coast Ranges and
Sierra Nevada). Pollen: Unknown, but exhibits a decided preference for the flowers of
the Rosaceae; visits a wide variety of flowers including Arctostaphylos patula,
Baccharis, Ceanothus cordulatus, C. cuneatus, C. greggii, C. integerrimus, C. soredatus,
Chamaebatia foliolosa, Cryptantha, Eriodictyon angustifolium, E. californicum,
Phacelia distans, Prunus emarginata, Ranunculus occidentalis, Rhamnus californicus,
R. crocea, Rhus diversiloba, Salix lasiolepis, Solidago californica, Thysanocarpus
curvipes.

Hylaeus Nunemacheri Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 157. 9.

Genus HYLAEUS Subgenus METZIELLA Michener

Hylaeus subg. Metziella Michener, 1942. N. Y. Ent. Soc., Jour. 50: 273.
Type-species: Prosopis potens Metz. Monotypic and orig. desig. (=Prosopis sparsa
Cresson).

Revision: Snelling, 1968. Los Angeles Co. Mus., Contrib. Sci. 144: 3-4.
sparsus (Cresson). Canada (Quebec), eastern U.S. (Mich. to N. Y. south to Ga.) west to Tex.
Pollen: Unknown, but visits flowers of Hulophus americanus, Osmorrhiza longistylis,
Thaspium aureum trifoliatum.
Prosopis sparsa Cresson, 1869. Boston Soc. Nat. Hist., Proc. 12: 271. 2.
Prosopis thaspii Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 43. °.
Prosopis potens Metz, 1911. Amer. Ent. Soc., Trans. 37: 103. d.

Genus HYLAEUS Subgenus Unassigned

formosus Krombein. Fla. (Key Largo, Stock Is.); West Indies (Bahamas). Pollen: Unknown,
but visits flowers of Metopium toxiferum.
Hylaeus formosus Krombein, 1953. Amer. Mus. Novitates 1633: 19. 2.
Hylaeus metopii Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 80. @.

1770 Hymenoptera in America North of Mexico

Taxonomy: Krombein, 1967. U. S. Dept. Agr., Agr. Monog. 2, Sup. 2, p. 423.

graenicheri Mitchell. Fla. (Miami, Key Largo, Plantation Key, and Saddlebunch Keys). Pollen:
Unknown, but visits flowers of Metopium toxiferum, Flaveria linearis.
Hylaeus graenicheri Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 240. °.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 79. —Snelling, 1966. South.
Calif. Acad. Sci., Bul. 65: 174.

suffusus (Cockerell). Nev. Possibly a Paraprosopis.
Prosapis suffusa Cockerell, 1896. Psyche 7 (sup.): 32. ¢.

Taxonomy: Snelling, 1966. South. Calif. Acad. Sci., Bul. 65: 174. —Snelling, 1970. Los Angeles
Co., Mus., Contrib. Sci. 180: 58.

tuertonis (Cockerell). N. Mex. (Tuerto Mts. near Santa Fe), Colo. (Florissant). Possibly a
Paraprosopis and may be a synonym of H. wootoni (Cockerell).
Prosopis tuertonis Céckerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 423. 2.

Family OXAEIDAE

This small family of moderately large-sized bees occurs only in the Western Hemisphere.
Although as a group the Oxaeidae are primarily centered in the tropics, a few species are
present in the warm temperate areas of both North and South America. Some of the species
commence flight activity about sunrise and at least the males of one species, Protoxaea gloriosa
(Fox), cluster in large aggregations on selected plants where they spend the night.

The known intrafloral relationships of these bees suggest a rather narrow dependence upon
relatively few sources of pollen. These include only certain genera within the plant families
Leguminosae, Solanaceae, and Zygophyllaceae. However, the males and females seek nectar
from a comparatively wide variety of plants since the flowers of some of the preferred pollen
sources produce little or no nectar and large quantities of nectar are required to meet the
bioenergetic needs of these fast-flying bees.

Revision: Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 1-75, 68 figs., 3 pls., 2 maps, 2
tables (included genera and spp., behavior, intrafloral ecology, summary of biological
literature).

Taxonomy: Friese, 1898. K. K. Naturhist. Hofmus., Ann. 13: 59-86 (tax. characters, tax.
position). —Schrottky, 1913. Soc. Cient. Argentina, An. 75: 114-115, 180-286 (tax. position).
—Michener, 1944. Amer. Mus. Nat. Hist., Bul. 82: 246 (tax. characters, tax. status).
—Moure, 1946. Bol. Agr. 4: 12-13 (tax. characters). —Moure, 1950. Dusenia 1: 303-306 (tax.
characters, tax. status). —Rozen, 1964. N. Y. Ent. Soc., Jour. 72: 223-230, 7 figs. (tax.
characters, tax. status). —Rozen, 1965. Amer. mus. Novitates 2224: 1-18 (immature stages,
tax. status). —Michener, 1965. Amer. Mus. Nat. Hist., Bul. 180: 10 (tax. status). —Graf,
1966. Ciencia e Cultura 18: 137-138 (tax. characters, tax. status). —Linsley and Cazier,
1972. Amer. Mus. Novitates 2509: 1-25, 4 tables (tax. status). —Roberts, 1973. Kans. Ent.
Soc., Jour. 46: 437-446 (tax. status).

Biology: Linsley and Cazier, 1972. Amer. Mus. Novitates 2509: 1-25, 4 tables (behavior).
—Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 5-16, 2 tables (flower preferences,
territoriality of males, sleeping aggregations of males, nest sites, nest architecture, female
aggressiveness at nest site, parasites, immature stages).

Morphology: Popov, 1941. Acad. Sci. USSR, Compt. Rend. 30: 82-85, 4 figs. (male genitalia).
—popov, 1945. Zool. Zhur. 24: 329-336, 3 figs. (male genitalia). —Rozen, 1951. Kans. Ent.
Soc., Jour. 24: 142-150, 17 figs. (male genitalia).

Genus PROTOXAEA Cockerell and Porter

Protoxaea Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 410.
Type-species: Megacilissa gloriosa Fox. Monotypic and orig. desig.

This genus is composed of three species, two of which (P. australis Hurd and Linsley and P.
micheneri Hurd and Linsley) occur in Mexico below the elevation of Mexico City and well to the
south of P. gloriosa (Fox) which ranges over much of northern Mexico and adjacent
southwestern United States.

c

Superfamily APOIDEA 1771

Revision: Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 23-40, figs. 2-4, 17-40, map 1,
plates 1-2 (included spp. with summary of biological literature).

Biology: Linsley and Michener, 1962. Kans. Ent. Soc., Jour. 35: 385-389 (habits).
gloriosa (Fox). Tex. west to Ariz.; Mexico (Baja California, Chihuahua, Coahuila, Durango,

Nuevo leon, Sinaloa, Sonora and Tamaulipas). Parasite: Triepeolus sp. Pollen: Narrowly
polylectic, obtains pollen chiefly from the flowers of Acacia angustissima, Cassia
bauhinioides, C. leptocarpa, C. wislizenii, Desmanthus cooleyi, Kallstroemia
grandiflora, Larrea tridentata, Prosopis glandulosa var. torreyanum, Solanum
elaeagnifolium, S. rostratum, but visits some of these and a wide variety of other
flowers for nectar including Aloysia gratissima, A. wrightii, Asclepias subverticillata,
Baccharis glutinosa, Baileya multiradiata, Caesalpinia jamesii, Casuarina, Cevallia
sinuata, Conyza coulteri, Croton pottsii, Dalea scoparia, Gaillardia pulchella,
Gossypium, Guardiola tulocarpa, Helianthus annuus, Hoffmanseggia glauca,
Koeberlinia spinosa, Lepidium lasiocarpum, L. montanum, Lygodesmia juncea,
Medicago sativa, Melilotus alba, Menodora scabra, Mentzelia, Mimosa, Parkinsonia
aculeata, Psilostrophe tagetina, Salvia azurea, Solidago occidentalis, Sphaeralcea,
Tamarix gallica, T. ramosissima, Verbesina encelioides.

Megacilissa gloriosa Fox, 1893. Psyche 6: 421. @.

Oxaea Tristis Gribodo, 1894. Soc. Ent. Ital., Bol. 26: 278. 6.

Protoxaea gloriosa pallida Cockerefl, 1934. Canad. Ent. 66: 153. 2, 6.

Taxonomy: Rozen, 1964. N. Y. Ent. Soc., Jour. 72: 223-230, 7 figs. (larva and taxonomic
significance, parasite). —Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 29-38, figs. 2,
3, 25-36, map 1, plates 1-3 (redescription, synonymy, geogr. and floral records).

Biology: Linsley and Michener, 1962. Kans. Ent. Soc., Jour. 35: 385-389 (habits). —Cazier and
Linsley, 1963. Canad. Ent. 95: 547-556, 2 figs. (male territorial behavior). —Linsley and
Cazier, 1963. Pan-Pacifiec Ent. 39: 1-18, 6 figs., 2 tables (floral relationships, diurnal
activity). —Linsley and Cazier, 1972. Amer. Mus. Novitates 2509: 1-25, 4 tables (adult
behavior). —Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 1-20, figs. 1-6, tables 1-2
(foraging behavior). —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 27 (floral
relationships, territorial behavior). —Cazier and Linsley, 1975. Pan-Pacifie Ent. 51:
248-253, 6 figs., 2 tables (visitation to flowers of Kallstroemia grandiflora after two years
drought). —Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 14-16 (adult behavioral

characteristics).
Genus MESOXAEA Hurd and Linsley

Mesoxaea Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 41.
Type-species: Oxaea nigerrima Friese. Orig. desig.

As presently understood this genus contains seven species which occupy much of the Mexican
mainland to the northeast, south, and west of the Mexican Plateau although one species inhabits
the Valley of Mexico.
arizonica (Cockerell). South. Ariz.; Mexico (Nayarit, Sinaloa and Sonora). Pollen: Collects

pollen of Kallstroemia grandiflora, but also visits other flowers for nectar including
Acacia angustissima, Buddleja sessiliflora, Tephrosia leiocarpa, Turnera diffusa.
Protoxaea nigerrima arizonica Cockerell, 1936. Amer. Mus. Novitates 831: 6. 2.

Taxonomy: Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 61-64, figs. 61-64, map 3,
table 2 (redescription, geogr. records and floral preferences).
rufescens Hurd and Linsley. South. Ariz.; Mexico (Nayarit, Sinaloa and Sonora). Pollen:
Collects pollen from the flowers of Mimosa polyantha, Solanum, but visits other
flowers for nectar including Acacia angustissima, Buddleja sessiliflora, Croton
culiacanensis, Kallstroemia grandiflora, Turnera diffusa, Verbesina encelioides.
Mesoxaea rufescens Hurd and Linsley, 1976. Smithsn. Contrib. Zool. 220: 47, figs. 45-48,
map 2, table 2. d, °.
texana (Friese). La., Tex.; Mexico (Nuevo Leon). Ecology: Nests in large aggregations in areas
of sparse vegetation. Pollen: Collects pollen from flowers of Solanum rostratum, but

1772 Hymenoptera in America North of Mexico

visits other flowers for nectar including Aloysia gratissima, Croton, Eupatorium
serotinum, Gossypium, Monarda punctata, Polygonum, Sesamum indicum, Verbena

halei, Vernonia texana.
Oxaea texana Friese, 1898. K. K. Naturhist. Hofmus., Ann. 13: 85. 6.

Taxonomy: Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 6: 275. 6, 9. —Hurd and Linsley,
1976. Smithsn. Contrib. Zool. 220: 54-58, figs. 53-56, map 2, table 2 (redescription, geogr.
and floral records).

Biology: Birkmann, 1932. Giddings News, 2 September 1932: 9, p. 7, columns 6 and 7 (habits).
—Cockerell, 1933. Amer. Nat. 67: 286-288 (nest site, habits). —Hurd and Linsley, 1976.
Smithsn. Contrib. Zool. 220: 57-58 (summary of literature).

Family ANDRENIDAE

This is a large family of pollen-collecting bees which is present on all the continents except
Australia. Nearly all of the species excavate their nests in the ground and most are solitary
although a few species in each of the two subfamilies live in colonies. Their intrafloral relation-
ships are varied and range from numerous examples of oligolecty in varying degrees to broad
patterns of polylecty.

Even though both subfamilies are well represented in America north of Mexico by a great
many species, the majority of our fauna consists of the chiefly Holarctic genus Andrena and the
Nearctic genus Perdita.

Taxonomy: Rozen, 1951. Kans. Ent. Soc., Jour. 24: 142-150, 17 figs. (male genitalia).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 86-330, figs. 1, 2, 4, 15-79, tables 3-9
(eastern U.S. spp.). --Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 1-13, 18 figs.

(pupae).
SUBFAMILY ANDRENINAE
Taxonomy: Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 5-6 (pupae).

Genus ANDRENA Fabricius

Since studies of the relationships of the numerous subgenera within the genus Andrena are
incomplete, the subgenera are arranged alphabetically in this catalog. Those species currently
not assigned to any subgenus are placed after the recognized subgenera.

Taxonomy: Cockerell, 1898. N. Mex. Univ., Bul. 1: 48 (N. Mex. spp.) (reprint: Denison Univ.
Sci. Labs., Bul. 11: 48). —Cockerell, 1899. Ent. News. 10: 253 (misc.). —Cockerell, 1901.
Ent. News 12: 74 (misc.). —Cockerell, 1901. N. Y. Ent. Soc., Jour. 9: 132 (N. J. spp.).
—Robertson, 1902. Amer. Ent. Soc., Trans. 28: 187 (Ill. spp.). —Bruner, 1903. Amer. Ent.
Soc., Trans. 29: 240 (N. Amer. spp.). —Viereck, 1904. Canad. Ent. 36: 157, 189, 221 (Pacific
Northwest spp.). —Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 429-437 (Colo. spp.).
—Viereck, 1907. Ent. News 18: 280 (Conn. spp.). —Cockerell, 1907. Colo. Univ., Studies 4:
244 (Colo. spp.). —Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 10 (Colo. spp.).
—Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 1 (Rocky Mts. spp.). —Viereck,
1917 (1916). Conn. State Geol. and Nat. Hist. Survey, Bul. 22: 709-720 (Conn. spp.).
—Viereck, 1924. Canad. Ent. 56: 20 (subgenera). —Cockerell, 1930. Pan-Pacific Ent. 7: 7
(misc.). —Cockerell, 1931. Amer. Mus. Novitates 458: 1-20 (Rocky Mts. spp.). —Cockerell,
1932. Canad. Ent. 64: 155-158 (Micrandrena). —Cockerell, 1932. Pan-Pacifice Ent. 8:
173-177 (mise. Calif. spp.). —Cockerell, 1932. Canad. Ent. 64: 285-288 (sladeni group).
—Atwood, 1934. Canad. Jour. Res. 10 (2): 206-214, figs. 31-91 (Nova Scotia spp.).
—Cockerell, 1937. Amer. Mus. Novitates 948: 15 (Diandrena, pars). —Linsley, 1938. Calif.
Acad. Sci., Proc. (4) 23: 263-282 (black spp.). —Lanham, 1941. Ent. Soc. Amer., Ann. 34:
702-713 (Colo. spp.). —Lanham, 1949. Pan-Pacific Ent. 25: 33-35 (females of carlini group).
—Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 183-238 (subgenera). —Timberlake, 1951. U.S.
Natl. Mus., Proc. 101: 373-414 (mise. Calif. spp.). —Linsley and MacSwain, 1955.
Pan-Pacifie Ent. 31: 163-172 (N. Amer. spp. of Melandrena). —Linsley and MacSwain,
1956. Pan-Pacifie Ent. 32: 111-121 (Oenothera visiting spp.). —Mitchell, 1960. N. C. Agr.

Superfamily APOIDEA 1773

Expt. Sta. Tech. Bul. 141: 86-257, figs. 2, 4, 6-57, tables 3-7 (eastern U. S. spp.). —Linsley
and MacSwain, 1961. Pan-Pacifie Ent. 37: 31-41 (Diandrena spp. associated with flowers
of Oenothera). —Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 117-130
(Onagrandrena spp. associated with the flowers of Clarkia and Oenothera). —Linsley and
MacSwain, 1962. Pan-Pacific Ent. 38: 49-52 (N. spp. of Onagrandrena). —Linsley and
MacSwain, 1963. Pan-Pacific Ent. 39: 189-198 (N. spp. of Onagrandrena). —Knerer and
Atwood, 1964. Ent. Soc. Ontario, Proc. 94: 41-56 (Ontario spp.). —LaBerge, 1964. Nebr.
Univ. State Mus., Bul. 4: 279-316 (N. Amer. subgenera). —Ribble, 1967. Nebr. Univ. State
Mus., Bul. 6: 27-42, figs. 1-5, 1 table, 1 map (subg. Larandrena). —LaBerge, 1967. Nebr.
Univ. State Mus., Bul. 7: 1-316, 340 figs., 4 tables, 14 maps (subg. Callandrena).
—Warncke, 1968. Coimbra Univ. Est. Mus. Zool., Mem. 307: 1-110 (west. Palaearctic
subgenera). —Ribble, 1968. Kans. Ent. Soc., Jour. 41: 220-236, 15 figs., 1 map, 1 table (subg.
Belandrena). —Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 237-394, 124 figs., 18 maps
(subgenera Derandrena and Micrandrena). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52:
1-46, 88 figs., 5 tables, 13 maps (subg. Diandrena). —LaBerge, 1969. Amer. Ent. Soc.,
Trans. 95: 1-47, 38 figs., 3 tables (subgenera Aporandrena, Charitandrena, Plastandrena).
—LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 543-605, 75 figs., 5 maps
(subg. Tylandrena). —Warncke, 1970. Bayerischen Ent. Nachr. 19: 28-32 (lectotypic
designations). —LaBerge, 1971. Pan-Pacifie Ent. 47: 47-57, 15 figs. (subg. Nemandrena).
—LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 441-520, 68 figs., 1 table (subgenera
Rhaphandrena, Scrapteropsis, Xiphandrena). —LaBerge and Ribble, 1972. Amer. Ent.
Soc., Trans. 98: 27-358, 76 figs. (subgenera Geissandrena, Gonandrena, Parandrena,
Pelicandrena). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 235-371, 69 figs., 2 tables, 15
maps (subg. Trachandrena). —Ribble, 1974. Amer. Ent. Soc., Trans. 100: 101-189, 99 figs.,
5 tables (subg. Scaphandrena). —LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101:
371-446, 74 figs. (subg. Euandrena). —Donovan, 1977. Calif. Univ. Pubs. Ent. 81: 1-107,
130 figs., 17 maps (subg. Cnemidandrena; received too late for inclusion in this catalog).
—LaBerge, 1977. Amer. Ent. Soc., Trans. 103: 1-148, 128 figs., 3 tables (subgenera
Dasyandrena, Euandrena, Oxyandrena, Psammandrena, Rhacandrena, Thysandrena;
received too late for inclusion in this catalog).

Biology: Graenicher, 1905. Wis. Acad. Sci., Arts and Letters 15: 89-97 (floral relationships).
—Linsley, 1937. Brooklyn Ent. Soc., Bul. 32: 125-127 (occurrence of double broods in N.
Amer. spp.). —Linsley, MacSwain and Smith, 1955. Pan-Pacifie Ent. 31: 173-185 (floral
relationships of Melandrena). —Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16:
1-46, 9 plates, 6 textfigs. (Ranunculus visiting spp.). —Linsley, MacSwain and Raven,
1963. Calif. Univ. Pubs. Ent. 33: 1-58, 6 plates, 6 textfigs. (comparative behavior of spp.
visiting flowers of Onagraceae in the Colorado Desert and Great Basin). —Linsley,
MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 59-98, 3 plates, 17 tables
(comparative behavior of spp. visiting flowers of Onagraceae in the Mojave Desert).
—Stephen, 1966. Kans. Ent. Soc., Jour. 39: 42-50, 2 figs. (comparative bionomics).

— Youssef and Bohart, 1968. Kans. Ent. Soc., Jour. 41: 442-444 (literature references to
nesting habits). —Cruden, 1972. Madrono 21: 505-515, 1 fig., 2 tables (Nemophila visiting
spp.). —Cruden, 1972. Evolution 26: 373-389, 8 figs., 9 tables (oligolectie spp. of Nemophila
menziesii, evolution of oligolectic bees). —MacSwain, Raven and Thorp, 1973. Calif. Univ.
Pubs. Ent. 70: 1-80, 3 plates, 10 textfigs., 18 tables (comparative behavior of Clarkia
visiting spp.). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71:
1-68, 6 plates, 15 textfigs., 10 tables (comparative behavior of Camissonia and Oenothera
visiting spp.). —Davis and LaBerge, 1975. Ill. Nat. Hist. Survey, Biol. Notes 95: 2
(literature references to nesting habits).

Genus ANDRENA Subgenus ANDRENA Fabricius

Andrena Fabricius, 1775. Systema Ent., p. 376.
Type-species: Apis helvola Linnaeus. Desig. by Viereck, 1912.
Anthrena Illiger, 1801. Mag. Insektenk. 1: 127. Emend.
Anthocharessa Gistel, 1850. Isis (Eneyel. Ztschr.) 6: 82.
Type-species: Apis helvola Linnaeus. Autobasic; proposed unnecessarily for
Andrena Fabricius which Gistel considered to be preoccupied by Anthrenus
Geoffroy, 1764.

1774 Hymenoptera in America North of Mexico

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 205 (tax. characters). —Mitchell, 1960.
N. C. Agr. Expt. Sta. Tech. Bul. 141: 104-110, figs. 15-16, 20-22, table 3 (eastern U. S. spp.).
—Lanham, 1965. Kans. Ent. Soe., Jour. 38: 198-201 (Salix visiting spp. of the Colorado
Front Range).

advarians Viereck. B. C.
Andrena advarians Viereck, 1904. Canad. Ent. 36: 192, 224. 2, do.
albihirta (Ashmead). Rocky Mts. and Pacific Coast. Pollen: Collects pollen from Salix,
including S. laevigata, S. lasiolepis.
Cilissa albihirta Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 5. 2.
Andrena bebbiana Viereck and Cockerell, 1914. U. S. Natl. Mus., Proe. 48: 39. ?.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 8 (tax. characters). —Lanham, 1941.
Ent. Soc. Amer. Ann. 34: 705, 708. (tax characters, as bebbiana) female, male (key).
—Lanham, 1965. Kans. Ent. Soc., Jour. 38: 199-200 (tax. status, synonymy).

Biology: Lanham, 1974. Kans. Ent. Soc., Jour. 47: 373-377, 1 fig., 1 table (effect of melanism
on adult behavior, floral relationships, stylopization).
albosellata Cockerell. Wyo., Colo. Predator: Philanthus crabroniformis Sm.
Andrena albosellata Cockerell, 1931. Amer. Mus. Novitates 458: 7. °.
asmi Viereck. Wash.
Andrena asmi Viereck, 1904. Canad. Ent. 36: 192, 225. 2, 6.
banffensis Viereck. Alta.
Andrena (Andrena) banffensis Viereck, 1924. Canad. Ent. 56: 32. 9.
bella Viereck. B. C.
Andrena (Andrena) bella Viereck, 1924. Canad. Ent. 56: 22. °.
birtwelli Cockerell. N. Mex., Colo., Calif. Pollen: Unknown, but visits flowers of Antennaria,
Eriogonum, Potentilla, Solidago.
Andrena birtwelli Cockerell, 1901. Psyche 9: 283. 9.
Andrena birtwelli subatrata Cockerell, 1931. Amer. Mus. Novitates 458: 8. °.

Taxonomy: Viereck, 1903. Amer. Ent. Soc., Trans. 29: 54. 6. —Lanham, 1941, Ent. Soe.
Amer., Ann. 34: 705. 2 (key).
buckelli Viereck. B. C.
Andrena (Andrena) buckelli Viereck, 1924. Canad. Ent. 56: 22. 2, 6.
carrikeri Viereck and Cockerell. Nebr.

Andrena carrikeri Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 14. ¢.
ceanothifloris Linsley. South. Calif. Pollen: Possibly an oligolege of Ceanothus including C.
cordulatus, C. integerrimus, C. leucodermis, but visits other flowers including

Arctostaphylos patula, Eriodictyon californicum, Horkelia, Ribes.
Andrena ceanothifloris Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 270. 2, ¢.
clarkella (Kirby). Holarctic; Alaska to Ont., Que., Mass. and Maine, south to Colo., Minn. and
N. Y.; Europe. Pollen: Apparently an oligolege of Salix.
Melitta Clarkella Kirby, 1802. Monog. Apum Angliae, v. 2, p. 130. @.

Taxonomy: Sladen, 1919. Canad. Ent. 51: 124. 2, 6. —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 104-105, figs. 15-16, 21, table 3 (redescription). —Lanham, 1963. Pan-Pacific
Ent. 40: 9 (geogr. records). —Lanham, 1965. Kans. Ent. Soc., Jour. 38: 199 (floral and
geogr. records).
clypeoporaria Viereck. Wash., Oreg.

Andrena clypeoporaria Viereck, 1904. Canad. Ent. 36: 192. °.
cristata Viereck. Nev.

Andrena (Andrena) cristata Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 556. °.
diversicolor Viereck. Alta.

Andrena (Andrena) diversicolor Viereck, 1924. Canad. Ent. 56: 76. °.
edwardsi Viereck. Idaho.

Andrena edwardsi Viereck, 1916. Amer. Mus. Nat. Hist., Bul. 35: 731. 9.
edwiniae Cockerell. Colo.

Andrena edwiniae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 310. 2.

Superfamily APOIDEA 1775

Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 326: 249 (ecology).

enigmatica Viereck and Cockerell. Nebr.
Andrena enigmatica Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 33. d.
erecta Viereck. B. C. to Calif. Pollen: Unknown, but visits flowers of Chamaebatia foliolosa,
Clarkia dudleyana, Erigeron glaucus, Paeonia browni, Phacelia imbricata, Rubus,
Thysanocarpus curvipes.
Andrena (Andrena) erecta Viereck, 1924. Canad. Ent. 56: 28. 2, 6.
excellens Viereck. B. C.
Andrena (Andrena) excellens Viereck. 1924. Canad. Ent. 56: 76. °.
frigida Smith. Alaska to N. S., south to Va., Ill., Wis., Minn., Wash. and Oreg. Pollen:
Unknown, but visits flowers of Cornus, Prunus, Salix discolor.
Andrena frigida Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 115. 2.
Andrena moesta Smith, 1879. Desc. New Species Hym. Brit. Mus. p. 54. 2, d.
Andrena cockerelli Graenicher, 1903. Canad. Ent. 35: 168. 2, d.
Andrena hirticeps Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 116. 3. Preoce.
Andrena hirtignatha Linsley, 1951. Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1068. N. name.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 149. 2 (type). —Cockerell, 1906.

Psyche 13: 33 (type). —Atwood, 1934. Canad. Jour. Res. 10: 207, 209. 2, ¢ (key). —Morice
and Cockerell, 1901. Canad. Ent. 33: 149 (type of hirticeps). —Morice and Cockerell, 1901.
Canad. Ent. 33: 153 (type of moesta). —Cockerell. 1906. Psyche 13: 26 (type of hirticeps)
—Cockerell, 1906. Psyche 13: 34 (type of moesta). —Viereck, 1907. Ent. News 18: 283,
285. 2, 3 (key, as cockerelli). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
105-106, figs. 15-16, table 3 (redescription, synonymy).

fulvicrista Viereck. B. C.
Andrena (Andrena) fulvicrista Viereck, 1924. Canad. Ent. 56: 30. 2, 6.

harveyi Viereck. B. C., Oreg.
Andrena harveyi Viereck, 1904. Canad. Ent. 36: 192, 194. °, d.

hemileuca Viereck. Wash.
Andrena hemileuca Viereck, 1904. Canad. Ent. 36: 192, 198. 2, ¢.

hitei Cockerell. Colo.
Andrena hitei Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 369. °.

Taxonomy: Nelson, 1930. Ent. News 41: 322, fig. 9, d. —Lanham, 1941. Ent. Soc. Amer.,
Ann. 34: 705. 2 (key).

impuncta Kirby. Maine, N. Y.
Andrena impuncta Kirby, 1837. Faun. Bor.-Amer., v. 4, p. 268. 2.

irana Cockerell. Colo. Parasite: Nomada morrisoni Cress., N. vallesina Ckll.?
Andrena irana Cockerell, 1931. Ann. and Mag. Nat. Hist. (10) 3: 392. 9.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 265. 2 (key). —Lanham 1941. Ent.
Soc. Amer., Ann. 34: 705. 2 (key).
Biology: Hicks, 1934. Colo. Univ., Studies 21: 267 (nest, parasite).

laminibucca Viereck and Cockerell. Colo.

Andrena laminibucca Viereck and Cockerell, 1914. U.S. Natl. Mus., Proce. 48: 37. 6.
lewisii Cockerell. Colo. Pollen: Unknown, but visits flowers of Rubus.

Andrena lewisti Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 435. °.

Taxonomy: Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 57. 2 (var.). —Cockerell,
1927. Ent. Soc. Amer., Ann. 20: 397. °.
Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology).

lillooetensis Viereck. B. C.
Andrena (Andrena) lillooetensis Viereck, 1924. Canad. Ent. 56: 237. ¢.

lummiorum Viereck. B. C.
Andrena (Andrena) lummiorum Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 564. 9.

1776 Hymenoptera in America North of Mexico

macoupinensis Robertson. IIl., Mass., N. C. Pollen: Unknown, but visits flowers of Prunus,
Salix, Vicia.
Andrena macoupinensis Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 48. 2.
Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191. 2? (key). —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 106, figs. 15-16, 21, table 3 (redescription).
magnifica Viereck. B. C.
Andrena (Andrena) magnifica Viereck, 1924. Canad. Ent. 56: 77. @.
mandibularis Robertson. N.S. and Ont., south to Ga., west to Ill. and Minn. Pollen: Unknown,
but visits flowers of Amelanchier, Claytonia, Cornus, Crataegus, Hepatica, Kalmia,
Malus, Prunus, Pyrus, Rhamnus, Rhus, Salix, Staphylea, Uvularia, Viburnum,
Zanthoxylum.
Andrena mandibularis Robertson, 1892. Amer. Nat. 26: 272. 3.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191, 192. 2, d (key). —Atwood,
1934. Canad. Jour. Res. 10: 209. ¢ (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
141: 106-107, figs. 15, 22, table 3 (redescription).

Biology: Salt, 1927. Jour. Expt. Zool. 48: 252 (stylopization).
mesoleuca Cockerell. Calif. -
Andrena mesoleuca Cockerell, 1924. Pan-Pacific Ent. 1: 60. 9.
metea Cockerell. Wyo.
Andrena metea Cockerell, 1924. Ent. News 35: 350. 6.
milwaukeensis Graenicher. N. S. to Ga., west to Minn. Pollen: Unknown, but visits flowers of

Aruncus, Cornus, Ilex, Pyrus malus, Rubus, Viburnum.
Andrena milwaukeensis Graenicher, 1903. Canad. Ent. 35: 164. ?, 6.

Taxonomy: Viereck, 1907. Ent. News 18: 283, 285. 2, d (key). —Atwood, 1934. Canad. Jour.
Res. 10: 207, 209. 2, ¢ (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
107-108, figs. 15-16, 20, table 3 (redescription).

Biology: Brittain, 1933. Canad. Dept. Agr., Bul. 162: 94, figs. (ecology).
moesticolor Viereck and Cockerell. Colo.
Andrena moesticolor Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 43: 31. 2, 6.
monogonoparia Viereck. Nev.
Andrena (Andrena) monogonoparia Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 566. 3.
nitidarum Viereck. Alta.
Andrena (Andrena) nitidarum Viereck, 1924. Canad. Ent. 56: 78. 9.
nodosa Viereck. Alta.
Andrena (Andrena) nodosa Viereck, 1924. Canad. Ent. 56: 238. 2.
paenefulva Viereck and Cockerell. Colo.
Andrena paenefulva Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 51. °.
pallida Viereck. Alta.
Andrena (Andrena) pallida Viereck, 1924. Canad. Ent. 56: 78. °.
palpalis Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Calandrinia
menziesii, Cryptantha barbigera, Larrea tridentata, Phacelia affinis, P. distans, Salix
gooddingii.
Andrena (Andrena) palpalis Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 400. 2, 6.
perarmata Cockerell. B. C. to Calif. Pollen: Oligolege of Salix including S. lasiolepis, but visits
other flowers for nectar including Baccharis.
Andrena perarmata Cockerell, 1898. Entomologist 31: 88. 6.

Taxonomy: Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 7: 232. 6. —Lanham, 1941. Ent. Soe.
Amer., Ann. 34: 705, 708. 2, d (key). —Lanham, 1965. Kans. Ent. Soc., Jour. 38: 199-200
(tax. status, geogr. range).

pyrrhacita mosina Cockerell. Colo. Pollen: Oligolege of Salix.
Andrena pyrrhacita var. mosina Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 330. 2.

Taxonomy: Lanham, 1965. Kans. Ent. Soc., Jour. 38: 201 (tax. status).

Superfamily APOIDEA 1777

pyrrhacita pyrrhacita Cockerell. B. C. to Colo., Utah, Calif. Pollen: Oligolege of Salix.
Andrena pyrrhacita Cockerell, 1917. Ann. and Mag. Nat. Hist. (7) 19: 536. 9.
Andrena pyrrhacita var. coloradensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc.
48: 39. 2.

Taxonomy: Lanham, 1965. Kans. Ent. Soc., Jour. 38: 200-201 (tax. status).
revelstokensis Viereck. B. C.
Andrena (Andrena) revelstokensis Viereck, 1924. Canad. Ent. 56: 239. 2.
rhodotricha Linsley. Calif.
Andrena rhodotricha Linsley, 1939. Pan-Pacifie Ent. 15: 157. 2, 6.

Taxonomy: MacSwain, 1945. Pan-Pacific Ent. 21: 134 (nesting habits).
ribesina Cockerell. Colo. Pollen: Unknown, but visits flowers of Ribes.
Andrena ribesina Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 433. °.
Andrena jacobaea Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 267. 9.

Taxonomy: Linsley, 1951. Jn Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr.
Monog. 2: 1079 (synonymy).
ribifloris Viereck and Cockerell. Colo. Pollen: Unknown, but visits flowers of Ribes.
Andrena ribifloris Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 32. 3 (9
misdet.).
saceata Viereck. Wash. to Calif. Pollen: Unknown, but visits flowers of Eriogonum.
Andrena saccata Viereck, 1904. Canad. Ent. 36: 192, 195. 9, d.

Taxonomy: Cockerell, 1937. Amer. Mus. Novitates 899: 3 (tax. characters).
salicicola Viereck and Cockerell. Colo. Pollen: Unknown, but visits flowers of Salix.
Andrena salicicola Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 52. 9.
semifulva Viereck. Calif.
Andrena (Andrena) semifulva Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 575. 2.
singularis Viereck. B. C.
Andrena (Andrena) singularis Viereck, 1924. Canad. Ent. 56: 80. 9.
thaspii Graenicher. N. S. to N. J. west to Ind., Wis. and Minn. Pollen: Unknown, but visits
flowers of Angelica atropurpurea, Brassica, Chrysanthemum, Diervilla, Ledum, Pyrus
malus, Raphanus, Rosa, Rubus, Solidago, Stellaria, Taraxacum, Thaspium trifoliatum
aureum, Trifolium, Vaccinium.
Andrena thaspii Graenicher, 1903. Canad. Ent. 35: 162. 2, 6.

Taxonomy: Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 9: 382. 2 (var.). —Atwood, 1934.
Canad. Jour. Res. 10: 208, 209. 2, ¢d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 108-109, table 3 (redescription).

thaspiiformis Viereck. Calif. Pollen: Oligolege of Salix including S. gooddingii, S. hindsiana, S.
laevigata, S. lasiolepis, S. nigra, but visits other flowers presumably for nectar
including Sambucus caerulea, Spiraea prunifolia.

Andrena (Andrena) perezana thaspiiformis Viereck, 1917. Amer. Ent. Soc., Trans. 43: 386.
oF

tincta Viereck. Colo.

Andrena (Andrena) tincta Viereck, 1917. Amer. Ent. Soc., Trans. 48: 387. °.
topazana Cockerell. Colo., Idaho, Wyo.
Andrena topazana Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 434. °.

Taxonomy: Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 44, 45. 2. —Cockerell,
1927. Ent. Soc. Amer., Ann. 20: 397. @.
tridens Robertson. New England States, south to Tenn. and Ga., west to Mich. and III. Pollen:
Unknown, but visits flowers of Azalea, Dentaria, Hepatica, Hydrangea, Rhododendron,
Rhus.
Andrena tridens Robertson, 1902. Amer. Ent. Soc., Trans. 28: 192. d.
Andrena cornelli Viereck, 1907. Ent. News 18: 282. °.

1778 Hymenoptera in America North of Mexico

Taxonomy: Mitchell, 1958. Jn Krombein, U.S. Dept. Agr., Agr. Monog. 2, First Suppl. pp.
213, 216 (synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 109-110, figs.
15-16, 20-21, table 3 (redescription).

washingtoni Cockerell. B. C. to Wash. and Colo.
Andrena washingtoni Cockerell, 1901. Psyche 9: 284. 2, 6.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 192, 195. 2, 3 (key).
Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology).

Genus ANDRENA Subgenus APORANDRENA Lanham

Andrena subg. Aporandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 201.
Type-species: Andrena coactipostica Viereck. Monotypic and orig. desig.

Revision: LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 34-40, figs. 29-33 (N. Amer. spp.).
coactipostica Viereck. Calif. Pollen: Polylectic, apparently prefers flowers of the families
Papaveraceae and Boraginaceae in that order, but visits a variety of flowers including
Calandrinia menziesii, Capsella bursa-pastoris, Cryptantha intermedia, Encelia
farinosa, Eschscholzia californica, Lasthenia gracilis, Phacelia distans, Plagiobothrys
californicus, P. nothofulvus, Platystemon californicus.
Andrena (Andrena) coactipostica Viereck, 1917. Amer. Ent. Soc., Trans. 43: 372. 9.

Genus ANDRENA Subgenus BELANDRENA Ribble

Andrena subg. Belandrena Ribble, 1968. Kans. Ent. Soc., Jour. 41: 221.
Type-species: Andrena nemophilae Ribble. Orig. desig.

The species of this subgenus exhibit a strong preference for the flowers of the families
Hydrophyllaceae and Malvaceae.

Revision: Ribble, 1968. Kans. Ent. Soc., Jour. 41: 220-236, 15 figs., 1 map, 1 table (N. Amer.
spp.).
Biology: Cruden, 1972. Madrono 21: 505-515, 1 fig., 2 tables (floral relationships). —Cruden,
1972. Evolution 26: 373-389, 8figs., 9 tables (floral relationships).
nemophilae Ribble. Calif. Parasite: Caloglyphus sp., Meloe sp. Pollen: Oligolectic on flowers of
Nemophila including N. maculata, N. menziesti atomeria, N. m. integrifolia, N. m.
menziesii, N. pulchella, but visits other flowers for nectar including Arctostaphylos,
Ceanothus, Gilia, Montia, Pholistoma auritum.
Andrena (Belandrena) nemophilae Ribble, 1968. Kans. Ent. Soc., Jour. 41: 223, figs. 1-5,
map 1, table 1. °, d.
sagittagalea Ribble. Tex. Parasite: Vidia sp. Pollen: Apparently oligolectic on flowers of
Phacelia, but also visits flowers of Salix.
Andrena (Belandrena) sagittagalea Ribble, 1968. Kans. Ent. Soc., Jour. 41: 230, figs. 6-10,
map 1, table 1. 2, 6.
sphaeralceae Linsley. Ariz. west to south. Calif. and south. Nev. Pollen: Apparently oligolectic
on flowers of Sphaeralcea including S. ambigua, S. laxa, S. orcutti, S. subhastata, but
visits other flowers presumably for nectar including Koeberlinia, Phacelia distans.
Andrena (Opandrena) sphaeralceae Linsley, 1939. Pan-Pacifie Ent. 15: 160. 2, ¢.

Genus ANDRENA Subgenus CALLANDRENA Cockerell

Callandrena Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 186.
Type-species: Panurgus manifestus Fox. Monotypic.

Pterandrena Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193.
Type-species: Andrena pulchella Robertson. Orig. desig.

Most species of this subgenus obtain pollen from the flowers of the Compositae.

Revision: LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 1-316, 340 figs., 4 tables, 14 maps
(N. Amer. spp.).

Superfamily APOIDEA 1779

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 198 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 294 (tax. characters).
accepta Viereck. N. J. to S. C., west to Alta., Oreg., east. Calif. (Big Pine), and Ariz.; Mexico
(Chihuahua). Parasite: Myopa? or Zodion obliquefasciatum (Macq.). Pollen: Oligolege of
Compositae, especially Helianthus including H. angustifolius, H. annuus, H.
divaricatus, H. giganteus, H. grosseserratus, H. maximilliani, H. petiolaris, H.
salicifolius, H. scabberimus, H. strumosus, H. tuberosus, but visits other flowers
including Aster ericoides villosus, Bidens aristosa, Cassia chamaecrista,
Chrysothamnus, Coreopsis tripteris, Gaillardia pulchella, Grindelia squarrosa,
Gutierrezia sarothrae, Haplopappus, Machaeranthera, Medicago sativa, Prionopsis
ciliata, Silphium integrifolium, S. perfoliatum, S. speciosum, Solidago nemoralis,
Verbesina occidentalis, V. oreophila.
Andrena pulchella Robertson, 1891. Amer. Ent. Soc., Trans. 18: 57. 2. Preoce.
Andrena accepta Viereck, 1916. Biol. Soc. Wash., Proc. 29: 127. N. name.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 198, 194. 2, d (key). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 137-138, figs. 24-26, 28, table 4 (redescription).
—Rozen, 1973. N. Y. Ent. Soc., Jour. 81: 58-61, figs. 3-10 (mature larva).

Biology: Rozen, 1973. N. Y. Ent. Soc., Jour. 81: 54-58, 2 figs. (nest architecture, life history,
parasite).
afimbriata LaBerge. Tex. Pollen: Unknown, but visits flowers of Berlandiera, Polygala alba,
Pyrrhopappus carolinianus.
Andrena (Callandrena) afimbriata LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 165,
figs. 169-172. 2, 6.
aliciae Robertson. N. Y. to Ga., west to Minn. Nebr. and Kans. Pollen: Oligolege of
Compositae, especially Helianthus including H. angustifolius, H. divaricatus, H.
giganteus, H. microcephalus, H. strwmosus, H. tuberosus, but visits other flowers
including Aster azurella, Bidens aristosa, B. chrysanthemoides, B. laevis, B.
trichosperma, Cucurbita, Heliopsis helianthoides, Rudbeckia fulgida, R. laciniata, R.
lanceolata, R. triloba, Silphium perfoliatum, Solidago occidentale.
Andrena aliciae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 57. 2, ¢.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193. 2, ¢ (key). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 138-139, figs. 24-25, table 4 (redescription).
aliciarum Cockerell. N. Mex., Ariz.; Mexico (Sonora). Pollen: Unknown, but visits flowers of
Eriogonum, Pectis papposa.
Andrena aliciarum Cockerell, 1897. Entomologist 30: 138. ?.
Taxonomy: Cockerell, 1899. Ent. News 10: 254. 6.
aridis LaBerge. Colo., Nebr., N. Mex., Tex. Pollen: Unknown, but visits flowers of
Chrysothamnus nauseosus, Gutierrezia microcephala, G. sarothrae.
Andrena (Callandrena) aridis LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 249, figs.
277-281. 2, d.
asteris Robertson. Que. to Ga., west to Man., N. Dak., Nebr., and Mo. Pollen: Oligolege of
Compositae, especially flowers of Aster including A. ericoides, A. paniculatus, but visits
other flowers including Echinacea, Eupatorium altissimum, Polygonum
pennsylvanicum, Solidago nemoralis, S. puberula, S. rigida, S. ulmifolia. Predator:
Philanthus solivagus Say.
Andrena asteris Robertson, 1891. Amer. Ent. Soc., Trans. 18: 56. 2, ¢.°
Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 194. 2, d (key). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 139-140, figs. 24-26, 28, table 4 (redescription).

Biology: Salt, 1927. Expt. Zool., Jour. 48: 251 (stylopization). —Britton, 1937. Conn. (State)
Agr. Expt. Sta., Bul. 408: 261 (economic).
asteroides Mitchell. N. J. and Pa., south to S. C. and Miss. Pollen: Apparently an oligolege of
Aster including A. ericoides, A. dwmosus, A. paniculatus, but visits other flowers
including Chrysopsis, Haplopappus.
Andrena (Pterandrena) asteroides Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 140,
figs. 24, 25. 2, 6.

1780 Hymenoptera in America North of Mexico

auripes LaBerge. Ariz. (Graham Mts.); Mexico (Distrito Federal and Mexico).
Andrena (Callandrena) auripes LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 204, figs.
219-223. 2, 6.
balsamorhizae LaBerge. Nev. (Lake Mead, Overton Beach and Rodgers Springs). Pollen:
Unknown, but visits flowers of Balsamorhiza.
Andrena (Callandrena) balsamorhizae LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 252,
figs. 282-286. 2, 6.
barberi Cockerell. N. Mex. (Eagle Creek and Ruidoso Forks, White Mts.); Mexico (Durango,
Guadalajara, Guanajuato, Hidalgo, Mexico, Puebla, Tlaxcala and Vera Cruz). Pollen:
Apparently an oligolege of Compositae, including Bidens triplinervia var. macrantha,
Heterotheca, Rudbeckia laciniata, Solidago trinervata, but visits other flowers including

Argemone, Prunus.
Andrena barberi Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 448. 2°, 6.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 14. 2 (key).
beameri LaBerge. Ark., Kans. Pollen: Unknown, but visits flowers of Coreopsis grandiflora,
Echinacea angustifolia, Helenium nudiflora.
Andrena (Callandrena) beameri LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 135. 9.
berkeleyi Viereck and Cockerell. Colo., Okla., Tex. Pollen: Unknown, but visits flowers of

Actinea, Engelmannia pinnatifida.
Andrena berkeleyi Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 19. ¢.

Taxonomy: Cockerell, 1933. Pan-Pacifie Ent. 9: 153. ¢.
biscutellata Viereck. Colo., Kans., N. Mex., Tex. Pollen: Unknown, but visits flowers of
Gaillardia, Monarda. :
Andrena (Ptilandrena) biscutellata Viereck, 1917. Amer. Ent. Soc., Trans. 43: 393. 2.
braccata Viereck. New England States to Va. Pollen: Apparently an oligolege of Solidago
including S. altissima, S. nemoralis, but also visits flowers of Aster.
Andrena braccata Viereck, 1907. Ent. News 18: 284, 286. 2, do.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 141-142, figs. 24-25, table 4
(redescription).

Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology).
bullata LaBerge. Tex. (Brazos and Lee Counties). Pollen: Unknown, but visits flowers of
Gutierrezia texana, Heterotheca subaxillaris.
Andrena (Callandrena) bullata LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 154, figs.
154-158. 2, ¢.
calvata LaBerge. Ariz. (22 miles west of Springerville). Pollen: Unknown, but visits flowers of
Viguiera annua.
Andrena (Callandrena) calvata LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 74, figs.
60-64. 3.
crawfordi Viereck. Kans., Okla., Tex. Pollen: Oligolege of Compositae, tribe Cichoriae, visits
flowers of Berlandiera, Coreopsis, Engelmannia bipinnatifida, Lindheimeria texana,
Polygala alba, Pyrrhopappus carolinianus, P. geiseri, P. grandiflora, P. multicaulis,
Serinea oppositifolia, Sitilias.
Andrena crawfordi Viereck, 1909. Ent. Soc. Wash., Proc. 11: 143. 2, 6.
duplicata Mitchell. N. J., N. C., Ohio, IIl., Mo. Pollen: Apparently an oligolege of Compositae,
visits flowers of Bidens coronata, Helianthus divaricatus, H. grosseserratus, Solidago.
Andrena (Pterandrena) duplicata Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 144,
figs. 24-26, 28. 2, d.
fulvipennis Smith. N. J. to Fla. on coastal plain, also east. Tex. Pollen: Apparently an oligolege
of Compositae, visits flowers of Aster, Chrysopsis mariana, Coreopsis, Haplopappus,
Heterotheca latifolia, H. subaxillaris, Gutierrezia texana.
Andrena fulvipennis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 117. 9.

Taxonomy: Viereck, 1902. Ent. News 13: 237. 2. —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 145-146, figs. 24-25, table 4 (redescription).

Superfamily APOIDEA 1781

gardineri Cockerell. Md. to Ga., west to S. Dak., Colo. and Tex. Pollen: Apparently an oligolege

of early summer flowering Compositae, especially Senecio including S. awreus, S.
pauperculus var. balsamitae, S. smallii, S. multilobatus, S. triangularis, but visits other
flowers including Aster, Astragalus striatus, Baileya multiradiata, Crepis runcinata,
Geraea, Hymenoxys glabra, Lesquerella ovalifolia, Penstemon, Rubus.

Andrena (Pterandrena) gardineri Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 307. 9.

Andrena ashmeadi Viereck and Cockerell, 1914. U. S. Natl. Mus., Proe. 48: 45. 9.

Andrena (Opandrena) lamellicauda Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 629.
é.

Andrena campbelli Cockerell, 1933. Pan-Pacific Ent. 9: 153. 6.

Taxonomy: Lanham, 1941. Ent. Soc. Amer., Ann. 34: 711 (tax. characters). —Lanham, 1951.
In Linsley In Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr. Monog. 2: 1066
(synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 146-147, figs. 24-26,
table 4 (redescription).

Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 366: 249 (ecology).
haynesi Viereck and Cockerell. Alta. and N. Dak., south to Tex. Pollen: Oligolege of
Helianthus, mostly H. petiolaris, but also visits other flowers presumably for nectar
including Campanula, Echinacea pallida, Solidago rigida.
Andrena haynesi Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 26. °.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 14. 2 (key).
helianthi Robertson. Que. south to Va., west to Alta., Oreg., Calif. and Ariz. Pollen: Oligolege

of Compositae, apparently preferring the flowers of Helianthus including H. annuus, H.
coloradinus, H. coronatus, H. divaricatus, H. giganteus, H. grosseserratus, H.
maximilliani, H. petiolaris, H. rigidus, H. subrhomboideus, H. tuberosus, but also visits
other flowers including Aster novaeangliae, Bidens aristosa, Chrysothamnus nauseosus,
Cirsium undulatum, Cleome serrulata, Gilia, Gutierrezia sarothrae, Medicago sativa,
Rudbeckia laciniata, Silphium perfoliatum, Solidago canadensis, S. rigida, Verbesina
encelioides, V. oreophila.

Andrena helianthi Robertson, 1891. Amer. Ent. Soc., Trans. 18: 55. 2, d.

Andrena nitidior Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 406. @.

Andrena graenicheri Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 104. @.

Andrena lincolnella Viereck and Cockerell, 1914. U. S. Natl. Mus., Proe. 48: 46. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 194. 2, 6. —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 147-148, figs. 24-25 (redescription). —LaBerge, 1967. Nebr.
Univ. State Mus., Bul. 7: 94-101, figs. 8, 94-98, table 3 (redescription, synonymy).

Biology: Hicks, 1926. Colo. Univ., Studies 15: 222 (nesting habits).
helianthiformis Viereck and Cockerell. Mont. and N. Dak., south to Kans. Pollen: Apparently
an oligolege of Echinacea including E. angustifolia, E. pallida, but also visits other
flowers including Amorpha canescens, Gaillardia, Heliopsis helianthioides, Melilotus
officinalis.
Andrena helianthiformis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 26. 2.
humeralis LaBerge. Tex. Pollen: Unknown, but visits flowers of Aster, Baccharis.
Andrena (Callandrena) humeralis LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 76. 2,
3.
ignota LaBerge. S. C. (Florence).
Andrena (Callandrena) ignota LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 83. 2.
irrasus LaBerge. IIl., Nebr., Kans., Wyo., N. Mex. Pollen: Unknown, but visits flowers of
Amphiachyris dracunculoides, Aster, Chrysothamnus.
Andrena (Callandrena) irrasus LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 131, figs.
129-133. 2, d.
isocomae Timberlake. Calif. Pollen: Oligolege of Compositae, visits flowers of Chrysothamnus,
Eriogonum, Gaillardia, Gutierrezia californica, G. lucida, G. sarothrae, Haplopappus
bernardinus, H. vernonioides, Heterotheca grandiflora, Senecio douglasii, Solidago.
Andrena (Pterandrena) isocomae Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 378. °, 6.

1782 Hymenoptera in America North of Mexico

krigiana Robertson. Conn. to Fla., west to Minn., Okla. and Tex. Pollen: Oligolege of Krigia
including K. biflora, K. virginia, but visits other flowers presumably for nectar including
Cornus, Hieraceum venosum, Ranunculus acris.
Andrena krigiana Robertson, 1901. Canad. Ent. 33: 229. 2, d.
Andrena (Ptilandrena) parakrigiana Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
157. 3.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 194. 2, 6. —Viereck, 1917 (1916).
Conn. State Geol. and Nat. Hist. Survey, Bul. 22: 777. 2, ¢. —LaBerge, 1967. Nebr. Univ.
State Mus., Bul. 7: 157-162, figs. 12, 159-163 (redescription, synonymy).

levipes LaBerge. Oreg., Calif. Pollen: Collects pollen from the flowers of Lasthenia
chrysostoma and Linanthus aureus, but visits other flowers including Arenaria,
Coreopsis bigelovii, Cryptantha, Gilia, Phacelia distans.
Andrena (Callandrena) levipes LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 289, figs.
336-340. 2, 3.
melliventris Cresson. Kans. to Tex., Ariz.? Pollen: Unknown, but visits flowers of Aster
tanacetifolium, Coreopsis cardaminefolia, Gaillardia pulchella, Monarda punctata.
Andrena melliventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 257. 2.
monticola LaBerge. Ariz. (Chiricahua Mts.); Mexico (Chihuahua, Durango, Guanajuato and
Mexico). Pollen: Unknown, but visits flowers of Aster, Cirsium, Heliopsis, Solidago.
Andrena (Callandrena) monticola LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 125, figs.
119-128. 2, 6.
neomexicana LaBerge. Ariz., N. Mex. Pollen: Unknown, but visits flowers of Baileya
pleniradiata, Erigeron.
Andrena (Callandrena) neomexicana LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 246,
figs. 272-276. 2, d.
ofella LaBerge. Tex., N. Mex., Ariz. Pollen: Unknown, but visits flowers of Aplopappus
gracilis, Chrysothamnus, Gutierrezia microcephala, G. sarothrae.
Andrena (Callandrena) ofella LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 267, figs.
307-311. 2, d.
pecosana Cockerell. N. Mex., Colo., Utah, Ariz.; Mexico (Chihuahua, Hidalgo, Jalisco and
Zacatecas). Pollen: Unknown, but presumably an oligolege of Compositae, visits flowers
of Grindelia, Gutierrezia sarothrae, Helianthus, Heliopsis Viguiera annua.
Andrena pecosana Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 12: 104. 9.
Andrena townsendi Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 49. 9.
Andrena colletoides Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 27. d.

Taxonomy: LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 112-116, figs. 9, 84-88
(redescription, synonymy, floral relationships).
pectidis (Cockerell). N. Mex., Tex. Pollen: Unknown, but visits flowers of Pectis papposa.
Panurgus pectidis Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 148. °.

Taxonomy: Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 418 (tax. characters).
perpunctata LaBerge. Ariz. (Chiricahua Mts.); Mexico (Guerrero, Mexico and Morelos). Pollen:
Unknown, but visits flowers of Heterotheca subavxillaris.
Andrena (Callandrena) perpunctata LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 222,
figs. 243-247. 2, do.
placata Mitchell. Que. and Ont., south to N. C. west to Minn. and III. Pollen: Oligolege of
Solidago, but visits other flowers including Aster lateriflorum, A. macrophyllum,
Eupatorium serotinum, Fagopyrum esculentum. This species evidently hybridizes
occasionally throughout its range with the closely related A. simplex Smith.
Andrena (Pterandrena) placata Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 148,
figs. 24, 25. 6.
reflexa Cresson. Tex.; Mexico (Chihuahua, Jalisco and Michoacan). Pollen: Unknown, but visits
flowers of Crotalaria longirostrata.
Andrena reflexa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 256. ¢.
Andrena permitis Cresson, 1872. Amer. Ent. Soe., Trans. 4: 257. °.

Superfamily APOIDEA 1783

Taxonomy: LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 70-74, figs. 55-59 (redescription,
synonymy).
rudbeckiae Robertson. N. C. to Mich., Wis., Minn. and Colo., south to Tex. in the Great Plains
States. Pollen: Oligolege of Compositae, especially Ratibida including R. columnaris, R.
columnifera, R. pinnata and Rudbeckia including R. hirta, R. lanceolata, R. triloba, but
visits other flowers including Centaurea americana, Chrysopsis, Coreopsis, Echinacea
angustifolia, Gaillardia pulchella, Lepachys, Verbesina helianthioides.
Andrena rudbeckiae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 56. 2, ¢.

Taxonomy: Cockerell, 1899. Ent. News 10: 255. ¢ (key). —Robertson, 1902. Amer. Ent. Soc.,
Trans. 28: 194. ?, 3 (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 149, 151,
figs. 25-28, table 4 (redescription). —LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 90-94,
figs. 7 (redescription).

senticulosa LaBerge. Tex. Pollen: Unknown, but visits flowers of Berlandiera, Engelmannia
bipinnatifida, Pyrrhopappus, Serinia oppositifolia.
Andrena (Callandrena) senticulosa LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 177,
189-193. 2, d.

simplex Smith. N. H. to N. C., west to Minn., Nebr. and Kans. Pollen: Oligolege of Compositae,

especially Aster and Solidago, visits flowers of Amphiachyris dracunculoides, Aster
anomalus, A. azureus, A. commutatus, A. ericoides, A. e. vilosus, A. laterifolius, A.
multiflorus, A. novaeangeliae, A. paniculatus, A. praelatus, Boltonia asteroides,
Eupatorium perfoliatum, Euthamia graminifolia, Gnaphalium polycephalum,
Grindelia, Helianthus tuberosus, Polygonum hydropiperoides, P. scandens, Solidago
altissima, S. canadensis, S. graminifolia, S. lanceolata, S. nemoralis, S. rigida, S.
serotina. Predator: Philanthus solivagus Say. This species evidently hybridizes
occasionally throughout its range with the closely related A. placata Mitchell.

Andrena simplex Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 114. 9.

Andrena solidaginis Robertson, 1893. Amer. Ent. Soc., Trans. 18: 55. 2, ¢.

Andrena radmitricha Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 51. @.

Andrena (Ptilandrena) determinata Viereck, 1917. Amer. Ent. Soc., Trans. 43: 394. °.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 124 (type). —Cockerell, 1906. Psyche
13: 8 (type). —LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 137-148, figs. 10, 134-138,
table 4 (redescription, synonymy, floral relationships).

simulata Smith. Ariz., N. Mex., Colo.; Mexico. Pollen: Unknown, but visits flowers of Aster,
Viguiera annua.
Andrena simulata Smith, 1879. Deser. n. spp. Hymen. Brit. Mus., p. 52. 2.
Andrena aureocincta Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 88. 2, 6.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48. 5 (as awreocincta).
—LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 189-193, figs. 204-208 (redescription,
synonymy).

sitiliae Viereck. Okla., Tex. Pollen: Possibly an oligolege of Pyrrhopappus including P.
carolinianus, but also visits Coreopsis tinctoria.
Andrena sitiliae Viereck, 1909. Ent. Soc. Wash., Proc. 11: 144. 2.

sonorensis LaBerge. Ariz. (Show Low), N. Mex. (Carizozo). Pollen: Unknown, but visits
flowers of Gutierrezia microcephala.
Andrena (Callandrena) sonorensis LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 110,
figs. 99-103. 2, ¢.
tegularis LaBerge. Ariz. (Chiricahua and Santa Rita Mts.). Pollen: Unknown, but visits flowers
of Heliopsis.
Andrena (Callandrena) tegularis LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 195, figs.
209-213. 2, do.
texana Cresson. Tex. (Roosevelt and Victoria); Mexico (San Luis Potosi). Pollen: Unknown, but
visits flowers of Amphiachyris, Aster.
Andrena texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 258. ¢.
tonkaworum Viereck. Nebr., Kans., Colo., Tex., N. Mex. Pollen: Possibly an oligolege of
Engelmannia.
Andrena (Ptilandrena) tonkaworum Viereck, 1917. Amer. Ent. Soc., Trans. 43: 396. °.

1784 Hymenoptera in America North of Mexico

trimaculata LaBerge. Ariz. (Canelo); Mexico (Durango).
Andrena (Callandrena) trimaculata LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 68,
figs. 50-54. 2, d.
utahensis LaBerge. Utah, Colo., Ariz. Pollen: Unknown but visits flowers of Geraea.
Andrena (Callandrena) utahensis LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 258, figs.
292-296. 2, 6.
verbesinae Viereck and Cockerell. Tex. (Cotulla and San Antonio). Pollen: Unknown, but visits

flowers of Verbesina encelioides.
Andrena verbesinae Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 1. 2, d.

verecunda Cresson. Tex. Pollen: Unknown, but visits flowers of Engelmannia bipinnatifida,
Gaillardia, Pyrrhopappus carolinianus, P. grandiflora.
Andrena verecunda Cresson, 1872. Amer. Ent. Soc., Trans. 4: 257. °.

vulpicolor Cockerell. Wyo., south to N. Mex., west to Oreg. and Calif. Pollen: Apparently an
oligolege of Chrysothamnus including C. nauseosus, C. n. albicaulis, C. n. consimilis, C.
parryi, C. viscidiflorus pumilis, C. v. stenophyllus, but also visits flowers of Erigeron

neomexicana.
Andrena vulpicolor Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 512. °.
Andrena nubilipennis Viereck, 1904. Canad. Ent. 36: 193. °.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 49. 2 (key). —Cockerell, 1931.
Amer. Mus. Novitates 458: 13. 2 (key). —LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7:
106-110, figs. 6, 79-83 (redescription, synonymy).

Genus ANDRENA Subgenus CHAULANDRENA LaBerge

Andrena subg. Chaulandrena LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 314.
Type-species: Andrena porterae Cockerell. Orig. desig.
Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 284, 288, 314-315 (tax. characters).

porterae Cockerell. Sierra Nevada, Cascade and Rocky Mts. to N. Mex. Parasite: Stylops sp.
Pollen: Apparently oligolectic, presumably gathers pollen from the flowers of Ribes.
Andrena porterae Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 401. °.
Andrena leptanthi Viereck and Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 27. d.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 264, 267. 2, d.
Biology: Linsley, 1937. Pan-Pacific Ent. 13: 157 (stylopization).
Genus ANDRENA Subgenus CNEMIDANDRENA Hedicke
Andrena subg. Cnemidandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 212.
Type-species: Melitta nigriceps Kirby. Orig. desig.

Revision: Donovan, 1977. Calif. Univ. Pubs. Ent. 81: 1-107, 130 figs., 17 maps (N. Amer. spp.;
received too late for inclusion in this catalog).

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 211 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 303 (tax. characters).

antonitonis Viereck and Cockerell. Colo.
Andrena antonitonis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 56. 2.

apacheorum Cockerell. Rocky Mts. to N. Mex., Ariz.
Andrena apacheorum Cockerell, 1897. Entomologist 30: 306. °.
Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 14, 15. 2, d (key).
Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology). —Hicks,
1926. Univ. Colo., Studies 15: 221 (habits).

autumnalis Viereck and Cockerell. Nebr.
Andrena autumnalis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 44. 9°.

beulahensis Viereck. N. Mex.
Andrena beulahensis Viereck, 1903. Amer. Ent. Soc., Trans. 29: 53. 9.

Superfamily APOIDEA 1785

canadensis canadensis Dalla Torre. N.S. to N. Y. and N. J., west to Minn. Pollen: Unknown,
but visits flowers of Aster, Epilobiwm, Solidago.
Andrena simulata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 318. 2, d.
Preoce.
Andrena canadensis Dalla Torre, 1896. Cat. Hym., v. 10, p. 107. N. name.
Andrena (Pterandrena) persimilis Graenicher, 1904. Ent. News 15: 66. @.

Taxonomy: Viereck, 1907. Ent. News 18: 282, 286. 2, ¢ (key). —Atwood, 1934. Canad. Jour.
Res. 10: 207, 209, fig. 2, ¢ (key). —Timberlake, 1951. In Muesebeck, Krombein and
Townes, U.S. Dept. Agr., Agr. Monog. 2: 1059 (synonymy). —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 172, figs. 30-31, 33, table 4 (redescription).

Biology: Salt, 1927. Jour. Expt. Zool. 48: 245 (stylopization).
canadensis oslarella Viereck and Cockerell. Colo., N. Mex.
Andrena canadensis oslarella Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 28.
3.

Biology: Clements and Long, 1923. Carnegie Inst. Wash. Pub. 336: 249 (ecology).
chromotricha Cockerell. N. Mex., Wis.
Andrena chromotricha Cockerell, 1899. Entomologist 32: 128. 2.
Andrena clypeonitens Cockerell, 1902. Canad. Ent. 34: 47. 2.
Andrena truncata Viereck, 1903. Amer. Ent. Soc., Trans. 29: 53. 9.

Taxonomy: Graenicher, 1910. Canad. Ent. 42: 160. ¢. —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 172-173, table 4 (redescription of female).
citrinihirta Viereck. Calif. Pollen: Unknown, but visits flowers of Chrysothamnus,
Heterotheca grandiflora, Solidago californica.
Andrena (Andrena) citrinihirta Viereck, 1917. Amer. Ent. Soc., Trans. 43: 371. °.
colletina Cockerell. B. C. to Calif., east to Colo. Pollen: Unknown, but visits flowers of
Achillea millefolium, Chrysothamnus nauseosus consimilis, C. viscidiflorus typicus,
Haplopappus bloomeri var. angustatus.
Andrena colletina Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 454. 6.

Taxonomy: Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 5: 260. 2. —Cockerell, 1931. Amer.
Mus. Novitates 458: 14, 15. 2, d (key).
columbiana Viereck. B. C. to Calif., N. Mex. Pollen: Unknown, but visits flowers of Achillea
lanulosa, Erigeron coulteri, E. pygmaeus, Microseris nutans, Senecio aureus var.
subnudus, S. canus, S. integirrimus, S. scorzonella.
Andrena (Andrena) columbiana Viereck, 1917. Amer. Ent. Soc., Trans. 43: 374. °.
costillensis Viereck and Cockerell. Colo., N. Mex.
Andrena costillensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 50. °.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 14. °, 6.

davidsoni Viereck and Cockerell. Calif.
Andrena davidsoni Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 52. 2.

Taxonomy: Cockerell, 1930. Pan-Pacifie Ent. 7: 8. 2 (key).
hirticincta Provancher. Alta. to N. S., south to Ga. Pollen: Unknown, but visits flowers of
Aster, Epilobium, Solidago.
Andrena fimbriata Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 116. 2, ¢. Preoce.
Andrena hirticincta Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 308. 2,
3.
Anthrena americana Dalla Torre, 1896. Cat. Hym., v. 10, p. 102. N. name.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 149 (type of fimbriata). —Cockerell,
1906. Psyche 13: 9 (types of fimbriata). —Atwood, 1934. Canad. Jour. Res. 10 (2): 207, 209,
fig. 2, d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 173-174, figs. 30-31,
33-34, table 4 (redescription).

Biology: Salt, 1927. Jour. Expt. Zool. 48: 233, figs. (stylopization).

indecisa Cockerell. B. C., Alta., Mont.
Andrena costillensis indecisa Cockerell, 1937. Canad. Ent. 69: 34. °.

1786 Hymenoptera in America North of Mexico

mentzeliae Cockerell. Colo., N. Mex. Pollen: Unknown, but visits flowers of Mentzelia.
Andrena mentzeliae Cockerell, 1897. Entomologist 30: 307. 2.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 14, 15. 2, ¢. —Lanham, 1941. Ent.
Soe. Amer., Ann. 34: 706, 708. 2, 3d (key).
nubecula Smith. East. Canada., south to Ga., west to B. C. Pollen: Unknown, but visits flowers
of Aster, Euthamia, Solidago. Predator: Philanthus solivagus Say.
Andrena nubecula Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 117. 2.
Andrena nubecula tristicornis Cockerell, 1931. Canad. Ent. 63: 22. 9.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 150 (type). —Robertson, 1902. Amer.
Ent. Soc., Trans. 28: 191, 192. 2, 3 (key). —Cockerell, 1906. Psyche 13: 9 (type).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 174-175, figs. 31, 33-34, table 4
(redescription).

pacta Viereck. N. Mex.
Andrena pacta Viereck, 1903. Amer. Ent. Soc., Trans. 43: 54. 2, 6.
peckhami Cockerell. Minn. to Vt., south to N. C. Pollen: Unknown, but visits flowers of
Baptisia, Solidago.
Andrena peckhami Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 105. @.

Taxonomy: Graenicher, 1910. Canad. Ent. 42: 159. 2, 6. —LaBerge and Ribble, 1972. Amer.
Ent. Soc., Trans. 98: 272 (subgeneric position).

ramaleyi Cockerell. Colo.
Andrena ramaleyi Cockerell, 1931. Ann. and Mag. Nat. Hist. (10) 7: 346. 9.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 14 (key).
robervalensis Mitchell. Que., Mich., Minn. Pollen: Unknown, but visits flowers of Melilotus
alba.
Andrena (?Cnemidandrena) robervalensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
14 e175 soe
scutellinitens Viereck. Oreg., Calif., Nev. Pollen: Unknown, but visits flowers of
Chrysothamnus nauseosus speciosus, C. viscidiflorus typicus, Haplopappus bloomeri
var. angustatus, Oxypolis occidentalis, Solidago elongata.
Andrena (Andrena) scutellinitens Viereck, 1916. Acad. Nat. Sci. Phila, Proc. 68: 573. 2, 3.

Taxonomy: LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 294 (subgeneric position).
segregans Cockerell. N. Mex.
Andrena segregans Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 404. 2, d.
surda Cockerell. Wyo., Colo., Oreg., Calif., N. Mex. Pollen: Unknown, but visits flowers of
Chrysothamnus nauseosus, C. viscidiflorus typicus, Erigeron pygmaeus,
Sphenosciadium capitellatum.
Andrena hirticincta var. surda Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 264. 6.
Andrena pertarda Cockerell, 1916. Entomologist 49: 157. ¢.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 13, 14. 2, d (key). —Lanham, 1941.
Ent. Soc. Amer., Ann. 34: 706, 708. 2, ¢ (key). —Timberlake, 1951. Jn Linsley In
Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2: 1082 (synonymy).

xanthigera Cockerell. N. Mex.
Andrena xanthigera Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 402. 9.
Andrena albovirgata Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 403. 9.

Taxonomy: Cockerell, 1931. Canad. Ent. 63: 22. 2. —Cockerell, 1931. Amer. Mus. Novitates
458: 19. 2.
Genus ANDRENA Subgenus CONANDRENA Viereck

Andrena subg. Conandrena Viereck, 1924. Canad. Ent. 56: 20.
Type-species: Andrena bradleyi Viereck. Monotypic and orig. desig.

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 206 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 284, 288, 300 (tax. characters).

Superfamily APOIDEA 1787

bradleyi Viereck. Colo. to N. S., south to Ga. Pollen: Unknown, but visits flowers of Prunus,
Pyrus, Rhododendron, Ribes, Vaccinium.
Andrena bradleyi Viereck, 1907. Ent. News 18: 285, 286. 2, d.
Andrena saccharina Cockerell and Rohwer, 1907. Ann. and Mag. Nat. Hist. (4) 20: 128. d.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 206, 208, figs. 2, ¢ (key). —Timberlake,
1951. Jn Linsley In Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2:
1059 (synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 226-228, figs. 40,
46-47, 49-50 (redescription).

Biology: Salt, 1927. Jour. Expt. Zool. 48: 249 (stylopization).

carolina Viereck. Minn. to N.S., south to Tenn. and Ga. Pollen: Unknown, but visits flowers of
Azalea, Claytonia, Dentaria, Ledum, Pyrus, Rhodora, Vaccinium.
Andrena carolina Viereck, 1909. Ent. News 20: 126. °.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 208, 209. 2, ¢6. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 228-229, figs. 40, 45-46, 49 (redescription, flower records).

cheyennorum Viereck and Cockerell. Nebr., Wyo., Idaho.
Andrena cheyennorum Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 20. 2, d.
Andrena angustifrons Cockerell, 1934. Pan-Pacific Ent. 9: 155. 2.

Taxonomy: Timberlake, 1951. Jn Linsley In Muesebeck, Krombein and Townes, U. S. Dept.

Agr., Agr. Monog. 2: 1061 (synonymy).

durangoensis Viereck and Cockerell. Colo. to N. S., south to W. Va. Ecology: Unknown, but

visits flowers Ilex, Rubus.

Andrena durangoensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 36. 6.
Andrena (Andrena) media Viereck, 1922. Boston Soc. Nat. Hist., Occas. Papers 5: 41. d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 229-230, figs. 40, 45, 46
(redescription, synonymy).
rufosignata Cockerell. Minn. to N. B. and New England states.
Andrena rufosignata Cockerell, 1902. Canad. Ent. 34: 46. °.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 230, fig. 49 (redescription,
subgeneric assignment). —Knerer and Atwood, 1963. Canad. Ent. 95: 585, figs. 5, 7. d.

Genus ANDRENA Subgenus DACTYLANDRENA Viereck

Andrena subg. Dactylandrena Viereck, 1924. Canad. Ent. 56: 20.
Type-species: Andrena (Dactylandrena) maura Viereck. Monotypic and orig. desig.

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 204 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 285, 288, 299 (tax. characters).
berberidis Cockerell. Colo. Pollen: Unknown, but visits flowers of Berberis.
Andrena milwaukeensis var. berberidis Cockerell, 1905. Canad. Ent. 37: 371. 9.
Taxonomy: Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 13: 9. ¢. —Lanham, 1941. Ent. Soe.
Amer., Ann. 34: 705, 707. 2, d (key).
caliginosa Viereck. Calif. Pollen: Unknown, but possibly an oligolege of Ribes.
Andrena (Andrena) caliginosa Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 552. 9.
Andrena (Dactylandrena) maura Viereck, 1924. Canad. Ent. 56: 31. ¢ (2 misdet.).
Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 267. 2, d (key).

submaura Linsley. South. Calif. (The Gavilan and Sandbergs). Pollen: Unknown, but possibly
gathers pollen from Ribes indecorum.
Andrena submaura Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 269. d, °@.

Genus ANDRENA Subgenus DERANDRENA Ribble

Andrena subg. Derandrena Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 333.
Type-species: Andrena vandykei Cockerell. Orig. desig.

Revision: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 333-367, figs. 90-124, maps 11-18,
tables 12-14 (N. Amer. spp.).

1788 | Hymenoptera in America North of Mexico

arctostaphylae Ribble. Calif. Pollen: Apparently an oligolege of Arctostaphylos including A.
crustacea, A. glauca, A. mariposa, A. pungens.
Andrena (Derandrena) arctostaphylae Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 344,
figs. 95-99, map 12. 2, 6.
californiensis Ribble. Calif. Pollen: Unknown, most often collected on flowers of Salix
including S. argophylla, S. lasiolepis, but visits a wide variety of flowers including
Arctostaphylos pungens, Cryptantha intermedia, Descourainia sophia, Eriodictyon
californicum, Gilia multicaulis, Lasthenia chrysostoma, Lomatium dasycarpum,
Orthocarpus, Plagiobothrys nothofulvus, Quercus, Rhamnus crocea, Rhus trilobata,
Ribes indecorum, Salvia columbariae, Sisymbrium irio, Tamarix gallica, Trifolium.
Andrena (Derandrena) californiensis Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 348,
figs. 100-104, map 18, table 13. 2, d.
hermosa Ribble. Calif. (Central Valley). Pollen: Unknown, but possibly oligolectic on
Lasthenia.
Andrena (Derandrena) hermosa Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 360, figs.
115-119, map 18. 2, 6.
murietae Ribble. Calif. (11 miles northwest of California Hot Springs). A. penutiani Ribble
may be the male.
Andrena (Derandrena) murietae Ribble, 1968. Nebr. Univ. State Mus. Bul. 8: 346. °.
penutiani Ribble. Cent. Calif. Pollen: Unknown, but visits flowers of Lasthenia. A. murietae
Ribble may be the female.
Andrena (Derandrena) penutiani Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 352, map
15. 3.
timberlakei Cockerell. Calif. (San Benito and Tulare Cos., south to Riverside Co.). Pollen:
Based upon the mouth parts of the female, this species evidently collects pollen from
the flowers of Cryptantha including C. intermedia, but visits other flowers presumably
for nectar including Descourainia sophia, Encelia farinosa, Lomatium dasycarpum,
Plagiobothrys californicus, P. nothofulvus, Rhamnus crocea.
Andrena timberlakei Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 300. ?, 6.

Taxonomy: Cockerell, 1931. Canad. Ent. 64: 158. 9, d.
vandykei Cockerell. South. Oreg. (12 miles southwest of Keno), Calif.; Mexico (Baja California).
Pollen: Unknown, most often collected on flowers of Arctostaphylos including A.
crustacea, A. glauca, A. mariposa, A. pungens as well as Ceanothus including C.
arboreus, C. cuneatus, C. foliosus, but visits other flowers including Baccharis,
Phoradenron villosum, Rhamnus crocea, Rhus diversiloba, R. ovata, Salix, Tamarix,
Xylococcus bicolor.
Andrena (Micrandrena) solutula Cockerell, 1936. Pan-Pacific Ent. 12: 150. °.
Andrena (Micrandrena) vandykei Cockerell, 1936. Pan-Pacific Ent. 12: 151. 6.
viridissima Ribble. Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Brassica, Lasthenia tenella, Nemophila racemosa, Phacelia ciliata, P. distans, P.
douglasii, Pholisma racemosa, Ranunculus canus.
Andrena (Derandrena) viridissima Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 360, figs.
115-119, map 18. °, d.
ziziaeformis Cockerell. Mass. to Ga., Tenn., Ill. Pollen: Unknown, but visits flowers of
Potentilla including P. canadensis, Rubus, Waldsteinia.
Andrena ziziaeformis Cockerell, 1908. Canad. Ent. 40: 234. 2.

Taxonomy: Cockerell, 1932. Canad. Ent. 64: 157. 2 (key). —Mitchell, 1960. N. C. Agr. Expt.
Sta. Tech. Bul. 141: 169-171, figs. 29-32 (redescription).
Genus ANDRENA Subgenus DIANDRENA Cockerell

Diandrena Cockerell, 1903. Psyche 10: 75.
Type-species: Panurgus chalybaeus Cresson. Orig. desig.

Revision: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 1-146, 88 figs, 5 tables, 13 maps (N. Amer.
spp., includes comparative ecological information).

Superfamily APOIDEA 1789

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 219-220 (tax. characters). —Linsley and
MacSwain, 1961. Pan-Pacific Ent. 37: 31-41 (Camissonia visiting spp.). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 288, 287, 309-310 (tax. characters). —Linsley, MacSwain,
Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 13-26, figs. 6-11 (Camissonia visiting

spp. of Calif. and Baja Calif.).

Biology: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 74-75, pls. 1-3 (floral
relationships of Mojave Desert spp.). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 14-43,
124-128, figs. 1-2, tables 1-5 (nest architecture, life history, comparative ecology, parasites,
predators). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 13-26,
47-50, tables 1-2 (life history, floral relationships of Camissonia visiting spp.).

ablegata (Cockerell). Idaho, Wyo., Colo., Utah Calif. Pollen: Oligolectic on ligulate Compositae,
most likely collects pollen from early morning-opening flowers of the subtribe
Microseridinae including Agoseris glauca, Microseris nutans, Taraxacum officinale, but
visits other flowers including Camissonia subacaulis, Cardaria draba, Lomatium
dissectum.
Diandrena ablegata Cockerell, 1922. Amer. Mus. Novitates 40: 1. 9.
Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 32-33 (tax. characters).
—Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 93-95, figs. 1, 29-30, 56, map 7 (redescription).
agoseridis Thorp. Calif. Pollen: Collects pollen from ligulate Compositae, primarily from
Agoseris heterophylla, Anisocoma acaulis, but visits other flowers including Calycoseris
parryi, Camissonia campestris campestris, C. pallida pallida, Descourainia sophia,
Erodium cicutarium, Euphorbia albomarginata, Hesperochiron, Lasthenia
chrysostoma, Layia glandulosa, Malacothrix californica, M. glabrata, Phacelia ciliata,

Rafinesquia neomexicana, Sisymbrium, Tamarix gallica.
Andrena (Diandrena) agoseridis Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 116, figs. 1, 49-50,

56, map 12. @, d.

anatolis Linsley and MacSwain. South. Calif.; Mexico (Baja California). Pollen: Collects pollen
from morning-opening flowers of Camissonia including C. bistorta, C. campestris, C.
cheiranthifolia suffruticosa, but visits other flowers including Calandrinia ciliata,
Camissonia crassifolia, Ceanothus crassifolius, Cryptantha intermedia, Potentilla,

Sitanion for nectar.
Andrena (Diandrena) anatolis Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 36. 2, d.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 98-99, figs. 1, 33-34, 56, map 8. 2, d

(redescription).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 99, fig. 1 (floral relationships). —Linsley,
MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 21-22, 49, fig. 9 (floral
relationships).

apasta Linsley and MacSwain. Calif. Pollen: Collects pollen from morning-opening flowers of
Camissonia, principally from C. campestris, but occasionally from C. ovata in
northernmost localities; also visits other flowers including Agoseris heterophylla,
Brassica, Descourainia sophia, Layia, Monolopia lanceolata for nectar.
Andrena (Diandrena) apasta Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 34. °.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 118-119, figs. 1, 51-52, 56, 68, 80, map 10

(redescription).

Biology: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 75, fig. 11 (nest,
floral relationships). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 99 (floral relationships).
—Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 26, 50, fig. 11

(floral relationships). .
blennospermatis Thorp. Calif. Pollen: Collects pollen from flowers of Blennosperma including

B. bakeri, B. nanuwm.
Andrena (Diandrena) blennospermatis Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 81, figs. 1,

19-20, 56, 63, 75. 2, d.
chalybaea (Cresson). Calif. (Central coast, from Mendocino to San Luis Obispo Counties).

Parasite: Nomada sp. Pollen: Collects pollen only from the early morning-opening

1790 Hymenoptera in America North of Mexico

flowers of Camissonia ovata, but visits other flowers for nectar including
Blennosperma bakeri, Brassica, Cryptantha, Ranunculus californicus.
Panurgus chalybaeus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 61. ?, d.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 57-61, figs. 1, 3-4, 55-58, 69, 85-86, map 1
(redescription, post defecating larva).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 5-14, 59, fig. 2 (nest architecture, floral
relationships, parasite). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs.
Ent. 71: 15-17, 48, fig. 6, pl. 1 (nest, floral relationships, mating).

chalybioides (Viereck). Wash., Oreg., Calif. Parasite: Myopa rubida (Bigot)? Pollen: Collects

pollen almost exclusively from ligulate Compositae in the early morning, primarily from
Agoseris including A. heterophylla, occasionally from introduced weedy species of
Cichorieae including Hypochoeris radicata, Taraxacum officinale, but also visits other
flowers for nectar including Blennosperma, Ceanothus, Convolvulus, Daucus, Lasthenia
chrysostoma, Layia platyglossa, Linum, Ranunculus orthorhynchus.

Parandrena chalybioides Viereck, 1904. Canad. Ent. 36: 229. 9, 6.

Andrena (Parandrena) perchalybea Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 591. 2.

Diandrena purdyi Cockerell, 1936. Pan-Pacific Ent. 12: 155. 6.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 31, 33 (synonymy). —Thorp,
1969. Calif. Univ. Pubs. Ent. 52: 83-85, figs. 1, 21-22, 56 (redescription).

Biology: MacSwain and Bohart, 1947. Pan-Pacific Ent. 23: 30 (parasite). —Thorp, 1969. Calif.
Univ. Pubs. Ent. 52: 34, 85 (floral relationships, parasite).

chlorosoma Linsley and MacSwain. Calif. (Central Coast Ranges, from Mendocino County to
Ventura County). Pollen: Collects pollen from early morning opening ligulate
Compositae including Agoseris grandiflora, A. heterophylla as well as from introduced
weedy species of Cichorieae including Hypochoeris radicata, but visits these and other
flowers for nectar including Camissonia ovata, Hemizonia, Lasthenia chrysostoma,
Layia chrysanthemoides, Ranunculus californicus, Taraxacum officinale.

Andrena (Diandrena) subchalybea chlorosoma Linsley and MacSwain, 1961. Pan-Pacific
lDyati, BIR B¥4, Oe Se

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 107-111, figs. 1, 41-44, 56 (redescription).

Biology: Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 18, 30, 31 (floral

relationships, as purdyi). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 110, 111 (floral
relationships).

cuneilabris Viereck. Oreg., Calif. Parasite: Myopa perplexa Camras, M. rubida (Bigot), Stylops
pacifica Bohart? Pollen: Collects pollen and nectar chiefly from Ranunculus including
R. californicus, R. occidentalis, R. orthorhynchus, occasionally collects pollen from
Platyglossa californicus during periods when Ranunculus pollen is scarce or absent;
visits other flowers for nectar including Amsinckia, Agoseris grandiflora, A.
heterophylla, Blennosperma bakeri, Brassica, Calandrinia ciliata, Camissonia ovata, C.
tanacetifolia, Geum macrophyllum, Lasthenia, Lomatium, Wyethia.

Andrena (Parandrena) cuneilabris Viereck, 1926. Calif. Acad. Sci., Proe. (4) 15: 400. @.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 64-67, figs. 1, 7-8, 56, 59, 70, 81-82, map 2
(redescription).

Biology: MacSwain and Bohart, 1946. Pan-Pacifie Ent. 23: 30 (parasite). —Camras and Hurd,
1957. Calif. Ins. Survey, Bul. 6: 39 (parasite). —Linsley and MacSwain, 1959. Calif. Univ.
Pubs. Ent. 16: 2-31, figs. 4, 6 (floral relationships, parasites). —Thorp, 1969. Calif. Univ.
Pubs. Ent. 52: 34, 66-67 (floral relationships, parasites).

cyanosoma (Cockerell). Calif.; Mexico (Baja California). Pollen: Collects pollen exclusively
from the flowers of Camissonia including C. bistorta, C. campestris, C. cheiranthifolia
suffruticosa, C. dentata, C. sierrae sierrae, C. spiralis, but visits these and other flowers
for nectar including Cryptantha intermedia, Lasthenia aristata, L. coronaria, Rhus
trilobata, Salix.

Diandrena cyanosoma Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 49. 2, 3.

Superfamily APOIDEA 1791

Andrena (Parandrena) austrocalifornica Viereck, 1917. Acad. Nat. Sci. Phila., Proce. 68:
587. 9.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 36 (synonymy). —Thorp, 1969.
Calif. Univ. Pubs. Ent. 52: 99-102, figs. 1, 35-36, 56, 65, 77, map 9 (redescription).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 101 (floral relationships). —Linsley,
MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 23-24, fig. 10 (floral
relationships).

eothina Linsley and MacSwain. Calif. (Merced County to Ventura County). Pollen: Collects

pollen principally from Camissonia campestris and C. sierrae sierrae and also from C.
boothii decorticans, C. contorta, C. graciliflora, but visits these and other flowers for
nectar including Agoseris heterophylla, Amsinckia, Encelia virginiensis actoni,
Lasthenia chrysostoma, Layia, Monolopia lanceolata, Tropidocarpum gracile.

Andrena (Diandrena) eothina linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 39. 2, 6.

Andrena (Diandrena) anatolis matutina Linsley and MacSwain, 1961. Pan-Pacific Ent. 37:
38: 2, 6.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 95-97, figs. 1, 31-32, 56, map 8
(redescription, synonymy).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 97 (floral relationships). —Linsley,
MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 22-23, fig. 9 (nest
architecture, floral relationships).

evoluta Linsley and MacSwain. Wash., Idaho and Wyo., south to northeast. Calif., Nev. and
northern Ariz. (Grand Canyon). Pollen: Collects pollen primarily from microseridine
Compositae including Agoseris glauca, A. g. var. monticola, Microseris nutans and
locally obtains pollen from some of the crepidine Compositae such as Crepis
occidentalis, the onagraceous Camissonia including C. pallida, C. tanacetifolia, and the
introduced weed Taraxacum officinale; visits other flowers for nectar including
Ranunculus californicus.

Andrena (Diandrena) evoluta Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 33. 2.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 85-88, figs. 1, 23-24, 56, map 6
(redescription).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 40-41 (floral
relationships). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 88 (floral relationships).
foxii Cockerell. Calif. Pollen: Collects pollen exclusively from flowers of Camissonia, chiefly C.

campestris, but also from C. contorta, C. dentata parishii and C. pallida pallida when
C. campestris is seasonally or locally rare or absent; visits these and other flowers for
nectar including Coreopsis californica, Cryptantha, Hemizonia, Isomeris arborea,
Lupinus, Malacothrix.

Andrena foxii Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 188. 2, d.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 130 (subgeneric position).
—Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 67-69, figs. 1, 9-10, 56, 60, 71, 87, map 2
(redescription).

Biology: Linsley and MacSwain, 1956. Pan-Pacific Ent. 32: 120-121 (floral relationships).
—Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 75 (floral relationships).
—Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 69 (floral relationships). —Linsley, MacSwain,
Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 18-20, 48-49, fig. 7, pl. 1 (floral
relationships).

gnaphalii (Cockerell). South. Calif. (Catalina Island and Transverse and Peninsular Ranges).
Pollen: Collects pollen apparently only from early morning opening flowers of A goseris
including A. heterophylla, A. retrorsa, but visits these and other flowers for nectar
including Anisocoma acaulis, Ceanothus integerrimus, Descourainia sophia,
Gnaphalium bicolor, Lasthenia chrysostoma, Potentilla, Sisymbrium, Sonchus

oleraceus.
Diandrena gnaphalii Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 148. d.

1792 Hymenoptera in America North of Mexico

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 91-93, figs. 1, 27-28, 56, map. 13
(redescription).
Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 92-93 (floral relationships).
lewisorum Thorp. Calif. Pollen: Collects pollen from flowers of the genus Clarkia including C.
breweri, C. concinna, C. cylindrica, C. douglasiana, C. dudleyana, C. imbricata, C.
purpurea, C. speciosa, C. unguiculata, C. williamsonii, C. xantiana, but visits these and
other flowers for nectar including Baccharis, Cryptantha flaccida, Datura, Eriogonum
fasciculatum, Hemizonia kelloggii, Lonicera interrupta, Mimulus, Ranunculus
occidentalis, Rhamnus californica.
Andrena (Diandrena) lewisorum Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 72, figs. 1, 13-14,
56, 61, 73, map 3. 9, d.

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 73-74 (floral relationships). —MacSwain,
Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 10-13, fig. 1 (floral relationships).
macswaini Linsley. Calif. (San Joaquin Valley). Pollen: Collects pollen from the flowers of
Camissonia campestris campestris and C. sierrae sierrae, but visits these and other
flowers for nectar including Camissonia boothii decorticans, Erodium, Isomeris
arborea, Malacothrix californica.
Andrena (Diandrena) macswaini Linsley, 1960. Pan-Pacifie Ent. 36: 97. 2, 6.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 36 (tax characters). —Thorp,
1969. Calif. Univ. Pubs. Ent. 52: 102-105, figs. 1, 37-38, 56, 66, 78, map 10 (redescription).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 105 (floral relationships). —Linsley,
MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 24-26, 50, pl. 2, fig. 11 (nest
architecture, floral relationships).

malacothricidis Thorp. South. Calif.; Mexico (Baja California). Pollen: Collects pollen from
stephanomerine Compositae, primarily Malacothrix including M. californica, M.
glabrata, but visits other flowers including Anisocoma acaulis, Calochortus,
Camissonia campestris campestris, Convolvulus malacophyllus, Cryptantha,
Descourainia sophia, Euphorbia albomarginata, Lasthenia chrysostoma, Linanthus
aureus, Phacelia ciliata.

Andrena (Diandrena) malacothricidis Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 113, figs. 1,
47-48, 56, map 11. 2, ¢.
Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 115-116 (floral relationships).

nothocalaidis (Cockerell). B. C. to north. Calif., east to Wyo. and Colo. Pollen: Collects pollen
from flowers of ligulate Compositae, primarily Microseridinae including Agoseris glauca,
Microseris nutans, Nothocalais cuspidata, Taraxacum officinale, but visits other
flowers for nectar including Antennaria, Camissonia tanacetifolia tanacetifolia, Crepis

occidentalis, Erigeron, Lepidium, Mimulus.
Diandrena nothocalaidis Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 183. 9, 6.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 33-35 (tax. characters and
status). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 105-107, figs. 1, 39-40, 56, map 11
(redescription).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 107 (floral relationships).
olivacea Viereck. Ariz. to south. Nev. and south. Calif. Pollen: Collects pollen almost

exclusively from the flowers of stephanomerine Compositae, primarily Anisocoma
acaulis and Malacothrix including M. californica, M. glabrata, but visits other flowers
including Agoseris heterophylla, Astragalus, Baileya, Calycoseris wrightti, Camissonia
campestris, Coreopsis bigelovii, C. californica, Cryptantha barbigera, Encelia farinosa,
Fallugia paradowa, Geraea canescens, Hemizonia, Hyptis emoryi, Layia glandulosa,
Phacelia, Rafinesquia neomexicana. Predator: Lestomyia sp.

Andrena (Parandrena) olivacea Viereck, 1916. Acad. Nat. Sci. Phila., Proce. 68: 590. 2, 6.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 35 (tax. characters). —Thorp,
1969. Calif. Univ. Pubs. Ent. 52: 88-91, figs. 1, 25-26, 56, 64, 76, map 7 (redescription).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 91 (floral relationships).

Superfamily APOIDEA 1793

parachalybea Viereck. South. Calif.; Mexico (Baja California). Pollen: Collects pollen from

Camissonia ineluding C. bistorta, C. campestris campestris, C. cheiranthifolia
suffruticosa, C. dentata, C. pallida pallida, but visits these and other flowers for nectar
including Agoseris heterophylla, Astragalus, Baccharis viminea, Ceanothus crassifolius,
Coreopsis bigelovii, C. californica, Cryptantha intermedia, Descourainia sophia,
Erodium cicutarium, Haplopappus linearifolius, Eriophyllum confertiflorum,
Lasthenia, Layia glandulosa, L. platyglossa campestris, Lomatium dasycarpum,
Monolopia lanceolata, Plagiobothrys californicus, Rhus trilobata, Salix laevigata, S.
lasiolepis, Sisymbrium irio, Tamarix.

Andrena (Parandrena) parachalybea Viereck, 1917. Amer. Ent. Soc., Trans. 43: 391. 9.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 61-63, figs. 1, 5-6, 56, map 1
(redescription).

Biology: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 74-75 (floral
relationships). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71:
17-18, 48, fig. 6 (floral relationships).

puthua (Cockerell). Calif.; Mexico (Baja California). Parasite: Stylops sp. Pollen: Collects pollen

from Lasthenia including L. aristata, L. chrysostoma, L. coronaria, L. debilis, L.
gracilis, L. minor, L. tenella, but visits these and other flowers for nectar including
Calandrinia ciliata, Chaenactis glabriuscula, Cryptantha, Eschscholzia californica,
Layia platyglossa, Rhamnus crocea.

Diandrena puthua Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 24. d (2 misdet.).

Diandrena beatula Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 48. 9.

Diandrena clariventris Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 50. 3.

Diandrena sanctorum Cockerell, 1941. San Diego Soc. Nat. Hist., Trans. 9: 346. 9.

Taxonomy: Timberlake, 1951. Jn Linsley Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1078 (synonymy). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 74-78,
figs. 1, 15-16, 56, map 4 (redescription, synonymy).

Biology: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 34, 77 (floral relationships, parasite).

sperryi (Cockerell). Calif. Pollen: Collects pollen from the flowers of Camissonia including C.
campestris, C. clavaeformis, C. pallida pallida, but visits these and other flowers for
nectar including Anisocoma acaulis, Coreopsis californica, C. bigelovii, Malacothrix

glabrata. Predator: Lestomyia sp.
Diandrena sperryi Cockerell, 1937. Amer. Mus. Novitates 948: 14. 2.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 69-72, figs. 1, 11-12, 56, 72, map 3
(redescription).

Biology: Linsley, MacSwain and Thorp, 1964. Calif. Univ. Pubs. Ent. 33: 74, table 3 (floral
relationships). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 33, 72 (floral relationships,
predator). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 20, 49,
fig. 8 (floral relationships).

subapasta Thorp. Calif. Pollen: Apparently collects pollen primarily from Arenaria californica,
but visits other flowers including Lasthenia, Orthocarpus erianthus.

Andrena (Diandrena) subapasta Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 120, figs. 1, 53-54,

56, map 13. 2, 6.

subchalybea Viereck. Calif.; Mexico (Baja California). Parasite: Dalmannia picta Will. Pollen:
Collects pollen primarily from ligulate Compositae, especially Agoseris heterophylla;
visits flowers of Amsinckia, Brassica, Camissonia campestris campestris, Coreopsis
californica, Crepis vesicaria, Descourainia sophia, Erodium, Euphorbia albomarginata,
Haplopappus Linearifolius, Hesperochiron californicus, Lasthenia chrysostoma, Layia
chrysanthemoides, Layia platyglossa, Limnanthes douglasii, Madia gracilis,
Malacothrix, Microseris douglasii, M. linearifolia, Mimulus, Monolopia lanceolata,
Phacelia ciliata, Platystemon californicus, Ranunculus californicus, Sisymbrium,

Sonchus oleraceus.
Andrena (Parandrena) subchalybea Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 5938. 9.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 32 (tax. status). —Thorp, 1969.
Calif. Univ. Pubs. Ent. 52: 111-113, figs. 1, 45-46, 56, 67, 79, 83-84, map 12 (redescription).

1794 Hymenoptera in America North of Mexico

Biology: Bohart, 1938. Pan-Pacific Ent. 14: 132 (parasite). —Thorp, 1969. Calif. Univ. Pubs.
Ent. 52: 34, 113 (floral relationships, parasite).
submoesta Viereck. Calif. Parasite: Stylops sp. Pollen: Collects pollen from the flowers of

Lasthenia including L. chrysostoma, L. coronaria, L. gracilis, L. minor, L. tenella, but
visits these and other flowers for nectar including Haplopappus, Hypochoeris radicata,
Layia, Monolopia lanceolata, Tamarix.

Andrena (Parandrena) submoesta Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 594. 9.

Diandrena marinensis Cockerell, 1936. Pan-Pacifie Ent. 12: 153. ¢.

Taxonomy: Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1082 (synonymy). —Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 78-81,
figs. 1, 17-18, 56, 62, 74, map 5 (redescription).

Biology: Thorp, 1969. Calif Univ. Pubs. Ent. 52: 34, 79-80 (floral relationships, parasite).
Genus ANDRENA Subgenus EREMANDRENA LaBerge

Andrena subg. Eremandrena LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 295.
Type-species: Pterandrena pallidiscopa Viereck. Monotypic and orig. desig.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 283, 293, 295-296 (tax. characters).
pallidiscopa pallidiscopa (Viereck). Calif., Oreg., Colo. Parasite: Myopa rubida (Bigot). Pollen:
Unknown, but visits flowers of Lomatium, Sanicula.
Pterandrena pallidiscopa Viereck, 1904. Canad. Ent. 36: 227. °.
Pterandrena nudimedicornis Viereck, 1904. Canad. Ent. 36: 227. o.

Taxonomy: Timberlake and Cockerell, 1933. Pan-Pacific Ent. 9: 28 (tax. characters).

Biology: Bohart, 1941. Pan-Pacific Ent. 17: 95 (parasite).
pallidiscopa trifasciata Timberlake and Cockerell. South. Calif. Parasite: Myopa rubida
(Bigot). Pollen: Unknown, but visits flowers of Calandrinia menziesii, Lasthenia
chrysostoma, Lomatium dasycarpum, L. utriculatum, Rhus trilobata, Sanicula
bipinnatifida.
Andrena pallidiscopa trifasciata Timberlake and Cockerell, 1933. Pan-Pacific Ent. 9: 28.
2.

Genus ANDRENA Subgenus EUANDRENA Hedicke

Andrena subg. Euandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 212.
Type-species: Andrena bicolor Fabricius. Orig. desig.

Andrena subg. Xanthandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 218.
Type-species: Andrena auricoma Smith. Orig. desig.

Andrena subg. Geandrena LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 313.
Type-species: Andrena caerulea Smith. Orig. desig.

Revision: LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101: 371-446, 74 figs. (N. Amer.
Spp.).

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 309 (tax. characters, as
Xanthandrena). —LaBerge, 1977. Amer. Ent. Soc., Trans. 103: 107-135, figs. (revised key
to included spp., synonymy; received too late for inclusion in this catalog).

algida Smith. N. S. to Alta. and N. W. T., south to N. J., N. Y., Mich., Minn., S. Dak., N. Mex.

and Ariz. Pollen: Apparently polylectic with some preference for the flowers of Salix
including S. brachycarpae, but visits other flowers including Cornus mas, Fragaria,
Pulsatilla hirsutissima, Ranunculus testiculatus, Ribes vallicola, Taraxacum
officinale, Thlaspi arvense.

Andrena algida Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 116. 2, d.

Andrena fragiliformis Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 435. 9.

Andrena fernaldiella Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 34. 3.

Andrena brachycarpae Viereck and Cockerell, 1914. U. S. Natl. Mus., Proe. 48: 55. ?.

Andrena (Andrena) fuscisignata Viereck, 1917. Amer. Ent. Soc., Trans. 45: 379. 2, d.

Andrena (Andrena) albisigna Viereck, 1922. Boston Soe. Nat. Hist., Occas. Papers 5: 38.
Oe

Superfamily APOIDEA 1795

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 150 (type). —Cockerell, 1906. Psyche
13: 7. 2 (type). —Atwood, 1934. Canad. Jour. Res. 10: 207, 209, figs. 2, d. —Mitchell, 1960.
N. C. Agr. Expt. Sta. Tech. Bul. 141: 218-219, figs. 42-44 (redescription). —LaBerge and
Ribble, 1975. Amer. Ent. Soc., Trans. 101: 396-399, figs. 3, 25-29 (redescription, synonymy).

astragali Viereck and Cockerell. Wash. to Calif. east to Utah, Wyo. and Nebr. Pollen:

Unknown, although frequently collected from the flowers of Zygadenus fremontii, but
visits other flowers including Agoseris glauca, Alliwm haematochiton, Astragalus,
Cardaria draba var. repens, Chamaebatia foliolosa, Cornus, Eriogonum, Oxytropis,
Salix, Taraxacum officinale, Tofieldia occidentalis.

Andrena astragali Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 46. °.

Andrena zygadeni Cockerell, 1932. Pan-Pacifie Ent. 8: 174. 9, 6.

auricoma Smith. B. C. to south. Calif., east to Wyo., Colo. and south to Ariz. Pollen: Polylectic,

most frequently collected at the flowers of Ranunculus, Descowrainia, Salix and
Potentilla in that order, but visits a great variety of flowers including Baccharis,
Barbarea orthoceras, Brassica nigra, Camissonia, Ceanothus, Chamaebatia foliolosa,
Cornus californicus, Crataegus, Cryptantha intermedia, C. muricata, Daucus carota,
Dendromecon rigida, Descourainia sophia, Eriodictyon californicum, Eriogonum
marifolium, Eriophyllum obovatum, Euphorbia albomarginata, Fragaria californica,
Fritillaria lanceolata, Geranium, Gilia capitata, Hackelia floribunda, H. patens,
Lasthenia chrysostoma, Layia, Linanthus montanus, Lomatium dissectum, L.
triternatum, Lupinus bicolor, Malacothamnus arcuatus, Malus, Mentzelia, Mirabilis
laevis, Montia perfoliata, Nemophila integrifolia, Orthocarpus erianthus, Phacelia
cicutaria, P. distans, P. hispida, P. ramosissima, P. tanacetifolia, Plagiobothrys
californicus, P. tenellus, Potentilla glandulosa, Prunus ilicifolia, P. melanocarpa, P.
subcordata, Pyracantha, Quercus agrifolia, Ranunculus californicus, R. occidentalis,
Rhamnus crocea, Rhus diversiloba, R. trilobata, Rubus ursinus, Salix laevigata, S.
lasiolepis, Salvia columbariae, Sanicula crassicaula, S. nevadensis, Saxifraga
integrifolia, Scrophularia californica, Sedum, Sisymbrium irio, Sisyrinchium bellum,
Stachys, Stenotopsis linearifolius, Tamarix gallica, Taraxacum officinale,
Thysanocarpus curvipes.

Andrena auricoma Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 56. ¢.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 155. d (notes on type). —Cockerell,
1906. Psyche 13: 8 (notes on type). —Viereck, 1904. Canad. Ent. 36: 193. 2, 3 (key).
—Lanham, 1941. Ent. Soc. Amer., Ann. 34: 706, 708. 2, ¢ (key).

caerulea Smith. B. C. (Vancouver Is.) to Calif., east to Idaho, Nev., and Utah (Mendon).

Parasite: Myopa rubida (Bigot), Myopa sp., Nomada obscurella Fowler, N. opacella
Timberlake, Stylops pacifica Bohart. Pollen: Oligolege of Ranunculus, especially R.
californicus, but visits other species of Ranunculus including R. alsimaefolius, R.
occidentalis, R. 0. dissectus, R. orthorhynchus as well as the flowers of Arctostaphylos,
Astragalus, Brassica, Ceanothus including C. cuneatus, Cryptantha, Fagus, Gilia,
Ipomoea, Lasthenia chrysostoma, Lomatium, Lysichiton kamtschatcense, Mimulus,
Nemophila, Phacelia, Polygonum bistortoides, Rhamnus, Ribes, Rubus, Salix, Senecio,
Sanicula nevadensis, Sisyrinchium bellum, Taraxacum officinale. Predator: Xysticus.

Andrena caerulea Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 55. °.

Andrena coerulea var. territa Cockerell, 1898. Entomologist 31: 89. °.

Pterandrena erigenoides Viereck. 1904. Canad. Ent. 36: 227. °.

Pterandrena crypta Viereck, 1904. Canad. Ent. 36: 227. °.

Pterandrena acrypta Viereck, 1904. Canad. Ent. 36: 227. °.

Pterandrena complexa Viereck, 1904. Canad. Ent. 36: 227. °.

Andrena (Ptilandrena) francisca Viereck, 1917. Acad. Nat. Sci. Phila., Proc.'68: 595. °.

Andrena supervirens Cockerell, 1924. Pan-Pacific Ent. 1: 64. 2.

Andrena supervirens var. aurescens Cockerell, 1924. Pan-Pacific Ent. 1: 64. 2.

Andrena (Andrena) innominata Viereck, 1926. Calif. Acad. Sci., Proc. (4) 15: 404. 6.

Andrena (Ptilandrena) tristis Linsley, 1951. In Muesebeck, Krombein and Townes, U. S.
Dept. Agr., Agr. Monog. 2: 1062. Nomen nudum.

1796 Hymenoptera in America North of Mexico

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 154 (type). —Cockerell, 1906. Psyche
13: 9 (type). —Viereck, 1904. Canad. Ent. 36: 227. ? (key, as territa). —Cockerell, 1932.
Canad. Ent. 64: 287. 2 (as complexa). —Michener, 1953. Kans. Univ. Sci. Bul. 35: 1034,
figs. 82, 83, 85 (larva, as complexa). —LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans.
101: 406-415, figs. 6, 35-39 (redescription, synonymy).

Biology: Bohart, 1936. Pan-Pacific Ent. 12: 9 (parasite). —Bohart, 1941. Pan-Pacific Ent. 17:
95 (parasite). —Bohart, 1941. Calif. Univ. Pubs. Ent. 7: 91 (parasite). —MacSwain and
Bohart, 1947. Pan-Pacifie Ent. 23: 30 (parasite). —MacSwain, 1949. Pan-Pacifie Ent. 25:
89-90 (parasite). —Linsley and MacSwain, 1955. Wasmann Jour. Biol. 13: 253 (parasite).
—Linsley and MacSwain, 1957. Calif. Univ. Pubs. Ent. 11: 395 (parasite). —Linsley and
MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 5-13, text figs. 1-5, plate figs. 1b, 3e and d
(nest construction, life history, overwintering, emergence, behavior, sex ratio, mating,
floral relationships, parasites).

chlorura Cockerell. Wash. to Calif., east to Utah and Airz.; Mexico (Baja California). Pollen:

Polylectic, visits a wide variety of flowers including Acer macrophyllum, Arbutus
menziesii, Arctostaphylos bicolor, A. crustacea, A. drupacea, A. glandulosa, A. glauca,
A. mariposa, A. patula, Berberis pinnata, Ceanothus arboreus, C. cordulatus, C.
crassifolius, C. cuneatus, C. greggii, C. integerrimus, C. leucodermis, C. palmeri, C.
soredatus, C. velutinus, C. verrucosa, Cercocarpus betulifolius, Claytonia spathulata,
Cryptantha, Dentaria californica, Fragaria californica, Gilia, Lasthenia chrysostoma,
Malus, Paeonia brownii, Petasites palmata, Prunus subcordata, Quercus, Ranunculus,
Rhamnus californica, R. crocea, R. ilicifolia, Rhus integrifolia, R. ovata, R. trilobata,
Ribes indecorum, Salix lasiolepis, Salvia mellifera, Tamarix.

Andrena chlorura Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 48. °.

Andrena (Andrena) complicata Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 554. 2.

Andrena ablusula Cockerell, 1936. Pan-Pacifie Ent. 12: 135. 9.

Andrena clementina Timberlake, 1941. South. Calif. Acad. Sci., Bul. 39: 193. 2.

Taxonomy: Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1061 (synonymy). —LaBerge and Ribble, 1975. Amer. Ent. Soc.,
Trans. 101: 402-406, figs. 5, 30-34 (redescription, synonymy).

dissimulans Timberlake. Calif. Pollen: Evidently an oligolege of Lasthenia including L.
aristata, L. chrysostoma, L. gracilis, L. tenella, but also visits the flowers of
Calandrinia menziesii, Salvia.

Andrena (Thysandrena) dissimulans Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 408. 2,
3

Anarene (Thysandrena) blandula Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 410. °.

Taxonomy: LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101: 426-429, figs. 3, 55-59
(redescription, synonymy).
hamulata LaBerge and Ribble. Calif. Pollen: Unknown, but visits flowers of Salix.
Andrena (Euandrena) hamulata LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101:
399, figs. 3, 20-24. 2, 6.
lawrencei Viereck and Cockerell. B. C. to Calif., east to Alta., Idaho, Wyo., Utah and Nebr.
Pollen: Unknown, but visits flowers of Balsamorhiza sagittata, Carnaria draba,
Helianthus nuttallii, Phlox hoodii, Silphium, Wyethia mollis.
Andrena lawrencei Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 15. °.
Andrena (Iomelissa) extensa Viereck, 1924. Canad. Ent. 56: 240. 9, d.

Taxonomy: Cockerell, 1932. Canad. Ent. 64: 286. ? (key, as extensa). —LaBerge and Ribble,
1975. Amer. Ent. Soc., Trans. 101: 386-390, figs. 3, 18-19 (redescription, synonymy).
misella Timberlake. Calif. Pollen: Unknown, but visits flowers of Amsinckia eastwoodae,
Anagallis arvensis, Brassica, Calandrinia ciliata, Lasthenia, Montia perfoliata,
Phacelia, Ranunculus.
Andrena misella Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 410. @.
nigrihirta (Ashmead). Alaska and transcont. south. Canada, south to Calif., Nev., Utah, Colo.,
N. Dak., Minn., Wis., Ill., Mich., N. Y. and Conn. Pollen: Apparently polylectic, visits
flowers of Angelica, Dentaria californica, Eriogonum, Eriophyllum confertiflorum,

Superfamily APOIDEA 1797

Erythronium grandiflorum, Fragaria, Fritillaria pumila, Lonicera, Lotus, Montia
perfoliata, Petasites palmata, Pulsatilla hirsutissima, Ranunculus, Rhododendron
occidentale, Rosa, Rubus, Smilacina stellata, Synthiridis plantagium.

Cillissa nigrihirta Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 6. d.

Andrena longihirtiscopa Viereck, 1904. Canad. Ent. 36: 191. 9.

Andrena decussata Viereck, 1904. Canad. Ent. 36: 1938, 194. @, d.

Andrena decussatula Viereck, 1904. Canad. Ent. 36: 193. 2.

Andrena synthiridis Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 436. 9.

Andrena novae-angliae Viereck, 1907. Ent. News 18: 283. 9.

Andrena nivaloides Graenicher, 1911. Pub. Mus. Milwaukee, Bul. 1: 235. 9°.

Andrena discolor Viereck, 1916. Amer. Mus. Nat. Hist., Bul. 35: 370. 9.

Andrena (Andrena) tumida Viereck, 1922. Boston Soc. Nat. Hist., Occas. Papers 5: 35. 2,
3.

Andrena (Andrena) vancouverensis Viereck, 1924. Canad. Ent. 56: 80. d.

Andrena (Andrena) nigrovaria Viereck, 1924. Canad. Ent. 56: 237. °.

Andrena (Andrena) marina Viereck, 1926. Calif. Acad. Sci., Proc. (4) 15: 405. 3.

Andrena ripariella Cockerell, 1936. Pan-Pacifie Ent. 12: 148. ¢.

Andrena (Thysandrena) crenata Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 221,
figs. 44, 51. 6.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 207. 2 (key, as novaeangliae). —Lanham,
1941. Ent. Soc. Amer., Ann. 34: 705. 2, 6 (key). —LaBerge and Ribble, 1975. Amer. Ent.
Soc., Trans. 101: 380-386, figs. 2, 8-12 (redescription, synonymy).

nigrocaerulea Cockerell. B. C. to Calif., east to Wyo. and Colo. Parasite: Stylops pacifica

Bohart. Pollen: Polylectic, visits a wide variety of flowers including Allium, Arbutus
menziesii, Arctostaphylos nevadensis, Arenaria kingti, Barbarea orthoceras, B. vulgaris,
Berberis aquilifolium, Brassica, Cakile edentula californica, Calyptridium umbellatum,
Ceanothus integerrimus, Cercocarpus betuloides, Chamaebatia foliolosa, Cryptantha
intermedia, Daucus carota, Descourainia sophia, Doronicum, Eriodictyon californicum,
Eriogonum lobbii, E. marifolium, Erodium botrys, Erysimum asperum, Fragaria
californica, Gilia capitata, G. multicaulis, G. tricolor, Hackelia patens, Hesperochiron
californicus, Horkelia tillingi, Lappula, Linanthus androsaceus, L. parviflorus,
Lithophragma affinis, Lomatium dissectum, Lonicera, Lotus, Mertensia, Microseris
nutans, Mimulus guttatus, Montia perfoliata, M. sibirica, Nemophila integrifolia, N.
spatulata, Oenothera, Penstemon cyananthus, Phacelia humilis, P. linearis, P. pusilla,
P. tanacetifolia, Plectritis macrocera, Potentilla glandulosa, Prunus virginiana, P.
subcordata, Pyracantha, Quercus agrifolia, Ranunculus californicus, Raphanus sativa,
Rhamnus californicus, R. crocea, Rubus, Salix, Sanicula nevadensis, Senecio canus, S.
integerrimus, Sisyrinchium bellum, Spraguea umbellata, Taraxacum officinale, T.
vulgare, Trifolium repens.

Andrena nigrocaerulea Cockerell, 1897. Entomologist 30: 309. 2, d.

Andrena seattlensis Viereck, 1904. Canad. Ent. 36: 191, 195. 9, 6.

Andrena epileuca Cockerell, 1924. Pan-Pacific Ent. 1: 62. 9, d.

Andrena lustrans Cockerell, 1924. Pan-Pacific Ent. 1: 638. 2.

Taxonomy: Cockerell, 1932. Canad. Ent. 64: 286 (key). —Cockerell, 1932. Canad. Ent. 64: 286.
2 (variation, as lwstrans). —Cockerell, 1932. Canad. Ent. 64: 286. 2 (key, as epileuca).
—LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101: 390-396, figs. 4, 13-17
(redescription, synonymy).

Biology: Linsley and MacSwain, 1957. Calif. Univ. Pubs. Ent. 11: 399 (stylopization).
—Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 19 (floral relationships,
stylopization).

penemisella LaBerge and Ribble. Calif. and Oreg. (Cayuse and The Dalles). Pollen: Unknown,
but visits flowers of Calandrinia, Cryptantha, Erodium, Montia perfoliata.
Andrena (Euandrena) penemisella LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101:
LMC
suavis Timberlake. Oreg. (Klamath Co.), Calif. Parasite: Nomada obliquella Timberlake, N.
opacellus Timberlake, Stylops pacifica Bohart. Pollen: Oligolectic, apparently collects

1798 Hymenoptera in America North of Mexico

pollen only from the flowers of Ranunculus californicus, but visits other flowers
presumably for nectar including Astragalus, Calandrinia menziesii, Ceanothus,
Lasthenia chrysostoma, Lomatium utriculatum, Montia perfoliata, Oenothera ovata,
Orthocarpus densiflorus, Platystemon californicus, Rhus trilobata, Wyethia
angustifolia. Predator: Xysticus sp.

Andrena (Ptilandrena) suavis Timberlake, 1938. Pan-Pacific Ent. 14: 24. 2, 6.

Biology: Bohart, 1941. Calif. Univ. Pubs. Ent. 7: 128 (parasite). —MacSwain, 1949.
Pan-Pacifie Ent. 25: 89 (stylopization). —Linsley and MacSwain, 1955. Wasmann Jour. Biol.
13: 253 (parasite). —Linsley and MacSwain, 1957. Calif. Univ. Pubs. Ent. 11: 395-422
(parasite, stylopization). —Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 13-16,
plate 3, figs. a and b (nest architecture, life history, overwintering, emergence, sex ratio,
floral relationships, parasites, predator).
subdepressa Timberlake. Calif. Pollen: Unknown, but visits flowers of Brassica, Dentaria
californica, Ranunculus californicus, Salix lasiolepis.
Andrena (Thysandrena) subdepressa Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 407. 2,
é.
Biology: Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 19 (floral relationships).
venata LaBerge and Ribble. Calif. (Sonora Pass). Pollen: Unknown, but visits flowers of

Ranunculus.
Andrena (Euandrena) venata LaBerge and Ribble, 1975. Amer. Ent. Soc., Trans. 101: 429,

figs. 60-64. 2, d.
Genus ANDRENA Subgenus GEISSANDRENA LaBerge

Andrena subg. Geissandrena LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 302.
Type-species: Andrena trevoris Cockerell. Monotypic and orig. desig.

Revision: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 302-307, figs. 1, 36-40.
trevoris Cockerell. B. C. and Alta., south to Calif., south to Idaho, Utah and ?Ariz. (Huachuca
Mts. and Nogales). Pollen: Unknown, but visits flowers of Aesculus, Cirsium, Lonicera,
Rosa, Symphoricarpos.
Andrena trevoris Cockerell, 1897. Entomologist 30: 306. d.
Andrena semipolita Viereck, 1904. Canad. Ent. 36: 192. °.
Andrena ricardonis Cockerell, 1916. Canad. Ent. 48: 272. 6.

Taxonomy: Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 380. 2.
Genus ANDRENA Subgenus GONANDRENA Viereck

Andrena subg. Gonandrena Viereck, 1917. Amer. Ent. Soc., Trans. 43: 390.
Type-species: Andrena (Gonandrena) persimulata Viereck. Monotypic.
Andrena subg. Tropandrena Viereck, 1924. Canad. Ent. 56: 21.
Type-species: Andrena fragilis Smith. Monotypic and orig. desig.
Revision: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 271-302, figs. 1-4, 10-35 (N.
Amer. spp.).
Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 210 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 302 (tax. characters).
avulsa LaBerge and Ribble. Tex. Pollen: Unknown, but visits flowers of Parkinsonia,
Phacelia.
Andrena (Gonandrena) avulsa LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 298,
figs. 4, 26-30. 2, d.
flocculosa LaBerge and Ribble. Wash., Calif. (Gold Lake and Meadow Valley), Nev. (Lamoille).
Andrena (Gonandrena) flocculosa LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98:
296, figs. 4, 24-25. 2, 6.
fragilis Smith. Que. to Ga., west to Wis., Iowa and Kans. Pollen: Apparently an oligolege of
Cornus including C. amonum, C. paniculata, but visits other flowers including Aruncus,

Superfamily APOIDEA 1799

Castanea pumila, Ceanothus, Cicuta maculata, Cirsium, Hydrangea, Pastinaca sativa,
Ptelea, Salix, Svida, Trifoliwm, Viburnum dentatum, V. molle.

Andrena fragilis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 115. ¢.

Andrena laticeps Provancher, 1888. Addit. Corr. Fauna Ent. Canada, Hym., p. 307. 6.
Preoce.

Andrena provancheri Dalla Torre, 1896. Cat. Hym. v. 10, p. 147. N. name.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 149 (type of fragilis). —Cockerell,
1906. Psyche 13: 7 (type of fragilis). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
231-2382, figs. 46-49, table 6 (redescription). —LaBerge and Ribble, 1972. Amer. Ent. Soc.,
Trans. 98: 278-284, figs. 2, 10-14 (redescription, synonymy).

integra Smith. Que. and Ont., south to N. C., west to Minn. and Kans. Pollen: Possibly an
oligolege of Cornus including C. stolonifera, but visits other flowers including Achillea,
Aruncus, Barbarea vulgare, Crataegus, Cryptotaenia, Melilotus alba, M. officinalis,
Pastinaca, Ptelea, Rhus, Viburnum, Vitis.
Andrena integra Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 114. 9.
Andrena lineata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 309. °.
Andrena (Gonandrena) lucifera Cockerell, 1932. Canad. Ent. 64: 155. °.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 124 (type of integra). —Cockerell,
1906. Psyche 13: 6 (type of integra). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
232-233, figs. 46-49, table 6 (redescription). —LaBerge and Ribble, 1972. Amer. Ent. Soc.,
Trans. 98: 284-287, figs. 2, 15-19 (redescription, synonymy).

nigrifrons (Cresson). Ont. and N. H. south to N. C., west to Man., Minn., Nebr., Kans. and Tex.

Pollen: Possibly polylectic, although seems to prefer pollen of Cornus including C.
asperifolia, C. paniculata, C. racemosa, C. sericea, C. stolonifera, but visits other
flowers including Achillea, Barbarea vulgaris, Ceanothus americanus, Cryptotaenia
canadensis, Erigeron philadelphus, Hieraceum, Lotus corniculatus, Melilotus alba, M.
officinalis, Ptelea trifoliata, Rhus glabra, Rubus, Sambucus americanus, Spiraea
aruncus, S. vanhouttei, Taraxacum, Toxicodendron, Viburnum dentatum, Zizia aurea.

Panurgus nigrifrons Cresson, 1878. Amer. Ent. Soc., Trans. 7: 62. ¢.

Andrena platyparia Robertson, 1895. Amer. Ent. Soc., Trans. 22: 119. 2, ¢.

Andrena (Andrena) barbarica Viereck, 1917. Amer. Ent. Soc., Trans. 43: 369. °.

Andrena (Gonandrena) monroensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
233. 2.

Taxonomy: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 287-292, figs. 3, 20-21
(redescription, synonymy).
persimulata Viereck. N.S. to N. W. T. and Alta., south to Conn., N. Y., Mich., Wis., Minn.,

Nebr., Colo. and Utah. Pollen: Possibly an oligolege of Cornus including C. asperifolia,
C. stolonifera, but visits other flowers including Angelica atropurpurea, Crataegus,
Deutzia gracilis, Fragaria, Lomatium dissectum, Polygonum convolvulus, Prunus,
Spiraea, Taraxacum officinalis, Viburnum.

Andrena (Gonandrena) persimulata Viereck, 1917. Amer. Ent. Soc., Trans. 43: 390. °.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 206, 209. 2, ¢ (key). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 235-236, figs. 45-46, 48, table 6 (redescription). —LaBerge
and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 292-296, figs. 4, 22-23 (redescription).

Genus ANDRENA Subgenus HESPERANDRENA Timberlake
Andrena subg. Hesperandrena Timberlake, 1949. Jn Lanham, Calif. Univ. Pubs. Ent. 8:
208.

Type-species: Andrena escondida Cockerell. Orig. desig.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 283, 293, 301 (tax. characters).
baeriae Timberlake. Calif. Pollen: Apparently an oligolege of the flowers of Lasthenia
including L. aristata, L. chrysostoma, L. gracilis, but visits other flowers including
Eriophyllum confertiflorum, Layia platyglossa, Lessingia germanorum, Senecio

californica.
Andrena baeriae Timberlake, 1941. South. Calif. Acad. Sci., Bul. 39: 194. 2, 6.

1800 Hymenoptera in America North of Mexico

duboisi Timberlake. North. Calif. Pollen: Unknown, but visits flowers of Lasthenia
chrysostoma, Layia platyglossa.
Andrena (Hesperandrena) duboisi Timberlake, 1951. U. S. Natl. Mus., Proce. 101: 390. 2, ¢.
escondida Cockerell, South. Calif.; Mexico (Baja California). Pollen: Unknown, but visits
flowers of Encelia, Layia platyglossa.
Andrena escondida Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 146. d.
lativentris Timberlake. Calif. Pollen: Unknown, but visits flowers of Amsinckia intermedia,
Coreopsis maritima, Hemizonia pungens, Lasthenia chrysostoma, L. gracilis, L. tenella,
Layia platyglossa.
Andrena (Hesperandrena) lativentris Timberlake, 1951. U.S. Natl. Mus., Proc. 101: 388. 2,
é.
limnanthis Timberlake. Calif. Pollen: Possibly an oligolege of Limnanthes including L.
douglasii, but visits other flowers including Chamaebatia foliolosa, Layia platyglossa,
Raphanus sativus.
Andrena (Hesperandrena) limnanthis Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 387.
oF ok

Genus ANDRENA Subgenus IOMELISSA Robertson

Iomelissa Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 50.
Type-species: Andrena violae Robertson. Monotypic.

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 207 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 284, 288, 300 (tax. characters).
violae Robertson. N. Y. to N. C., west to Ill. and Colo. Pollen: Obtains pollen solely from the
flowers of Viola, but visits other flowers for nectar including Cardamine, Ellisia,
Oxalis, Potentilla.
Andrena violae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 53. 2, 6.
Andrena davisi Viereck, 1907. Ent. News 18: 283. 2.

Taxonomy: Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 10. 2 (key). —Lanham, 1941.
Ent. Soc. Amer., Ann. 34: 706, 707. 2, 3 (key). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 252-254, figs. 54-56 (redescription, synonymy).

Genus ANDRENA Subgenus LARANDRENA LaBerge

Andrena subg. Larandrena LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 304.
Type-species: Andrena miserabilis Cresson. Orig. desig.

Revision: Ribble, 1967. Nebr. Univ. State Mus., Bul. 6: 27-42, figs. 1-5, 1 table, 1 map.

Taxonomy: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 308 (tax. status).

miserabilis Cresson. Transcont. south. Canada and U.S., south to Fla., Ala., Miss., Tex., N.
Mex., Utah and Calif.; Mexico (Tamaulipas). Parasite: Lewcophora obtusa (Zett.),
Nomada sp., Stylops bipunctata Pierce, S. oklahomae Pierce. Pollen: Polylectic,
principally obtains pollen from the flowers of Rosaceae and Salicaceae; visitation
records include Amelanchier, Antennaria, Arabis, Aronia, Berberis, Brassica, Capsella,
Cardamine, Ceanothus, Cercis, Cercocarpus, Claytonia, Comandra, Convolvulus,
Cornus, Crataegus, Cydonia, Dentaria, Erigenia, Exochorda, Fragaria, Hepatica,
Heracleum, Ilex, Isopyrum, Malus, Prunus, Ptelea, Pyracantha, Pyrus, Ranunculus,
Rhamnus, Rhus, Rubus, Salix, Solidago, Sorbaria, Spiraea, Stachys, Staphylea,
Stellaria, Taraxacum, Viburnum, Viola, Zanthoxylum.

Andrena clypeata Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 115. 2. Preoce.
Andrena miserabilis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 259. 2 (6 misdet.).
Andrena bipunctata Cresson, 1872. Amer. Ent. Soc., Trans. 4: 259. d.

Andrena flavoclypeata Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 54. d.
Andrena scutellaris Robertson, 1893. Amer. Ent. Soc., Trans. 10: 148. 2. Preoce.
Anthrena clypeolata Dalla Torre, 1896. Cat. Hym., v. 10, p. 133. N. name.
Anthrena scutellata Dalla Torre, 1896. Cat. Hym., v. 10, p. 151. N. name.

Andrena pennsylvanicola Viereck, 1907. Ent. News 18: 284. 9.

Andrena pronitens Cockerell, 1930. Ann. and Mag. Nat. Hist. (10) 5: 114. @.

Superfamily APOIDEA 1801

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 124, 152 (types of clypeata,
flavoclypeata). —Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193. 2, 5 (key).
—Cockerell, 1906. Psyche 13: 9, 36, (types of clypeata, flavoclypeata). —Atwood, 1934.
Canad. Jour. Res. 10 (2): 208, 209, fig. 2, ¢ (key). —Lanham and Timberlake, 1951. Jn
Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2: 1058 (synonymy).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 95, 160, fig. 29, table 4
(redescription). —Ribble, 1967. Nebr. Univ. State Mus., Bul. 6: 27-42, 5 figs., 1 table, 1 map
(redescription, synonymy).

Biology: Michener and Rettenmeyer, 1956. Kans. Univ. Sci. Bul. 37: 679-681, fig. 21 (nest
architecture, life history, parasites, as bipwnctata). —Ribble, 1967. Nebr. Univ. State
Mus., Bul. 6: 37, table 1 (floral relationships).

Genus ANDRENA Subgenus LEUCANDRENA Hedicke

Andrena subg. Leucandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 215.
Type-species: Apis sericea Christ. Orig. desig.

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 220 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 285, 290, 310 (tax. characters).
bifurcata Mitchell. Mich., Ohio.
Andrena (Leucandrena) bifurcata Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 211,
figs. 40-42. ¢.
chippewaensis Mitchell. Mich., N. B.
Andrena (?Leucandrena) chippewaensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
141: 212, fig. 43. 2.

Taxonomy: Knerer and Atwood, 1963. Canad. Ent. 95: 584, figs. 2, 4. d.
electrica Casad and Cockerell. N. Mex.
Andrena electrica Casad and Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 89. gd, @.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 49. 2, 3 (key).

erythronii Robertson. Que. to Mich., Ill. and Kans., south to Maine and Mass. Pollen: Primarily
collects pollen of Erythronium including E. mesochoreum, but also collects pollen from
the flowers of Malus, Prunus, Quercus, Taraxacum; visits other flowers presumably
for nectar including Amelanchier, Claytonia, Erigenia, Hepatica, Salix.
Andrena erythronti Robertson, 1891. Amer. Ent. Soc., Trans. 18: 53. 2, 6.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 193. 2, ¢ (key). —Michener,
1953. Kans. Univ. Sci. Bul. 35: 1035, figs. 88-91 (larva). —Mitchell, 1960. N. C. Agr. Expt.
Sta. Tech. Bul. 141: 212-213, figs. 40-42 (redescription). —Yager and Rozen, 1966. Amer.
Mus. Novitates 2265: 5 (pupa).

Biology: Michener and Rettenmeyer, 1956. Kans. Univ. Sci. Bul. 37: 645-679, figs. 1-20 (nest
architecture, life history, floral relationships, mating, number of females per nest,
immature stages).

lupini Cockerell. Calif. Pollen: Unknown, but visits flowers of Arctostaphylos glandulosa, A.
glauca, Ceanothus cuneatus, C. greggii, C. integerrimus, C. verrucosus, Clematis
lasiantha, Erodium botrys, Eschscholzia californica, Lasthenia chrysostoma, Lathyrus
alfeldii, Lupinus, Phacelia.

Andrena lupini Cockerell, 1936. Pan-Pacifie Ent. 12: 142. 9.

mariposorum Viereck. Calif.

Andrena (Andrena) mariposorum Viereck, 1917. Amer. Ent. Soc., Trans. 43: 382. 9.

parnassiae Cockerell. Mich., Wis., Vt. Pollen: Unknown, but visits flowers of Parnassia.
Andrena parnassiae Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 105. @.
Taxonomy: Graenicher, 1904. Ent. News 15: 66. 2, ¢. —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 213-214, figs. 40-43 (redescription).

perezana Viereck and Cockerell. Nebr.
Andrena perezana Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 47. °.

1802 Hymenoptera in America North of Mexico

picta Mitchell. N. Y. (Ithaca).
Andrena (Leucandrena) picta Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 214. °.
placida placida Smith. Alaska to N. S., south to Va., Ohio, Ill., Mich., Minn. and Calif. Parasite:
Meloe opacus LeC., M. strigulosus Mann., Stylops sp.?. Pollen: Polylectic, analyzed pollen
stores reveal that a given female, with one exception, provisions her cells with only one
kind of pollen which in California consisted of predominantly pollen of Eschscholzia
californica, while others stored pollen of a ligulate Compositae (Agoseris?), legume
(Lupinus?), and a crucifer (Raphanus?); visitation records for other flowers include
Acer, Amelanchier, Fragaria, Layia carnosa, Prunus, Pyrus malus, Taraxacum,
Viburnum.
Andrena placida Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 112. 9.
Andrena macilenta Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 313. 2, o.
Andrena macgillivrayi Cockerell, 1897. Entomologist 30: 308. °.
Andrena salicacea Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 48. 9.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 151 (type of placida). —Cockerell,
1906. Psyche 13: 7 (type of placida). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
214-216, figs. 40-42, table 6 (redescription). —Thorp and Stage, 1968. Ent. Soc. Amer., Ann.
61: 1583-1584, figs. 3-9. (immature stages, chromosome number).

Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology). —Thorp
and Stage, 1968. Ent. Soc. Amer., Ann. 61: 1580-1586, 9 figs., table (nest architecture, life
history, floral realtionships, parasites).

placida sapellonis Cockerell. Colo., N. Mex.
Andrena sapellonis Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 19. 9.
recta Mitchell. Minn. to N. Y. south to N. C. Pollen: Unknown, but visits flowers of Helianthus
annuus. Predator: Philanthus sp.
Andrena (Leucandrena) recta Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 217, fig.
43. 9.
seavillensis Mitchell. N. J. (South Seaville).
Andrena (?Leucandrena) seavillensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
217, fig. 43. 2.
trapezoidina Viereck and Cockerell. Nebr.
Andrena trapezoidina Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 58. @.

Genus ANDRENA Subgenus MELANDRENA Perez

Melandrena Perez, 1890. Soc. Linn. Bordeaux, Actes 44: 170.
Type-species: Andrena morio Brulle. Desig. by Hedicke, 1933.

Andrena subg. Gymnandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 213.
Type-species: Apis thoracica Fabricius. Orig. desig.

Andrena subg. Cryptandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 222. Preocc.
Type-species: Andrena carlini Cockerell. Orig. desig.

Andrena subg. Bythandrena Lanham, 1950. Ent. News 61: 140. N. name.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 310, 311 (tax. characters, as
Gymnandrena and Melandrena. —LaBerge, 1971. Pan-Pacific Ent. 47: 47 (synonymy).
bisignata Mitchell. Mass. to N. C., west to Minn. Pollen: Unknown, but visits flowers of
Erythronium americanum, Ilex verticillata, Prunus serotina, Pyrus malus, Salix
sericea, Viburnum acerifolium.
Andrena (Gymnandrena) bisignata Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
122; figs. 20, 22, 23. 2, d.
bryanti Cockerell. Alta.
Andrena bryanti Cockerell, 1938. Canad. Ent. 70: 5. 9.
carlini carlini Cockerell. N. S. to Ga., west to Minn., Colo., Wyo. and Oreg.?, B. C.? Pollen:
Apparently polylectic, visits flowers of Amelanchier, Anemonella, Arabis, Brassica,
Cercis, Chaerophyllum, Claytonia, Cornus, Dentaria, Dicentra, Erigenia, Erythronium,
Hepatica, Heracleum, Hydrophyllum, Isopyrum, Malus, Polemonium, Prunus, Pyrus,

Superfamily APOIDEA 1808

Rhus, Ribes, Rubus, Salix, Sanguinaria, Sassafras, Smilacina, Solidago, Thaspium,
Tragana, Trifolium, Uvularia, Vaccinium, Viburnum, Vicia.
Andrena carlini Cockerell, 1901. Canad. Ent. 23: 150, 153. 9.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 207, 209. 2, ¢ (key). —Lanham, 1941. Ent.
Soe. Amer., Ann. 34: 705, 708. 2, d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
141: 111-112, figs. 17, 19, table 3 (redescription).

Biology: Atwood, 1933. Canad. Jour. Res. 9: 456 (floral relationships). — Brittain, 1933. Canad.
Dept. Agr., Bul. 162: 94, figs. (floral relationships).
carlini neorhodura Mitchell. Mass., Pa., Mich.
Andrena (Bythandrena) carlini neorhodura Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 113. 6.

carliniformis Viereck and Cockerell. B. C. to Calif., east to Wyo., Colo. and Utah. Pollen:
Unknown, but visits flowers of Agastache urticifolia, Allium, Calochortus, Ceanothus,
Chamaebatia foliolosa, Eriogonum, Euphorbia crenulata, Gilia, Horkelia, Lappula,
Lotus nevadensis, Lupinus bicolor, Phacelia Plagiobothrys nothofulvus, Potentilla,
Prunus demissa, Ranunculus californicus, Rhamnus crocea, R. californica, Salix,
Salvia columbariae, Taraxacum officinale, Vicia, Wyethia.
Andrena carliniformis Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 25. ?.
carolinensis Mitchell. N. C. (Raleigh). Pollen: Unknown, but visits flowers of Vicia
caroliniana.
Andrena (Gymnandrena) carolinensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
123, fig. 20. 2.
cerasifolii Cockerell. Calif., Ariz., Utah, N. Mex. Pollen: Polylectic, visits flowers of
Arctostaphylos drupacea, A. glauca, A. mariposa, A. patula, Brassica arvensis, B.
campestris, Ceanothus cuneatus, C. greggii, C. palmeri, Cirsium californicum, Clarkia
biloba, C. nitens, C. unguiculata, Cryptantha intermedia, Emmenanthe penduliflora,
Eriogonum fasciculatum, Erodium, Erysimum asperum, Heteromeles arbutifolia,
Horkelia bolanderi, Hyptis emoryi, Penstemon spectabilis, Phacelia brachyloba, P.
cicutaria, Rhamnus californicus, R. crocea, Rhus trilobata, Ribes indecorum, Salix
hindsiana, S. lasiolepis, Sisymbrium irio, Swertia parryi, Trichostema parryi,
Viburnum.
Andrena cerasifolii Cockerell, 1896. Ent. Monthly Mag. (2) 7: 220. ¢.
Andrena mimetica Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 412. 2, 6.
Andrena mimetica var. falli Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 536.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48. ¢ (key). —Cockerell, 1924.
Pan-Pacifie Ent. 1: 58. 6. —Timberlake, 1951. Jn Muesebeck, Krombein and Townes, U. S.
Dept. Agr., Agr. Monog. 2: 1060 (synonymy).

Biology: Linsley, 1937. Brooklyn Ent. Soc., Bul. 32: 125 (floral relationships, number of
generations). —MacSwain, Raven, Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 46 (floral
relationships).

commoda Smith. Southeast. Canad, south to Ga., west to N. Dak. Pollen: Unknown, but visits
flowers of Cornus paniculata.
Andrena commoda Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 53. 2.
Andrena corni Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 50. °.
Andrena pyrura Cockerell, 1906. Ann. Mag. and Nat. Hist. (7) 17: 309. 9.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 152 (type). —Cockerell, 1906. Psyche
13: 6. 2 (type). —Viereck, 1907. Ent. News 18: 283, 286. 2, d (key). —Cockerell, 1931.
Amer. Mus. Novitates 458: 18, 19. —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
124, figs. 22-23, table 3 (redescription, synonymy).

confederata Viereck. Ohio to N. J., south to Fla. Pollen: Unknown, but visits flowers of
Amelanchier, Castanea, Crataegus, Fragaria, Ilex, Malus, Padus, Prunus, Pyracantha,
Rubus.
Andrena (Andrena) confederata Viereck, 1917. Amer. Ent. Soc., Trans. 43: 375. 2, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 125, figs. 22-23, table 3
(redescription).

1804 Hymenoptera in America North of Mexico

critica Mitchell. N. Y. (Catskill Mts.).
Andrena (Bythandrena) critica Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 113. °.
dunningi Cockerell. Que. to N. C., west to Minn. Pollen: Unknown, but visits flowers of
Claytonia, Gleditsia, Malus, Narcissus, Ribes, Rubus, Salix, Taraxacum, Viburnum.
Andrena dunningi Cockerell, 1898. Canad. Ent. 30: 103. °.
Andrena viciniformis Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 47. 2, 6.
Andrena annae Cockerell, 1931. Canad. Ent. 63: 200. °.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 192. 2, 6 (key). —Timberlake,
1951. In Linsley, In Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2:
1064 (synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 113-114, figs.
18-19, table 3 (redescription).

errans Smith. B. C. to Calif. and N. Mex. Pollen: Unknown, but visits flowers of

Arctostaphylos mariposa, Ceanothus, Eriogonum marifolium, Lappula, Ranunculus,
Salix.

Andrena errans Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 55. 2, d.

Andrena pluvialis Cockerell, 1901. Canad. Ent. 33: 154. 9.

Andrena junonia Viereck, 1904. Canad. Ent. 36: 191. 2.

Andrena compactiscopa Viereck, 1904. Canad. Ent. 36: 191. 2.

Andrena argentiniae var. trichomelaena Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 12:
SOsGrese.

Andrena spaldingi Cockerell, 1934. Amer. Mus. Novitates 697: 3. 2.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 153 (type). —Cockerell, 1903. Ann.
and Mag. Nat. Hist. (7) 12: 443. ¢ (as pluvialis). —Cockerell, 1906. Psyche 13: 34 (type).
—Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U.S. Dept. Agr.,
Agr. Monog. 2: 1065 (synonymy).

flexa Malloch. Ill. to Tex. Parasite: Nomada sp.
Andrena flexa Malloch, 1917. Brooklyn Ent. Soce., Bul. 12: 92. 2.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 126 (redescription).

Biology: Rozen, 1966. Amer. Mus. Novitates 2244: 26 (parasite).
gabrielsoni Mitchell. Maine, Conn., N. C., Mich.
Andrena (Bythandrena) gabrielsoni Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
115. @.
hilaris Smith. Mich. and Ohio to eastern Canada and New England States, south to Ga. and
Ala. Pollen: Unknown, but visits flowers of Aronia, Crataegus, Hydrangea, Ilex, Malus,
Prunus, Pyrus, Rubus, Salix, Vaccinium.
Andrena hilaris Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 112. @.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 151 (type). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 126-127, figs. 21-23, table 3 (redescription).
lupinorum helenae Cockerell. Colo.
Andrena lupinorum helenae Cockerell, 1934. Amer. Mus. Novitates 697: 2. 2.
lupinorum lupinorum Cockerell. Wyo., Colo., Nebr. Pollen: Unknown, but visits flowers of
Lupinus.
Andrena lupinorum Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 308. 9.
Andrena vicina argentiniae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 432. 9.
Taxonomy: Lanham, 1941. Ent. Soc. Amer., Ann. 34: 705. 2 (key). —Timberlake, 1951. In
Linsley, Jn Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr. Monog. 2: 1070
(synonymy).
macra Mitchell. N. C. Pollen: Unknown, but visits flowers of Crataegus, Pyracantha, Rubus.
Andrena macra Mitchell, 1951. Elisha Mitchell Sci. Soc., Jour. 67: 246, figs. 1-3. 2, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 127-128, figs. 22-23, table 3
(redescription).

Biology: Sivik, 1954. Ent. News 65: 253-255 (nest, supersedure).

Superfamily APOIDEA 1805

nigra Provancher. Ariz., south. Calif.; Mexico (Baja California). Pollen: Apparently an oligolege
of Phacelia including P. distans, but also visits flowers of Coreopsis maritima,
Cryptantha intermedia, Layia platyglossa, Malacothrix, Malus, Salix.
Andrena nigra Provancher, 1896. Nat. Canad. 23: 173. 9.
Andrena griseonigra Cockerell, 1905. Canad. Ent. 37: 371. 3.
Andrena subtristis Cockerell, 1905. Canad. Ent. 37: 372. 9.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 266, 279. 2, 3 (key).

Biology: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 280 (floral relationships).
nigripes Provancher. Calif.
Andrena nigripes Provancher, 1895. Nat. Canad. 22: 173. 2.
nivalis Smith. Que. to Ga., west to B. C., south to Calif. and N. Mex. Pollen: Unknown, but
visits flowers of Hydrangea, Rhododendron, Rubus, Vagnera.
Andrena nivalis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 118. 2, 6.
Andrena convexa Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 311. 9, ¢.
Andrena semirufa Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 407. °.
Andrena solidula Viereck, 1904. Canad. Ent. 36: 191, 194. 2, 6.
Andrena idahorum Viereck, 1916. Amer. Mus. Nat. Hist., Bul. 35: 732. ¢.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 151. 2 (type). —Cockerell, 1906.
Psyche 13: 9 (type). —Cockerell, 1931. Amer. Mus. Novitates 458: 14. 2 (key, as semirufa).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 128-129, figs. 22, 23, table 3
(redescription, synonymy).

obscuripennis Smith. N. J.(?), N. C., Ga., La.(?). Parasite: Nomada crudelis Cress.? Pollen:
Unknown, but visits flowers of Batodendron, Crataegus, Ilex, Padus.
Andrena obscuripennis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 188. °.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 150. 2 (type). —Cockerell, 1906.
Psyche 13: 36. ° (type). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 129-131,
figs. 21, 23, table 3 (redescription).

perimelas Cockerell. Coastal Calif. and Catalina Island Parasite: Nomada edwardsii edwardsii
Cress., Stylops pacifica Bohart, S. vandykei Bohart. Pollen: Apparently polylectic, visits
flowers of Brassica, Coreopsis maritima, Crepis, Limnanthes douglasii,
Mesembryanthemum edule, Prunus, Ranunculus californicus, Raphanus sativus, Ribes
cereum, Rubus, Salix, Sisymbrium bellum, Zygadenus fremontii.
Andrena perimelas Cockerell, 1905. Canad. Ent. 37: 371. °.
Andrena meadowsi Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 146. 6.

Biology: Linsley and MacSwain, 1955. Wasmann Jour. Biol. 13: 254-256, 270, pl. 1, fig. 5 (nest,
floral relationships, parasite). —Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 19,
pl. 5 (nest, floral relationships, parasite, stylopization).

pertristis Cockerell. Calif. Pollen: Unknown, but visits flowers of Brassica, Cryptantha
intermedia, Eriodictyon trichocalyx, Fremontodendron californica, Lupinus formosus,
Malus, Nemophila integrifolia, Phacelia heterophylla, Stanleya pinnata.
Andrena pertristis Cockerell, 1905. Canad. Ent. 37: 372. 2.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 265, 267. 2, d (key).
platyrhina Cockerell. Colo.
Andrena (Pterandrena) platyrhina Cockerell, 1930. N. Y. Ent. Soc., Jour. 37: 446. 9.
pruni Robertson. Minn. and III. to Mass., south to Ga. Pollen: Unknown, but visits flowers of
Aruncus, Claytonia, Crataegus, Dentaria, Heracleum, Pastinaca, Polemonium, Prunus,
Ribes, Rubus, Salix, Staphylea, Uvularia, Viburnum, Zanthoxylum.
Andrena pruni Robertson, 1891. Amer. Ent. Soc., Trans. 18: 51. 2, ¢.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191, 192. 2, ¢ (key). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 117-118, figs. 19-20, table 3 (redescription).

pulverulenta Viereck. Oreg.
Andrena pulverulenta Viereck, 1904. Canad. Ent. 36: 190, 195. °, d.

1806 Hymenoptera in America North of Mexico

regularis Malloch. B. C. to N. S., in eastern U.S. from Minn. to New England States, south to
Ga. Pollen: Unknown, but visits flowers of Aster, Brassica, Pyrus, Rubus, Syringa,

Vaccinium.
Andrena regularis Malloch, 1917. Brooklyn Ent. Soc., Bul. 12: 91. 6, °.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 207, 209. 2, d (key). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 118-119, figs. 18-19, table 3 (redescription).
sayi Robertson. N. B. to N. C., west to Minn., Ill. and Kansas. Pollen: Unknown, but visits
flowers of Amelanchier, Aronia, Brassica, Cercis, Crataegus, Malus, Padus, Prunus,
Pyrus, Rubus, Salix, Stellaria, Taraxacum, Vaccinium.
Andrena sayi Robertson, 1891. Amer. Ent. Soc., Trans. 18: 52. 2, 3.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191, 192. 2, 6 (key). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 131-182, fig. 23, table 3 (redescription).
sevierensis Mitchell. Tenn. (Sevier Co.).
Andrena (?Gymnandrena) seviensis(!) Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
182, fig. 20. 2.
Andrena (?Gymnandrena) sevierensis Krombein, 1967. Jn Krombein and Burks, U. S.
Dept. Agr., Agr. Monog. 2, Second Suppl. p. 434. Emend.
sola Viereck. B. C. to Calif., Ariz.; Mexico (Baja California). Pollen: Unknown, but visits
flowers of Amsinckia intermedia, Arctostaphylos, Baccharis, Brassica, Ceanothus
crassifolius, Chaenactis glabriuscula, Clematis lasiantha, Cryptantha intermedia,
Encelia farinosa, Eriodictyon, Gallium nuttallu, Gila, Layia platyglossa, Melilotus,
Papaver californica, Plagiobothrys californicus, Platystemon californica, Prunus
subcordatus, Quercus, Ranunculus, Rhamnus californica, R. crocea, Rhus diversiloba,
R. trilobata, Senecio californicus, Solanum umbelliferum, Tamaria.
Andrena (Andrena) sola Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 577. 2.
Andrena carissima Cockerell, 1924. Pan-Pacifiec Ent. 1: 58. 9.
Andrena microdonta Cockerell, 1924. Pan-Pacifie Ent. 1: 61. d.
Andrena ensenadensis Cockerell, 1941. San Diego Soc. Nat. Hist., Trans. 9: 345. @.

Taxonomy: Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1081 (synonymy).
transnigra Viereck. B. C. to Calif., Colo. and Wyo. Pollen: Unknown, but visits flowers of

Allium, Brassica, Ceanothus integerrimus, C. parviflorus, Chamaebatia foliolosa,
Lappula, Madia ramui, Ranunculus, Ribes, Salix, Taraxacum, Vaccinium, Wyethia.

Andrena transnigra Viereck, 1904. Canad. Ent. 36: 191. 9.

Andrena cyanura Cockerell, 1916. Canad. Ent. 48: 252. 9.

Andrena transnigra paysoni Cockerell, 1924. Ent. News 35: 349. 2.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 18. °.
vicina Smith. N. S. to Ga., west to B. C., Colo. and Oreg. Parasite: Nomada imbricata Sm., N.
obliterata Cress., N. vicina vicina Sm. Pollen: Unknown, but visits flowers of
Amelanchier, Azalea, Caragana, Castanea, Ilex, Malus, Narcissus, Padus, Prunus,
Pyrus, Rhododendron, Rhodura, Rubus, Salix, Scilla, Spiraea, Taraxacum, Trifolium,
Tulipa, Vaccinium, Viburnum.
Andrena vicina Smith, 1853. Cat. Hym. Brit. Mus., v. 1 p. 112. 2.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 151 (type). —Cockerell, 1906. Psyche
13: 9. —Atwood, 1934. Canad. Jour. Res. 10: 207, 210, figs. 2, d (key). —Lanham, 1941.
Ent. Soc. Amer., Ann. 34: 705, 708. 2°, ¢ (key). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 132-134, fig. 23, table 3 (redescription).

Biology: Packard, 1869. Guide Study Ins., p. 144 (nesting habits, parasites). —Clements and
Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology). —Salt, 1927. Jour. Expt. Zool.
48: 251 (stylopization).
victima Smith. N. S. to Ont. and Conn. Possibly a syn. of A. vicina Smith.
Andrena victima Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 113. 9.
Andrena desponsa Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 114. 6.

Superfamily APOIDEA 1807

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 123, 124. 2, 3 (types). —Cockerell,
1906. Psyche 13: 36. 2 (type). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
134-135 (stylopization and its possible effect on tax. characters).

Genus ANDRENA Subgenus MICRANDRENA Ashmead

Micrandrena Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 89.
Type-species: Andrena melanochroa Cockerell. Monotypic and orig. desig.
(=Micrandrena pacifica Ashmead).
Andrena subg. Andrenella Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 210.
Type-species: Melitta minutula Kirby. Orig. desig.

Revision: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 242-333, figs. 1-89, maps 1-10, tables
1-11 (N. Amer. spp.).

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 208 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 301 (tax. characters).

annectens Ribble. Calif. (San Benito Co.). Pollen: Unknown, but visits flowers of Salix.
Andrena (Micrandrena) annectens Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 323, figs.
80-84. 2, ¢.
candidiformis Viereck and Cockerell. B. C. and Wash. to Calif., east to Wyo., Colo. and Ariz.
Pollen: Polylectic, visits a wide variety of flowers including Arctostaphylos patula, A.
pungens, Arenaria douglasii, Baccharis, Ceanothus cordulatus, C. cuneatus, C.
divaricatus, C. greggii, C. integerrimus, Chamaebatia foliolosa, Crepis, Cryptantha,
Eriodictyon californicum, Eriogonum, Heracleum lanatum, Linum lewisii, Lomatium
dissectum, L. triternatum, Malus, Montia perfoliata, Nemophila menziesii, Physocarpa
intermedia, Prunus subcordata, P. virginiana, Pyrus, Ranunculus californicus,
Rhamnus californicus, Ribes nevadense, Salix foliosus, S. laevigata, S. lasiolepis,
Solanum umbelliferum, Taraxacum, Thysanocarpus curvipes, Trifolium, Veronica
americana, Wyethia.
Andrena candidiformis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 33. d.
Andrena nigritarsis Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 53. 2.
Andrena semotula Cockerell, 1936. Pan-Pacifie Ent. 12: 149. 6.

Taxonomy: Cockerell, 1936. Pan-Pacifie Ent. 12: 150. ¢. —Lanham, 1941. Ent. Soc. Amer.,
Ann. 34: 706, 708. 2, ¢ (key). —Timberlake, 1951. Jn Linsley, In Muesebeck, Krombein
and Townes, U.S. Dept. Agr., Agr. Monog. 2: 1060 (synonymy). —Ribble, 1968. Nebr.
Univ. State Mus., Bul. 8: 288-298, figs. 49-53, map 4 (redescription, synonymy).

Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 291-292, table 6 (floral relationships,
stylopizatiom).
chlorogaster Viereck. B. C. to Calif., east to Idaho, Utah and Ariz. Parasite: Stylops duboisi
Bohart, Imparipes sp. Pollen: Polylectic, visits a wide variety of flowers including
Alyssum maritimum, Amsinckia, Arctostaphylos crustacea, A. glauca, A. mariposa, A.
pungens, Baccharis, Brassica, Capsella bursa-pastoris, Ceanothus crassifolius, C.
cuneatus, C. greggii, C. orcuttii, C. soredatus, Cercocarpus betuloides, Collinsia
heterophylla, Cryptantha, Daucus, Descourainia sophia, Eriodictyon californicum,
Geranium molle, Gilia, Lasthenia chrysostoma, Layia glandulosa, Lomatiwm
dasycarpum, L. dissectum, L. triternatum, L. utriculatum, Montia perfoliata,
Nemophila menziesii, Phacelia ciliata, P. distans, Prunus subcordata, Ranunculus
californicus, R. canus, Rhamnus crocea, Rhus diversiloba, Salix gooddingii, S.
lasiolepis, S. leucodendroides, S. nigra, Sambucus, Sanicula bipinnatifida, S.
crassicaulis, S. nevadensis, S. tuberosa, Sedum, Sisymbrium irio, Sonchus, Tamarix,
Taraxacum officinale, Trifolium pratense.
Andrena chlorogaster Viereck, 1904. Canad. Ent. 35: 189. °.
Andrena subtilicornis Viereck, 1926. Pomona Jour. Ent. Zool. 18: 4. 6.
Andrena (Micrandrena) nitidicornis Cockerell, 1936. Pan-Pacifie Ent. 12: 144. 6.
Andrena catalinica Cockerell, 1939. Pomona Jour. Ent. Zool. 31: 25. 2.
Andrena (Micrandrena) chlorogaster nesiotes Timberlake, 1951. U. S. Natl. Mus., Proce.
101: 394. o.

1808 | Hymenoptera in America North of Mexico

Andrena (Micrandrena) chlorogaster gavilanica Timberlake, 1951. U. S. Natl. Mus., Proc.

101: 394. 3, @.
Andrena (Micrandrena) radialis Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 395. 6, °.
Taxonomy: Timberlake, 1951. U. S. Natl. Mus. Proc. 101: 396 (tax. characters, as
nitidicornis). —Ribble, 1965. Kans. Ent. Soc., Jour. 38: 87, fig. 1 (anomalous specimen).
—Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 279-288, figs. 41-48, map 2 (redescription,
synonymy).
Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 287-288, table 5 (floral relationships,
parasites).
illinoiensis Robertson. South. Canada (Man. to Alta.) and Ohio, Mich., west to Wash., Oreg.
and south. Calif. (Blythe), south to Tex., N. Mex. and Ariz. Parasite: Stylops bruneri
Pierce. Pollen: Polylectic, visits a wide variety of flowers including Amelanchier,
Amorpha fruticosa, Anemone, Antennaria plantaginifolia, Capsella, Cardamine,
Ceanothus ovatus, Cornus, Crataegus, Descourainia pinnata, Lepidium, Lomatium
daucifolium, L. foeniculaceum, Nasturtium, Prunus americana, Pyrus, Rhus
aromatica, Ribes missouriense, Rosa, Rubus, Salix amygdaloides, S. babylonica, S.
cordata, S. interior, S. nigra, Senecio, Shepherdia argentea, Sisymbriuwm, Sphaeralcea
coccinea, Spiraea thunbergii, Taraxacum officinale, T. vulgare, Thlaspi arvense, Zizia
aurea.
Andrena illinoiensis Robertson, 1891. Amer. Ent. Soe., Trans. 18: 54. 2, d.
Andrena salicinella Cockerell, 1895. Psyche 7 (sup.): 4. °.
Andrena placitae Cockerell, 1903. Ent. News 14: 215. ¢.
Andrena vegana Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 17. @.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 192. 2, ¢ (key). —Cockerell,
1914. Ann. and Mag. Nat. Hist. (8) 14: 10. 2 (key). —Lanham, 1941. Ent. Soe. Amer., Ann.
34: 706, 708. 2, ¢ (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 162-163,
figs. 29, 31-32, table 4 (redescription). —Ribble, 1968. Nebr. Univ. State Mus., Bul. 8:
299-308, figs. 59-63, map 6 (redescription, synonymy).
Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 306-307, table 8 (floral relationships,
stylopization).
ishii Ribble. Calif. Pollen: Unknown, but visits flowers of Salix.
Andrena (Micrandrena) ishii Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 319, figs. 70-74,
map 8. 9, 6.
labergei Ribble. Ariz. (Southwestern Research Station), N. Mex. (Fort Wingate). Pollen:
Unknown, but visits flowers of Salix.
Andrena (Micrandrena) labergei Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 321, figs.
75-79, map 8. 2, 6.
lamelliterga Ribble. Md., Pa., Ind., Tenn., IIl., Kans. Pollen: Unknown, but visits flowers of
Phacelia including P. dubia, P. ranunculacea.
Andrena (Micrandrena) lamelliterga Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 272,
figs. 31-35, map 4. 2, 6.
lepidii Ribble. South. Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Cryptantha, Lasthenia chrysostoma, Lepidium flavum.
Andrena (Micrandrena) lepidii Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 262, figs.
16-20, map 3. 2, 6.
melanochroa Cockerell. N. S. to N. C., west to B. C., Wash., Oreg and Calif. Pollen: Polylectic,
apparently with a preference for pollens of Fragaria and Potentilla, but visits a variety
of flowers including Ceanothus, Cerastium arvense, Cornus drummondi, Fragaria
virginiana, Kalmia, Potentilla canadensis, Ranunculus, Rubus, Salix, Taraxacum
officinale, Vaccinium, Waldsteinia.
Andrena melanochroa Cockerell, 1898. Entomologist 31: 89. 2, d.
Micrandrena pacifica Ashmead, 1899. Amer. Ent. Soc., Trans. 43: 54. 2, 6.
Andrena fragariana Graenicher, 1904. Ent. News 15: 64. 2, 6.
Andrena (Micrandrena) vagans Cockerell, 1932. Canad. Ent. 64: 157. 9.

Superfamily AROIDEA 1809

Taxonomy: Cockerell, 1932. Canad. Ent. 64: 157. 2 (key). —Cockerell, 1932. Canad. Ent. 64:
157, 158. 2, 3 (key, as fragariana). —Timberlake, 1951. In Linsley, In Muesebeck,
Krombein and Townes, U. S. Dept. Agr., Agr. Monog. 2: 1071 (synonymy). —Mitchell, 1960.
N. C. Agr. Expt. Sta. Tech. Bul. 141: 161, figs. 29-32, table 4 (redescription). —Ribble,
1968. Nebr. Univ. State Mus., Bul. 8: 293-299, figs. 54-58, map 5 (redescription, synonymy).

Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 297, table 7 (floral relationshiips,
stylopization).
micheneri Ribble. Tex. (Brownsville). Pollen: Unknown, but visits flowers of Lesquerella.
Andrena (Micrandrena) micheneri Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 264, figs.
21-25. 2, d.
microchlora Cockerell. B. C. to Calif., east to N. Dak., Wyo., Colo. and Ariz. Pollen: Polylectic
with some preference for the pollens of the Umbelliferae and to a lesser extent those of
the Cruciferae, Compositae, Caryophyllaceae and Ranunculaceae; visitation records
include Amsinckia, Arenaria californica, Brassica, Ceanothus cuneatus, Descourainia
sophia, Euphorbia albomarginata, Gilia, Helianthus, Lasthenia chrysostoma, Lomatium
dissectum, L. utriculatum, Musineon, Nemophila, Plectritis macrocera, Prunus
virginiana, Ranunculus californicus, Salix, Sanicula nevadensis, Taraxacum,
Tropidocarpum gracile.
Andrena microchlora Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 265. 3, 2.
Andrena microchlora var. subalia Cockerell, 1936. Pan-Pacific Ent. 12: 143. 9.

Taxonomy: Lanham, 1941. Ent. Soc. Amer., Ann. 34: 705, 707. 2°, 6 (key). —Timberlake,
1951. U. S. Natl. Mus., Proc. 101: 397 (tax. characters and status). —Ribble, 1968. Nebr.
Univ. State Mus., Bul. 8: 274-279, figs. 36-40, map 3 (redescription, synonymy).

Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 277, table 4 (floral relationships,
stylopization).
neonana Viereck. N. J. to Fla., west to Ark. and Tex. Pollen: Unknown, but visits flowers of
Fragaria virginiana, Ilex, Prunus persica, Pyrus.
Andrena (Scrapter) neonana Viereck, 1917. Amer. Ent. Soc., Trans. 43: 400. 9.

Taxonomy: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 259-262, figs. 11-15, map 3
(redescription).
nigrae Robertson. Conn. to north. Fla., west to Alta., Wash., Idaho, Utah and Tex. Pollen:
Apparently polylectic, although majority of collections have been made at the flowers of
Salix; visitation records include Alnus, Anemone virginiana, Antennaria
plantaginifolia, Arabis laevigata, Barbarea, Capsella, Cotoneaster, Crataegus,
Cymopterus acaulis, Descourainia pinnata, Erigenia bulbosa, Lomatium daucifolium,
L. dissectum, Prunus americana, P. angustifolia, P. serotina, P. virginiana, Pyrus,
Rhus aromatica, Ribes, Rubus, Salix amygdaloides, S. babylonica, S. cordata, S.
humilis, S. interior, S. nigra, S. sericea, Sassafras albidum, Sphaeralcea coccinea,
Spiraea, Taraxacum, Thlaspi arvense, Viburnum, Zizia aurea.
Andrena illinoensis var. bicolor Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 46. 9.
Preoce.
Andrena nigrae Robertson, 1905. Canad. Ent. 37: 237. N. name.
Andrena salicinellina Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 17. 3.
Andrena (Andrena) abacta Viereck, 1917. Amer. Ent. Soc., Trans. 43: 365. 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 165. 2. —Ribble, 1968. Nebr.
Univ. State Mus., Bul. 8: 313-319, fig. 69, map 8 (redescription, synonymy).

Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 317, table 10 (floral relationships,
stylopization).
personata Robertson. Pa. to Ga., west to Nebr., Kans., Okla. and Tex. Pollen: Polylectic, visits

a wide variety of flowers, especially those of the Rosaceae, Salicaceae, Umbelliferae,
and Hydrophyllaceae in that order; visitation records include Alliaria officinalis,
Apocynum cannabinum, Arenaria patula, Aruncus, Claytonia virginica, Cornus,
Crataegus crus-galli, C. mollis, Ellisia nyctelea, Erigeron, Euphorbia commutata,
Fragaria, Geranium carolinianum, Lesquerella, Osmorhiza longistylus, Oxalis violacea,

1810 Hymenoptera in America North of Mexico

Pastinaca sativa, Perideridea americana, Phacelia dubia, P. ranunculacea, P. purshii,
Potentilla canadensis, Prunus, Ranunculus abortivus, R. septentrionalis, Rubus
flagellaris, Salix nigra, Sanicula marilandica, Sisymbrium, Spiraea, Taenidia
integerrima, Thaspium barbinode, T. trifoliatum, Viburnum molle.

Andrena personata Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 336. ?, 6.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193. 2, ¢ (key). —Cockerell, 1932.
Canad. Ent. 64: 157, 158. 2, 3d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
165-167, figs. 28, 31-33, table 4 (redescription). —Ribble, 1968. Nebr. Univ. State Mus., Bul.
8: 254-259, figs. 6-10, map 2 (redescription).

Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 257-258, table 2 (floral relationships,
stylopization).
piperi Viereck. B. C. to Calif., east to Idaho, Colo. and Ariz. (Tucson); Mexico (Baja California).
Pollen: Polylectic, visits a wide variety of flowers, especially those of the Cruciferae;
visitation records include Agoseris auriantiaca, Alyssum, Amsinckia, Barbarea
orthoceras, Brassica alba, B. campestris, B. nigra, Capsella bursa-pastoris, Cardaria
draba, C. pubescens, Ceanothus, Cirsium, Cryptantha intermedia, C. muricata,
Descourainia pinnata, D. sophia, Erysimum repandum, Lasthenia chrysostoma,
Lepidium, Lobularia maritima, Lomatium dissectum, Lupinus, Medicago sativa,
Penstemon, Phacelia fremontii, Platystemon californicus, Polygonum, Prunus
subcordata, Ranunculus californicus, Raphanus sativus, Ribes, Rubus, Salix lasiolepis,
Salsola kali, Sisymbrium altissimum, S. irio, S. pinnatum, Stellaria media, Tamarix,
Taraxacum officinale, Thlaspi arvense, Tropidocarpum gracile.
Andrena piperi Viereck, 1904. Canad. Ent. 35: 189. 9.

Taxonomy: Timberlake, 1951. U.S. Natl. Mus., Proe. 101: 392. 6. —Ribble, 1968. Nebr. Univ.
State Mus., Bul. 8: 248-254, figs. 1-5, map 1 (redescription).

Biology: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 252-254, table 1 (floral relationships,
stylopization),
salictaria Robertson. Transcont. south. Canada (N. S. to B. C.) and Maine to N. C. west to

Wash., Utah, Colo. and N. Mex. Parasite: Stylops bruneri Pierce. Pollen: Apparently
narrowly polylectic, chiefly Salix including S. bebbiana, S. cordata, S. humilis, but visits
other flowers including Amelanchier, Antennaria, Barbarea, Chamaedaphne, Claytonia,
Cotoneaster, Crataegus, Lomatium, Prunus americana, Rhus, Sassafras, Viburnum
opulus.

Andrena salictaria Robertson, 1905. Canad. Ent. 37: 236. 2, 6.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 207, 210. 2, 6 (key). —Lanham, 1941. Ent.
Soc. Amer., Ann. 34: 705, 706, 708. 2°, 6 (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 167-168, figs. 30-32, table 4 (redescription). —Ribble, 1968. Nebr. Univ. State
Mus., Bul. 8: 309-313, figs. 64-68, map 7 (redescription).

Biology: Salt, 1927. Jour. Expt. Zool. 48: 252 (stylopization).

Taxonomy: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 311-312, table 9 (stylopization, floral
relationships).
ziziae Robertson. New England to Ga., west to Man., N. Dak., S. Dak., Nebr., Kans. and N.
Mex. Parasite: Imparipes sp. Pollen: Polylectic although nearly all floral records are
from Zizia including Z. aptera, Z. aurea, but visits other flowers including Amorpha
fruticosa, Aruncus dioicus, Barbarea vulgaris, Ceanothus ovatus, Echinacea
angustifolia, Hieracium, Pastinaca sativa, Polytaenia nuttallii, Rudbeckia
amplexicaulis, Sanicula marilandica, Taenidia integerrima, Thaspium barbinode, T.
trifoliatum.
Andrena ziziae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 55. 2, 6.
Andrena (Micrandrena) vernalis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 168,
fig2Z9n1?%

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 198. 2, ¢ (key). —Cockerell, 1932.
Canad. Ent. 64: 157, 158. 2, d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:

Superfamily APOIDEA 1811

168-169, figs. 31-32, table 4 (redescription). — Ribble, 1968. Nebr. Univ. State Mus., Bul. 8:
267-271, figs. 26-30, map 1 (redescription, synonymy).

Biology: Ribble, 1968. Nebr. State Mus., Bul. 8: 270, table 3 (floral relationships, parasite).

Genus ANDRENA Subgenus NEMANDRENA LaBerge

Andrena subg. Nemandrena LaBerge, 1971. Pan-Pacific Ent. 47: 48.
Type-species: Andrena torulosa LaBerge. Orig. desig.

Revision: LaBerge, 1971. Pan-Pacific Ent. 47: 47-57, 15 figs. (N. Amer. spp.).

Biology: Cruden, 1972. Evolution 26: 373-389, 8 figs., 9 tables (floral relationships). —Cruden,
1972. Madrono 21: 505-515, 2 tables (floral relationships).
crudeni LaBerge. Calif. Pollen: Apparently oligolectic on flowers of Nemophila including N.
maculata, N. menziesii menziesii.
Andrena (Nemandrena) crudeni LaBerge, 1971. Pan-Pacifie Ent. 47: 54, figs. 6-10. 2, ¢.
subnigripes Viereck. Calif. Pollen: Collects pollen from flowers of Nemophila menziesii
menziesii.
Andrena (Andrena) subnigripes Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 581. 2.
torulosa LaBerge. Oreg., Calif. Pollen: Apparently oligolectic on flowers of Nemophila
including N. menziesii atomeria, N. m. liniflora, N. m. menziesii, but visits other
flowers presumably for nectar including Arctostaphylos, Limnanthes douglasii,
Platystemon californica.
Andrena (Nemandrena) torulosa LaBerge, 1971. Pan-Pacific Ent. 47: 49, figs. 1-5. 2, 3.

Genus ANDRENA Subgenus OLIGANDRENA Lanham

Andrena subg. Oligandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 207.
Type-species: Andrena macrocephala Cockerell. Orig. desig.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 284, 290, 301 (tax. characters).
angelesia Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Cryptantha
intermedia, Encelia farinosa, Platystemon californicus.
Andrena (Oligandrena) angelesia Timberlake, 1951. U. S. Natl. Mus., Proce. 101: 404. 2, ¢.
anisochlora Cockerell. Oreg., Calif. Pollen: Unknown, but visits flowers of Amsinckia
douglasiana, Claytonia spathulata, Lasthenia, Montia perfoliata, Nemophila
heterophylla.
Andrena anisochlora Cockerell, 1936. Pan-Pacifie Ent. 12: 137. 2.
Andrena (Micrandrena) dinognatha Timberlake, 1938. Pan-Pacific Ent. 14: 26. ¢.

Taxonomy: Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U.S. Dept.
Agr., Agr. Monog. 2: 1056 (synonymy).
macrocephala macrocephala Cockerell. Calif. (lowlands); Mexico (Baja California). Parasite:

Stylops timberlakei Bohart. Pollen: Collects pollen from flowers of Nemophila including
N. menziesii, N. integrifolia as well as Pholistoma auritum, but visits other flowers
presumably for nectar including Calandrinia ciliata, Cryptantha intermedia,
Eriogonum fasciculatum, Erodium, Gilia, Orthocarpus densiflorus, Phacelia distans,
Ranunculus, Rhus trilobata, Sisymbrium irio.

Andrena macrocephala Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 278. 6.

Andrena peratra Cockerell, 1916. Pomona Jour. Ent. Zool. 7: 46. °.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 281. 2, d (synonymy, floral
relationships, parasite).

Biology: Cruden, 1972. Evolution 26: 378-380, fig. 7, tables 3, 4 (floral relationships).
—Cruden, 1972. Madrono 21: 505-515, 1 fig. 2 tables (floral relationships).

macrocephala tetleyi Linsley. South. Calif. (montane). Pollen: Collects pollen from flowers of
Nemophila integrifolia.
Andrena macrocephala tetleyi Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 282. 9.

1812 Hymenoptera in America North of Mexico

Biology: Cruden, 1972. Evolution 26: 379 (floral relationships). —Cruden, 1972. Madrono 21:
511 (floral relationships).
nigroclypeata Linsley. Calif. Pollen: Unknown, but visits flowers of Lasthenia, Layia
platyglossa, Marah, Phacelia alba, Platystemon californicus, Pholistoma racemosa,

Raphanus sativa, Tamarix.
Andrena nigroclypeata Linsley, 1939. Pan-Pacific Ent. 15: 155. 9, d.

Genus ANDRENA Subgenus ONAGRANDRENA Linsley and MacSwain

Andrena subg. Onagrandrena Linsley and MacSwain, 1956. Pan-Pacifie Ent. 32: 111.
Type-species: Andrena oenotherae Timberlake. Orig. desig.

The species of this subgenus are chiefly dependent upon the pollen and nectar of several
genera within the family Onagraceae, especially Camissonia, Clarkia, and Oenothera. These
bees are largely or entirely dark or black in coloration and their behavior is usually closely
synchronous with the diurnal flowering regimes of their host plants. Thus some of these bees
are matinal species while others are active at the flowers only in the late afternoon and early
evening and still others are active at the flowers both in the early morning and again in the late
afternoon and early evening hours.

Taxonomy: Linsley and MacSwain, 1955. Pan-Pacific Ent. 31: 163-172 (as Melandrena).
—Linsley and MacSwain, 1956. Pan-Pacifie Ent. 32-111-114 (tax. characters, included spp.).
—Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 117-130 (Oenothera and Clarkia
visiting spp.). —Linsley and MacSwain, 1962. Pan-Pacific Ent. 38: 49-52 (new spp. from
Calif., Nev., Wyo.). —Linsley and MacSwain, 1963. Pan-Pacifie Ent. 39: 189-198 (oenotherae
complex). —Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 5-6 (key to
Colorado Desert spp.). —Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33:
31 (key to Great Basin spp.). —Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent.
33: 64-65 (key to Mojave Desert spp.). —LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4:
284, 287, 290, 292, 315 (tax. characters). —Linsley, MacSwain, Raven and Thorp, 1973.
Calif. Univ. Pubs. Ent. 71: 27 (key to cismontane Calif. and Baja California spp.).

Biology: Linsley, MacSwain and Smith, 1955. Pan-Pacific Ent. 31: 173-185 (nesting habits,
flower relationships). —Linsley and MacSwain, 1956. Pan-Pacific Ent. 32: 114-121 (nesting
habits, floral relationships). —Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent.
33: 1-58, 6 plates, 6 text figs., 19 tables (comparative behavior of Oenothera visiting spp.
on the Colorado Desert and Great Basin). —Linsley, MacSwain and Raven, 1964. Calif.
Univ. Pubs. Ent. 33: 59-98, 3 plates, 17 tables (comparative behavior of Oenothera visiting
spp. on the Mojave Desert). —MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent.
70: 1-80, 3 plates, 10 text figs., 18 tables (comparative behavior of Clarkia visiting spp. in
western U. S.). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71:
1-68, 6 plates, 15 text figs., 10 tables (comparative behavior of Camissonia and Oenothera
visiting spp. in cismontane Calif. and Baja California).

anograe anograe Cockerell. Calif., Colo. Wyo. Pollen: Collects pollen from Oenothera
caespitosa, but visits other onagraceous flowers including Anogra, Oenothera albicaulis,
O. caespitosa var. montana, and may collect pollen from these as well as the large white
flowered evening primroses of the subgenus Pachylophis.
Andrena anograe Cockerell, 1901. Canad. Ent. 33: 154. °.
Andrena micranthophila Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 482. 2, 6.

Taxonomy: Cockerell, 1934. Amer. Mus. Novitates 697: 2 (tax. characters). —Linsley and
MacSwain, 1955. Pan-Pacific Ent. 31: 166 (synonymy, geogr. records). —Linsley and
MacSwain, 1961. Pan-Pacific Ent. 37: 126 (changed status). —Linsley, MacSwain, and
Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31 (tax. characters).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 39 (floral
relationships).

anograe knowltoni Linsley and MacSwain. Utah. Pollen: Unknown, but visits flowers of

Camissonia scapoidea scapoidea, Stanleya pinnata.
Andrena (Onagrandrena) anograe knowltoni Linsley and MacSwain, 1961. Pan-Pacific

Ent. 37: 126. 9.

Superfamily APOIDEA 1813

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 39 (floral
relationships).
bernardina Linsley. Calif. (San Bernardino Mts.). Pollen: Possibly oligolectie on Clarkia, but
visits other flowers for nectar including Potentilla glandulosa.
Andrena bernardina Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 275. 2, d.

Taxonomy: Linsley and MacSwain, 1955. Pan-Pacifie Ent. 31: 166 (tax. status). —Linsley and
MacSwain, 1961. Pan-Pacific Ent. 37: 128 (tax. status).

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 16, fig. 2, table 6
(floral relationships).
blaisdelli Cockerell. South. Calif. Pollen: Principal source of pollen is Camissonia campestris,
but also collects pollen from C. bistorta, C. californica, C. contorta; visits these and
other flowers for nectar including Cryptantha intermedia, Eriophyllum confertiflorum.
Andrena blaisdelli Cockerell, 1924. Pan-Pacifie Ent. 1: 59. 9.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 266, 267. 2, ¢ (key). —Linsley,
MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 27 (key).

Biology: Linsley, MacSwain, Raven and Thorp, 1978. Calif. Univ. Pubs. Ent. 71: 27-29, fig. 12
(floral relationships).
boronensis Linsley and MacSwain. Calif. (Mojave Desert). Parasite: Stylops sp. Pollen:
Collects pollen almost exclusively from the flowers of Camissonia campestris, but has
been observed collecting pollen from the flowers of C. clavaeformis and C. kernensis;
nectar requirements are chiefly satisfied at the flowers of C. campestris, but other
flowers are visited for nectar including Baccharis viminea, Camissonia bistorta,
Cryptantha intermedia, Encelia farinosa, Eriophyllum confertiflorum, Layia
glandulosa, Potentilla, Salix lasiolepis.
Andrena (Onagrandrena) boronensis Linsley and MacSwain, 1962. Pan-Pacifie Ent. 38: 49.
2. @-

Taxonomy: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 64, 65, 69 (key,
identity).

Biology: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 69-71, tables 3-4,
7-15 (nest architecture, male behavior, floral relationships, parasite).
camissoniae Linsley and MacSwain. Calif. (Santa Barbara County). Pollen: Apparently obtains
pollen from the flowers of Camissonia campestris.
Andrena (Onagrandrena) camissoniae Linsley and MacSwain, 1968. Pan-Pacific Ent. 44:
144. 2.

Taxonomy: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 27, 29
(key, tax. characters).

Biology: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 29 (floral
relationship).
chylismiae Linsley and MacSwain. Calif. (Owens Valley), Nev. Parasite: Nomada sp., Stylops

sp. Pollen: Collects pollen from the flowers of Camissonia clavaeformis cruciformis and
C. c. integrior, but visits these and other flowers for nectar including Haplopappus
acaulis, Sisymbrium altissimum, Stanleya pinnata, Taraxacum officinale.

Andrena (Onagrandrena) chylismiae Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37:
LUE Seid).

Taxonomy: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31 (key).
Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 37-38, fig. 6, tables
16, 17, 19 (floral relationships, parasites).
convallaria convallaria Linsley and MacSwain. Calif. (Central Valley and south. cent. Coast
Ranges). Pollen: Collects pollen principally from the flowers of Camissonia campestris

although occasionally obtains pollen from the flowers of C. californica, C.
cheiranthifolia, C. c. obispoensis, C. contorta, C. sierrae; visits these and other flowers

1814 Hymenoptera in America North of Mexico
for nectar including Brassica, Camissonia micrantha, Encelia actoni, Isomeris arborea,

Lasthenia, Lupinus, Malacothrix.
Andrena (Onagrandrena) convallaria convallaria Linsley and MacSwain, 1963.
Pan-Pacific Ent. 39: 193. 9, 6.
Taxonomy: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 27 (key).

Biology: Linsley, MacSwain, Raven and Thorp, 1973. Univ. Calif. Pubs. Ent. 71: 29-31, fig. 13,
tables 1, 3-8 (floral relationships).
convallaria subhyalina Linsley and MacSwain. Calif. (west. margin Mojave Desert). Parasite:
Stylops sp. Pollen: Collects pollen almost exclusively from the flowers of Camissonia
campestris although occasionally obtains pollen from the flowers of C. abramsii; visits
these and other flowers for nectar including Aniscoma acaulis, Camissonia

clavaeformis clavaeformis, Coreopsis bigelovii, C. californica, Layia californica.
Andrena (Onagrandrena) convallaria subhyalina Linsley and MacSwain, 1963. Pan-Pacific

Ent. 39: 193. 2, 6.
Biology: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 71-72, tables 3,
11-12, 14 (mating, floral relationships, parasite).

deserticola Timberlake. Calif. (Mojave Desert). Parasite: Nomada sp., Stylops sp. Pollen:
Collects pollen almost exclusively from the flowers of Camissonia campestris although
on one occasion collected pollen from the flowers of C. kernensis; visits these and other
flowers for nectar including Baileya multiradiata, Camissonia abramsu, C.
clavaeformis, C. boothii decorticans, Coreopsis bigelovii, C. californica, Isomeris

arborea.
Andrena deserticola Timberlake, 1937. Pan-Pacific Ent. 12: 173. @.

Taxonomy: Linsley and MacSwain, 1955. Pan-Pacific Ent. 31: 172 (geogr. and floral records).
—Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 64, 65 (key). —Linsley,
MacSwain, Raven and Thorp, 1978. Calif. Univ. Pubs. Ent. 71: 27 (key).

Biology: Linsley, MacSwain and Smith, 1955. Pan-Pacific Ent. 31: 182-183 (nest architecture,
parasites). —Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 68-69, tables
3-4, 7-11, 15 (nest architecture, floral relationships, male behavior, parasite). —Linsley,
MacSwain, Raven and Thorp, 19738. Calif. Univ. Pubs. Ent. 71: 31, table 1 (floral
relationships).

flandersi Timberlake. South. Calif (Mojave Desert). Pollen: Collects pollen from the flowers of

Camissonia campestris, C. clavaeformis, C. contorta, C. kernensis, but visits these and
other flowers for nectar including Chaenactis, Coreopsis bigelovii, C. californica,
Cryptantha, Ericameria, Lasthenia chrysostoma, Layia glandulosa.

Andrena flandersi Timberlake, 1937. Pan-Pacific Ent. 12: 72. °.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 266, 278. ° (key). —Linsley and
MacSwain, 1955. Pan-Pacific Ent. 31: 167 (geogr. and floral records). —Linsley, MacSwain
and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 64, 65 (key).

Biology: Linsley and MacSwain, 1956. Pan-Pacific Ent. 32: 118-120 (nest site, floral
relationships). —Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 72, tables

38, 7, 11-15 (floral relationships).
furva Linsley and MacSwain. South. Calif.; Mexico (Baja California). Pollen: Principally
gathers pollen from Camissonia campestris although it has been observed gathering

pollen from Clarkia cylindrica and occasionally with mixed pollen loads of Camissonia

and Clarkia.
Andrena (Onagrandrena) furva Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 117. 9.

Taxonomy: Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 64, 73 (key,
geogr. records). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71:

27 (key).
Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 46 (floral
relationships). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71:

32-33, fig. 15 (floral relationships).

Superfamily APOIDEA 1815

linsleyi Timberlake. South. Calif., southeast. Nev.; Mexico (Baja California). Parasite: Nomada
sp., Stylops sp. Pollen: Usually collects pollen from Oenothera deltoides, but occasionally
takes pollen from Camissonia clavaeformis including C. c. aurantiaca C. c. clavaeformis
and also occasionally from C. decorticans desertorum, Oenothera trichocalyx; visits
these and other flowers for nectar including Baileya, Dithyrea californica, Encelia,
Geraea canescens, Hyptis emoryi.
Andrena linsleyi Timberlake, 1937. Pan-Pacifie Ent. 12: 71. 9.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 266. 2 (key). —Linsley and
MacSwain, 1955. Pan-Pacific Ent. 31: 165, 166, 170-171 (key, geogr. and floral records).
—Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 6 (key).

Biology: Linsley, MacSwain and Smith, 1955. Pan-Pacific Ent. 31: 176 (floral relationships).
—Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 11-16, fig. 2, tables 1, 3,
7-12 (floral relationships, parasites). —Linsley, MacSwain and Raven, 1964. Calif. Univ.
Pubs. Ent. 3: 73 (floral relationships).

mojavensis Linsley and MacSwain. Calif. (jawbone and Short Canyons, Kern County). Pollen:
Principal pollen source is Camissonia kernensis, but also takes pollen from C.
campestris, C. clavaeformis, C. dentata johnstonii, C. d. parishii; visits these and other
flowers for nectar including Coreopsis bigelovii, Dithyrea californica, Salix.
Andrena (Melandrena) mojavensis Linsley and MacSwain, 1955, Pan-Pacific Ent. 31: 171.
Oo. GE

Taxonomy: Linsley and MacSwain, 1955. Pan-Pacific Ent. 31: 165 (key). —Linsley, MacSwain
and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 64, 65 (key).

Biology: Linsley, MacSwain and Smith, 1955. Pan-Pacifie Ent. 31: 179-181 (nest architecture,
floral relationships). —Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33:
65-67, tables 5-10 (nest architecture, floral relationships, male behavior).

nevadae Linsley and MacSwain. Nev. (Eureka and White Pine Counties). Pollen: Unknown,
presumably collects pollen from the flowers of the subgenera Anogra or Pachylophis;
visits flowers of Camissonia clavaeformis and Stanleya pinnata for nectar.
Andrena (Onagrandrena) nevadae Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 125.
oF

Taxonomy: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31 (key).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 39 (floral
relationships).
oenotherae Timberlake. Calif. (Coastal Santa Barbara Co. east to Colorado Desert); Mexico

(Baja California). Parasite: Lytta stygica (LeC.)?, Meloe sp.?, Stylops sp. Pollen:
Principally collects pollen from the flowers of Camissonia cheiranthifolia
cheiranthifolia on Santa Cruz Island, and C. c. suffruticosa along the mainland coast
while more interior populations collect from C. bistorta; visits these and other flowers
for nectar including Cakile edentula, Calandrinia, Chaenactis, Chrysanthemum
coronarium, Coreopsis californica, Croton californicus, Cryptantha intermedia,
Eriogonum fasciculatum, Haplopappus cooperi, Hemizonia kelloggii, Lantana, Layia
platyglossa, Lupinus, Salix lasiolepis, Sisymbrium.

Andrena oenotherae Timberlake, 1937. Pan-Pacific Ent. 12: 69. 2, d.

Taxonomy: Linsley, 1988. Calif. Acad. Sci., Proc. (4) 23: 266, 267. 2, d (key). —Linsley and
MacSwain, 1955. Pan-Pacific Ent. 31: 164, 166, 170 (key, geog. and floral records).
—Linsley and MacSwain, 1963. Pan-Pacific Ent. 39: 189-190. —Linsley, MacSwain, Raven
and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 27 (key).

Biology: Linsley, MacSwain and Smith, 1955. Pan-Pacific Ent. 31: 181-182 (nest architecture,
floral relationships, parasites). —Linsley and MacSwain, 1956. Pan-Pacifie Ent. 32: 116-118,
1 fig. (nest architecture, floral relationships, parasite). —Linsley and MacSwain, 1963.
Pan-Pacific Ent. 39: 189-190 (floral relationships). —Linsley, MacSwain, Raven and Thorp,
1973. Calif. Univ. Pubs. Ent. 71: 33-34, fig. 14, table 9 (floral relationships).

1816 Hymenoptera in America North of Mexico

omninigra clarkiae Linsley and MacSwain. Calif. (Monterey and San Luis Obispo Counties).
Pollen: Collects pollen from the flowers of Clarkia cylindrica, C. speciosa.
Andrena (Onagrandrena) omninigra clarkiae Linsley and MacSwain, 1961. Pan-Pacific
Hntyotelctie gene).
Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 15-16, fig. 1, tables
7-11 (floral relationships, mating behavior).
omninigra omninigra Viereck. Calif. (west. slopes of Sierra Nevada Mts.). Pollen: Collects
pollen from the flowers of Clarkia including C. biloba, C. cylindrica, C. dudleyana, C.
purpurea quadrivulnera, C. rhomboidea, C. speciosa polyantha, C. unguiculata, C.
xantiana; visits these and other flowers for nectar including Cryptantha flaccida, Gilia
capitata. Predator: Callinicus calcaneus (Loew).
Andrena (Andrena) omninigra Viereck, 1917. Amer. Ent. Soc., Trans. 43: 385. 9.
Andrena grundeli Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 274. 9, 6.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 265. 2 (key). —Linsley and
MacSwain, 1955. Pan-Pacifie Ent. 31: 164, 165, 166 (key, geogr. records). —Linsley and
MacSwain, 1961. Pan-Pacific Ent. 37: 27 (type).

Biology: Linsley, 1972. Pan-Pacific Ent. 48: 94-96 (predator). —MacSwain, Raven and Thorp,
1973. Calif. Univ. Pubs. Ent. 70: 14, fig. 1, tables 1, 6-9, 13-18 (floral relationships).
oraria actitis Linsley and MacSwain. Calif. (San Francisco). Pollen: Collects pollen from the
flowers of Camissonia cheiranthifolia cheiranthifolia.
Andrena (Onagrandrena) oraria actitis Linsley and MacSwain, 1963. Pan-Pacific Ent. 39:
1967 Od:
Biology: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 35, fig. 15
(floral relationship).
oraria oraria Linsley and MacSwain. Calif. (Marin and Sonoma Counties). Pollen: Collects
pollen from the flowers of Camissonia including C. cheiranthifolia cheiranthifolia, C.

ovata.
Andrena (Onagrandrena) oraria oraria Linsley and MacSwain, 1963. Pan-Pacific Ent. 39:

196) Oh de
Biology: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 35, fig. 15
(floral relationships).
raveni Linsley and MacSwain. Oreg., Calif., Nev., Utah. Parasite: Nomada sp., Stylops sp.
Pollen: Collects pollen from the flowers of Camissonia clavaeformis clavaeformis, C. c.
integrior, C. tanacetifolia, Oenothera deltoides piperi, but visits these and other flowers
for nectar including Agoseris, Madia, Stanleya pinnata, Taraxacum officinale.
Andrena (Onagrandrena) raveni Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37: 118. 2,
é.
Taxonomy: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31 (key).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31-36, fig. 4, tables
14-19 (nest architecture, floral relationships, parasites).
rozeni Linsley and MacSwain. Ariz., south. Calif., Nev.; Mexico (Baja California), deserts.

Parasite: Nomada sp., Stylops sp. Pollen: Collects pollen from the flowers of
Camissonia including C. clavaeformis aurantiaca, C. c. clavaeformis, C. c. integrior, C.
c. peirsonii, C. decorticans, C. latifolia, C. tanacetifolia, C. trichocalyx and Oenothera
deltoides piperi; visits these and other flowers for nectar including Agoseris glauca,
Brassica, Cryptantha clevelandii, C. intermedia, Encelia farinosa, Geraea canescens,
Madia ramii, Mirabilis, Sisymbrium altissimum, Stanleya pinnata, Taraxacum

officinale.
Andrena (Melandrena) rozeni Linsley and MacSwain, 1955. Pan-Pacific Ent. 31: 168. 2, d.

Taxonomy: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 5, 31 (key).
—Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 64, 65 (key).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 6-11, 36-87, figs. 1, 5,
tables 1-6, 17-18 (nest architecture, floral relationships, male behavior, parasites).

Superfamily APOIDEA 1817

—Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 72-73, table 16 (floral
relationships).
rubrotincta Linsley. South. Calif., northwest. Ariz., south. Nev., southwest. Utah. Pollen:
Collects pollen regularly only from Camissonia brevipes pallidula and C. parryi,
although females have been taken at flowers of C. brevipes brevipes and C. clavaeformis
aurantiaca with pollen in their scopae.
Andrena rubrotincta Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 278. 9.

Taxonomy: Linsley and MacSwain, 1955. Pan-Pacific Ent. 31: 165, 170 (key, geogr. records).
—Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 5, 6 (key).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 16-18, fig. 3, table 13
(floral relationships).
stagei Linsley and MacSwain. Wyo. (Sweetwater Co.). Pollen: Collects pollen from the flowers
of Oenothera trichocalyx.
Andrena (Onagrandrena) stagei Linsley and MacSwain, 1962. Pan-Pacific Ent. 38: 52. 2.

Taxonomy: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31 (key).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 38 (floral
relationships).
thorpi Linsley and MacSwain. Nev. (Humboldt Co.). Parasite: Stylops sp. Pollen: Collects
pollen from the flowers of Oenothera deltoides piperi.
Andrena (Onagrandrena) thorpi Linsley and MacSwain, 1962. Pan-Pacifie Ent. 38: 51. 9.

Taxonomy: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 31 (key).

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 38-39, table 15
(floral relationships, parasite).

vanduzeei Linsley. Calif. Pollen: Possibly an oligolege of Gayophytum.
Andrena vanduzeei Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 280. 2, ¢.

Taxonomy: Linsley and MacSwain, 1955. Pan-Pacifie Ent. 31: 165, 166, 172 (key, geogr.
records).
vespertina Linsley and MacSwain. Calif. (Kern and Stanislaus Counties). Pollen: Collects pollen
only from the flowers of Camissonia boothii decorticans; visits flowers of Isomeris

arborea for nectar.
Andrena (Onagrandrena) vespertina Linsley and MacSwain, 1961. Pan-Pacifie Ent. 37:

eS PRG):
Taxonomy: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 27 (key).

Biology: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 35-36, fig. 15,
table 10 (floral relationships).

yumorum Viereck. South. Calif.
Andrena (Andrena) yumorum Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 585. d.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 128-129 (type).
Genus ANDRENA Subgenus OPANDRENA Robertson

Opandrena Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193.
Type-species: Andrena cressonii Robertson. Orig. desig.

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 210 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 284, 289, 303 (tax. characters).
cressonii cressonii Robertson. U. S. and southern Canada. Pollen: Apparently polylectic, visits

a wide variety of flowers including Amelanchier, Anemonella, Arabis, Aronia, Aruncus,
Brassica, Capsella, Ceanothus, Celastrus, Claytonia, Cornus, Crataegus, Dentaria,
Fragaria, Heracleum, Ilex, Oxalis, Pastinaca, Philadelphus, Polytaenia, Potentilla,
Prunus, Ptelea, Pyracantha, Radicula, Ranunculus, Rhamnus, Rhus, Rosa, Rubus,
Salix, Sanicula, Sisymbrium, Smilacina, Smilax, Stellaria, Taenidia, Taraxacum,
Thaspium, Tradescantia, Trifolium, Veronica, Viburnum, Zanthoxylum, Zizia.

Andrena Cressonii Robertson, 1891. Amer. Ent. Soc., Trans. 18: 56. 2, d.

1818 Hymenoptera in America North of Mexico

Andrena kansensis Cockerell, 1899. Ent. News 10: 255. 6.

Andrena bridwelli Cockerell, 1899. Ent. News 10: 255. ¢.

Andrena dubia Robertson, 1902. Canad. Ent. 34: 48. 6.

Andrena (Trachandrena) trumani Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48:
ie, OQ

Andrena latisigna Viereck and Cockerell, 1914. U.S. Natl. Mus., Proce. 48: 22. d.

Andrena cressoni transformans Cockerell, 1934. Pan-Pacific Ent. 9: 157. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193. 2, ¢ (key). —Lanham, 1949.
Pan-Pacific Ent. 25: 147 (tax status). —Timberlake and Lanham, 1951. Jn Linsley, In
Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr. Monog. 2: 1063 (synonymy).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 248-249, figs. 50, 52-54
(redescription).

cressonii infasciata Lanham. Calif., Oreg., Wash. Pollen: Presumably gathers pollen from
Gilia, but visits other flowers for nectar including Brassica campestris, Ranunculus
californicus, Pyracantha, Salix lasiolepis, S. nigra.
Andrena cressonii infasciata Lanham, 1949. Pan-Pacifie Ent. 25: 147. @, d.

Biology: Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 19 (floral relationships).
cressonii ivanensis Mitchell. Va., N. C. Pollen: Unknown, but visits flowers of Brassica,
Padus.
Andrena (Opandrena) cressonti ivanensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
141: 249, fig. 52. ¢.
cressonii tallahasseensis Mitchell. Fla., N. C. Pollen: Unknown, but visits flowers of Castanea
umila.
anurans (Opandrena) cressonii tallahasensis(!) Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 249, figs. 50, 53. 3.
Andrena (Opandrena) cressonii tallahasseensis Mitchell, 1967. In Krombein and Burks, U.
S. Dept. Agr., Agr. Monog. 2, Second Suppl., p. 428. Emend.
prunifloris Cockerell. N. Mex. Pollen: Unknown, but visits flowers of Prunus.
Andrena prunifloris Cockerell, 1898. Canad. Ent. 30: 147. 9.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 49. 2. (key).
Genus ANDRENA Subgenus PARANDRENA Robertson

Parandrena Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 337.
Type-species: Panurgus andrenoides Cresson. Desig. by Cockerell, 1897.

The species of this Nearctic subgenus are apparently oligoleges of Salix.

Revision: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 307-346, figs. 5-9, 41-70 (N.
Amer. spp.).

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 202 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 298 (tax. characters).
andrenoides (Cresson). South. Canada (B. C., Sask., Man. and Ont.), south to Fla., Tex., N.
Mex. and Utah. Pollen: Oligelege of Salix including S. amygdaloides, S. cordata, S.
interior, S. nigra, but visits these and other flowers for nectar including Amelanchier
canadensis, Anemone patens, Antennaria plantaginifolia, Crataegus crus-galli,
Erigenia bulbosa, Heracleum maximum, Lomatium foeniculaceum, Prunus americana,
P. angustifolia, P. gracilis, Pyrus, Rubus, Sassafras variifolium, Sisymbrium, Stellaria
media, Viola.
Panurgus andrenoides Cresson, 1878. Amer. Ent. Soc., Trans. 7: 62. ¢.
Panurgus rufocinctus Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 4. °.
Andrena andrenoides f. bicolor Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 47. Preoce.
Parandrena andrenoides var. clarigastra Viereck, 1908. Ent. News 19: 42. N. name.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 238-239, figs. 49-50, 52, 55,
table 6 (redescription).

Biology: Salt, 1927. Jour. Exp. Zool. 48: 250 (stylopization).

Superfamily APOIDEA 1819

arenicola LaBerge and Ribble. Kans., Okla., Tex. Pollen: Apparently an oligolege of Salix
including S. amygdaloides, S. longifolia, S. nigra, but visits other flowers including
Pyrus pyrus.
Andrena (Parandrena) arenicola LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98:
328, figs. 7, 538-54. 2, ¢.
Biology: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 330 (stylopization).
bucculenta LaBerge and Ribble. Calif. Pollen: Presumably an oligolege of Salix, but visits
other flowers including E'schscholzia californica, Layia platyglossa, Prunus.
Andrena (Parandrena) bucculenta LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98:
334, figs. 6, 60-61. 2, 6d.
concinnula (Cockerell). Oreg., Calif., Ariz. (Chiricahua Mts.). Pollen: Oligolege of Salix
including S. argophylla, S. cordata, S. exigua, S. gooddingii, S. hindsiana, S. laevigata,
S. lasiolepis, but visits other flowers presumably for nectar including Amsinckia
douglasiana, Capsella bursa-pastoris, Eschscholzia californica, Gilia, Sambucus,
Spiraea prunifolia, Tamarix gallica.
Parandrena concinnula Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 189. 3.
Andrena (Parandrena) mendosa Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 588. °.

Taxonomy: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 331-334, figs. 8, 55-59
(redescription, synonymy).
gibberis Viereck. B. C. (Fairview), Oreg. (Irrigon), Calif. (Lassen and Mono Counties), Nev.
(Elko), Utah (Cedar City). Pollen: Presumably an oligolege of Salix.
Andrena (Parandrena) gibberis Viereck, 1924. Canad. Ent. 56: 241. 2, 6.
nevadensis (Cresson). B. C. to Calif., Nev. and Utah. Pollen: Presumably an oligolege of Saliz,
but visits other flowers including Acer, Ribes, Senecio, Tamarix.
Panurgus nevadensis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 214. 6.
Andrena (Parandrena) garretti Viereck, 1924. Canad. Ent. 53: 248. 6.
Andrena (Parandrena) triangularis Viereck, 1924. Canad. Ent. 55: 243. °.

Taxonomy: Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 188. ¢. —LaBerge and Ribble, 1972.
Amer. Ent. Soc., Trans. 98: 341-344, figs. 7, 67-68 (redescription, synonymy).
nida Mitchell. Vt. to Ga., west to Mich., Il., Mo. and Miss. Pollen: Presumably an oligolege of
Salix including S. interior, but visits other flowers including Amelanchier, Crataegus,
Prunus, Rubus, Viola.
Andrena (Parandrena) nida Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 240, figs.
49, 50, 52, 55. 2, d.
Taxonomy: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 321-324, fig. 7, 46-47
(redescription).
papagorum Viereck and Cockerell. Ariz., south. Calif., Utah. Pollen: Presumably an oligolege
of Salix, visits flowers of S. gooddingii, S. nigra.
Andrena papagorum Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 2. 2.
Taxonomy: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 338-341, figs. 9, 62-66
(redescription).
wellesleyana Robertson. Mass. and Conn., west to N. W. T., Alta., and Utah. Pollen: Unknown,
but visits flowers of Prunus virginiana, Salix.
Andrena wellesleyana Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 337. 2, 6.
Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 241-242, figs. 50, 52, 55, table
6 (redescription). —LaBerge and Ribble, 1972. Amer. Ent. Soe., Trans. 98: 325-328, figs. 8,

42-52 (redescription).
Genus ANDRENA Subgenus PELICANDRENA LaBerge and Ribble

Andrena subg. Pelicandrena LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 346.
Type-species: Parandrena atypica Cockerell. Monotypic and orig. desig.

Revision: LaBerge and Ribble, 1972. Amer. Ent. Soc., Trans. 98: 346-351, figs. 71-76.

1820 Hymenoptera in America North of Mexico

atypica (Cockerell). Oreg (Klamath Falls), Calif.; Mexico (Baja California). Pollen: Apparently

polylectic with some preference for flowers of Cryptantha, Salix, Rhus, Rhamnus and
Ceanothus in approximately that order; visitation records include Adenostoma
fasciculatum, Allium haematochiton, Arctostaphylos bicolor, Baccharis, Brassica,
Ceanothus crassifolius, C. cuneatus, C. greggii, C. soredatus, C. verrucosus, Cryptantha
intermedia, Emmenanthe penduliflora, Gilia capitata, G. multicaulis, Lomatium
dasycarpum, Marah macrocarpus, Oenothera, Plagiobothrys nothofulvus, Prunus
ilicifolia, Quercus, Rhamnus californica, R. crocea, Rhus diversiloba, R. integrifolium,
R. ovata, R. trilobata, Rosa minutiflora, Salix argophylla, Salvia columbariae, S.
mellifera, Tamarix gallica.

Parandrena atypica Cockerell, 1941. San Diego Soc. Nat. Hist., Trans. 9: 347. 2.

Taxonomy: Thorp, 1969. Calif. Univ. Pubs. Ent. 52: 123 (tax. characters and status).
Genus ANDRENA Subgenus PLASTANDRENA Hedicke

Andrena subg. Plastandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 217.
Type-species: Melitta tibialis Kirby. Orig. desig.

Andrena subg. Schizandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 218.
Type-species: Andrena aulica Morawitz. Orig. desig.

Revision: LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 2-34, figs. 1-23, 3 tables (N. Amer. spp.).

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 201 (tax. characters, as Schizandrena).
—LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 297 (tax. characters, synonymy).
argemonis Cockerell. Nebr. and Colo. south to N. Mex. and Ariz.; Mexico (Chihuahua,
Coahuila, and Sonora south to Oaxaca). Pollen: Polylectic, visits flowers of Acacia,
Argemone mexicana, A. platyceras, Cleome serrulata, Medicago sativa, Melilotus alba,
Penstemon occidentalis, Persicaris pennsylvanicus, Sisymbrium irio.
Andrena argemonis Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 80. 6.

Taxonomy: Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 449. 2. —Cockerell, 1931. Amer.
Mus. Novitates 458: 16. 2, ¢ (key). —Lanham, 1941. Ent. Soc. Amer., Ann. 34: 704, 707. 2,
36. —LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 12-15, figs. 24-28 (redescription).

crataegi Robertson. Transcont. in southern Canada (N. S. to B. C.) and northern U. S., south

to Ga., Ala., Ark., Tex., N. Mex., Utah, Nev. and north. Calif. Pollen: Polylectic with
some preference for the flowers of the family Rosaceae, visits flowers of Amelanchier,
Amorpha, Aruncus, Astragalus, Barbarea, Blephilia, Brassica, Caragana, Cardaria,
Carduus, Castanea, Ceanothus, Chamaebatia foliolosa, Chrysanthemum, Coniwm,
Cornus, Crataegus, Crepis, Cryptantha intermedia, Daucus, Deutzia, Eulophus,
Euphorbia, Fragaria, Frasera, Gilia achillaefolia, Hackelia, Heracleum lanatum,
Holodiscus, Hydrangea, Ilex, Iris, Ledum, Lomatium dasycarpum, Malvastrum,
Melilotus, Monarda, Narcissus, Opulaster, Pastinaca, Philadelphus, Polytaenia,
Potentilla, Prunus, Ptelea, Purshia, Pyracantha, Pyrus, Ranunculus, Raphanus,
Rhamnus, Rhododendron, Rhus, Rorippa nasturtium-aquaticum, Rosa, Rubus, Salia,
Senecio, Solidago, Sorbaria, Spiraea, Symphoricarpos, Taenidia, Tamarix, Taraxacum,
Thaspium, Trifolium repens, Vaccinium, Viburnum, Zizia.

Andrena crataegi Robertson, 1893. Amer. Ent. Soc., Trans. 20: 273. 2, 6.

Andrena subcommoda Cockerell, 1902. Canad. Ent. 34: 45. 2.

Andrena angusi Viereck, 1907. Ent. News 18: 284. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190. 2, 6 (key). —Atwood, 1934.
Canad. Jour. Res. 10: 208, 210. 2, ¢ (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 206-209, figs. 37-39, table 5 (redescription, synonymy). —LaBerge, 1967. Nebr.
Univ. State Mus., Bul. 7: 294 (subgeneric assignment, angusi). —LaBerge, 1969. Amer.
Ent. Soc., Trans. 95: 5-12, figs. 1, 24-28 (redescription).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 33-34 (nest). —Clements and Long, 1923.
Carnegie Inst. Wash., Pub. 326: 249 (ecology). —Brittain, 1933. Canada Dept. Agr., Bul.
162: 94, figs. (floral relationships). —LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 11, table 1
(floral relationships).

Superfamily APOIDEA 1821

fracta Casad and Cockerell. West. Tex. to south Calif.; Mexico (Distrito Federal, Puebla,
Zacatecas). Pollen: Polylectic, visits flowers of Acacia, Argemone, Baileya, Cryptantha
intermedia, Descourainia sophia, Dithyrea, Encelia farinosa, E'schscholzia, Fallugia,
Geraea canescens, Lepidium, Lesquerella, Medicago, Oenothera, Opuntia, Phacelia,
Physaria, Prosopis, Prunus, Pyrus, Salix, Senecio.
Andrena fracta Casad and Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 84. 6, 9.

Taxonomy: LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 30-34, figs. 14-18 (redescription).

Biology: LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 34, table 3 (floral relationships).
mellea Cresson. Tex., N. Mex., Ariz. Pollen: Unknown, but visits flowers of Argemone,
Asclepias subverticillata, Baccharis glutinosa, Fallugia paradoxa, Melilotus alba, Salix.
Andrena mellea Cresson, 1868. Amer. Ent. Soc., Trans. 1: 384. 9.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48 (key). —Cockerell, 1931.
Amer. Mus. Novitates 458: 16. 2 (key). —LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 16-18,
figs. 19-23 (redescription).

prunorum prunorum Cockerell. Sask., west to B. C., south to west. Tex., N. Mex., Ariz. and

Calif.; Mexico (Baja California). Pollen: Polylectic, visits a wide variety of flowers
including Acer, Actinea, Adenostoma, Amsinckia douglasiana, Arabis, Arctostaphylos,
Argemone, Baccharis glutinosa, B. viminea, Barbarea orthoceras, Berberis, Brassica
campestris, B. incana, B. nigra, Calochortus venustus, Calyptridium umbellatum,
Ceanothus integerrimus, Cercocarpus betuloides, Chaenactis glabriuscula, Chamaebatia
foliolosa, Chrysothamnus, Clarkia biloba, C. rhomboidea, Clematis, Cleome serrulata,
Crataegus, Crepis, Cryptantha intermedia, Cymopteris, Encelia farinosa, Eriastrum
virgata, Erigeron, Eriodictyon trichocalyx lanatum, Eriogonum fasciculatum, E.
nudum, Eriophyllum, Erysimum asperum, Eschscholzia californica, Fallugia
paradoxa, Gilia capitata, G. davyi, Hackelia, Helianthus, Heracleum, Hyptis emoryi,
Lappula, Larrea tridentata, Lepidium flavum, Lesquerella, Linaria, Lithospermum,
Lomatium, Malus, Melilotus, Mentzelia, Micrampelis, Monardella lanceolata,
Nemophila, Opuntia, Pastinaca, Penstemon, Petalostemon, Phacelia distans, P. minor,
Philadelphus lewisii californica, Platystemon, Plumbago, Potentilla, Prosopis, Prunus,
Purshia, Ranunculus, Raphanus sativa, Rhamnus californicus, Rhus, Ribes, Rorippa,
Rubus, Salix, Sambucus caerulea, Sanicula nevadensis, Sophia, Sphaeralcea ambigua,
Shepherdia, Sisymbrium irio, S. linifolia, Solidago, Spiraea, Symphoricarpos, Syringa,
Tamarix gallica, Taraxacum officinale, Thysanocarpus curvipes, Trichostema parishii,
Trifolium.

Andrena prunorum Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 81. 2, 6.

Andrena kincaidii Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 351. 9.

Andrena pascoensis Cockerell, 1897. Entomologist 30: 305. °.

Andrena prunorum gillettei Cockerell, 1898. Ent. News 9: 172. ¢.

Andrena vernoni Viereck, 1904. Canad. Ent. 36: 190, 195. 9, 6.

Andrena arizonensis Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 42. 6.

Andrena prunorum var. mariformis Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 46. °.

Andrena (Andrena) shasta Viereck, 1926. Calif. Acad. Sci., Proc. (4) 15: 402. 2.

Andrena prunorum var. pauperatula Cockerell, 1938. Canad. Ent. 70: 6. °.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 190, 195. 2, d (key, as kincaidii). —Cockerell,
1914. Ann. and Mag. Nat. Hist. (8) 14: 11. @ (key). —Cockerell, 1931. Amer. Mus. Novitates
458: 16. 2, d (key). —Lanham, 1941. Ent. Soc. Amer., Ann. 34: 704, 707. 2, d (key).
—LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 18-30, figs. 2-3, 9-13 (redescription,
synonymy, integumental color variation).

Biology: LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 28-30, table 2 (floral relationships).

prunorum sinaloa Viereck. Ariz.; Mexico (Baja California and Sinaloa). Pollen: Presumably

polylectic, visits flowers of Encelia farinosa, Larrea tridentata.
Andrena sinaloa Viereck, 1926. Calif. Acad. Sci., Proce. (4) 15: 403. °.

Taxonomy: LaBerge, 1969. Amer. Ent. Soc., Trans. 95: 30 (redescription).

Hymenoptera in America North of Mexico

1822
Genus ANDRENA Subgenus PTILANDRENA Robertson

Ptilandrena Robertson, 1902. Amer. Ent. Soc., Trans. 28: 192.
Type-species: Andrena erigeniae Robertson. Orig. desig.

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 215 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 283, 290, 307-308 (tax. characters). —Davis and LaBerge,

1975. Ill. Nat. Hist. Survey, Biol. Notes 95: 2 (included spp.).
distans Provancher. Canada; Minn. to New England, south to N. C. Pollen: Unknown, but
visits flowers of Geranium maculatum, Heracleuwm lanatum, Polemonium reptans.
Andrena distans Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 307. 2, d.
Andrena g. maculati Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 333. 2°, 6. Trinomial.
Andrena gerani-maculati Viereck, 1907. Ent. News 18: 282. °.
Taxonomy: Viereck, 1907. Ent. News 18: 282, 286. 2, 5 (key). —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 154-155, figs. 27, 29-30, table 4 (redescription, synonymy).
erigeniae Robertson. Ont. to Ga., west to Minn., Mo. and Okla. Parasite: Lewcophora obtusa
(Zett.), Stylops erigeniae Pierce. Pollen: Apparently collects pollen only from the flowers
of Claytonia virginica with which it is largely coterminous, but visits these and other
flowers for nectar including Barbarea vulgaris, Collinsia verna, Dicentra laciniata, D.
cucullaria, Erigenia bulbosa, Erythronium americanum, Hydrophyllum appendiculata,

Isopyrum biternatum, Prunus, Veronica hederaefolia.
Andrena erigeniae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 52. 2, 6.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 192. 9, d (key). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 155-156, figs. 27, 29-31, table 4 (redescription). — Davis
and LaBerge, 1975. Ill. Nat. Hist. Survey, Biol. Notes 95: 13, figs. 13, 16-24 (immature

stages).
Biology: Davis and LaBerge, 1975. Ill. Nat. Hist. Survey, Biol. Notes 95: 1-16, figs. 1-25, 1
table (nest, architecture, life history, provisioning, inquilinism, parasitism).
polemonii Robertson. IIl., Mich. Pollen: Unknown, but visits flowers of Geraniwm maculatum,

Polemonium reptans, Ranunculus septentrionalis.
Andrena polemonii Robertson, 1891. Amer. Ent. Soc., Trans. 18: 54. 2, ¢.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 192. 2, d (key). —Robertson
1904. Canad. Ent. 36: 278. (tax. characters). — Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.

Bul. 141: 157-158, figs. 27, 31, table 4 (redescription).
Genus ANDRENA Subgenus RHAPHANDRENA LaBerge
Andrena subg. Rhaphandrena LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 507.
Type-species: Andrena prima Casad and Cockerell. Orig. desig.
Revision: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 507-514, figs. 1, 3, 6, 59-68 (N. Amer.

spp.).
dapsilis LaBerge. Tex., Okla. Pollen: Unknown, but visits flowers of Descourainia pinnata

brachycarpa, Lesquerella.
Andrena (Rhaphandrena) dapsilis LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 511, figs. 6,

64-68. 2, d.

prima Casad. West. Tex. to south. Calif. and north. Utah. Pollen: Apparently an oligolege of
the brassicaceous flowers, especially Lesquerella including L. gordonii, but visits other
flowers presumably for nectar including Alyssum, Brassica oleracea, Cryptantha
probably jamesui, Dithyrea wislizenti, Geraea canescens, Prunus.

Andrena prima Casad, 1896. Ann. and Mag. Nat. Hist. (6) 18: 78. 9.
Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48. 2 (key). —Linsley and
MacSwain, 1961. Pan-Pacifie Ent. 37: 129 (subgeneric assignment).

Genus ANDRENA Subgenus SCAPHANDRENA Lanham

Andrena subg. Scaphandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 200.
Type-species: Andrena montrosensis Viereck and Cockerell. Orig. desig.

Superfamily APOIDEA 1823

Andrena subg. Elandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 203.
Type-species: Andrena amplificata Cockerell. Orig. desig.
Andrena subg. Truncandrena Warncke, 1968. Est. Mus. Zool. Coimbra Univ., Mem. 307:
46.

Type-species: Andrena truncatilabris Morawitz. Orig. desig.

Revision: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 101-189, 99 figs., 5 tables (N. Amer.
spp.).

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 296 (tax. characters). —Ribble,
1973. Ent. Soc. Amer., Ann. 66: 1281-1286 (tax. status of type species).
albiculta Viereck. Calif.

Andrena albiculta Viereck, 1917. Amer. Ent. Soc., Trans. 43: 366. 9.

Taxonomy: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 173-174 (tax. status).

arabis Robertson. N. Y. to N. C., west to Mich. and Ill. Pollen: Polylectic, often collected on
early spring flowering plants of the family Cruciferae; visitation records include Arabis
laevigata, Barbarea vulgaris, Brassica, Buxus, Capsella bursa-pastoris, Cardamine
bulbosa, Claytonia virginica, Dentaria diphylla, D. lanciniata, Erigenia bulbosa,
Erythronium americanum, Prunus, Salix microphylla, Spiraea, Stellaria media,
Taraxcum officinale, Tussilago farfara. Hybridizes with A. scurra Viereck and A.
capricornis Casad and Cockerell.

Andrena arabis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 334. 2.

Andrena (Pterandrena) brassicae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 142,
figs. 24, 25, 28. ¢.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191. 2 (key). —Viereck, 1907. Ent.
News 18: 282. ? (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 251-252, figs.
54-56, table 7 (redescription). —Ribble, 1973. Ent. Soc. Amer., Ann. 66: 1281-1286
(hybridization). —Ribble, 1974. Amer. Ent. Soc., Trans. 100: 115-118, figs. 1, 14-18
(redescription, synonymy).

Biology: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 118, table 1 (floral relationships).
bruneri Viereck and Cockerell. Nebr., Wyo., Colo., Nev. Pollen: Unknown, but visits flowers of
Astragalus, Erysimum.
Andrena bruneri Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 15. °.
Andrena laramiensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 22, 23. d.
Andrena viridibasis Cockerell, 1930. N. Y. Ent. Soc., Jour. 37: 444. °.

Taxonomy: Cockerell, 1931, Amer. Mus. Novitates 458: 9. —Timberlake, 1951. Jn Linsley, Jn
Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2: 1059 (synonymy).
—Ribble, 1974. Amer. Ent. Soe., Trans. 100: 161-164, figs. 1, 80-84 (redescription,
synonymy).

capricornis Casad and Cockerell. Tex. to south. Calif. Parasite: Stylops sp. Pollen: Apparently
polylectic, often collected at the flowers of the family Cruciferae; visitation records
include Actinea, Alyssum maritimum, Brassica oleracea, Descourainia sophia,
Lesquerella gordonii, Physaria, Prunus, Salvia. Hybridizes with A. scurra Viereck and
A. arabis Robertson.
Andrena capricornis Casad and Cockerell, 1896. Canad. Ent. 28: 182. ¢.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48. d (key). —Cockerell, 1899.
Ent. News 10: 255. ¢ (key). —Ribble, 1973. Ent. Soc. Amer., Ann. 66: 1281-1286
(hybridization). —Ribble, 1974. Amer. Ent. Soc., Trans. 100: 120-122, figs. 1, 19-23
(redescription).

chapmanae Viereck. Calif., Oreg., Idaho, Nev. and Utah. Pollen: Unknown, but visits flowers

of Ceanothus, Chamaebatia foliolosa, Clarkia dudleyana, Cryptantha ambigua, C.
flaccida, C. intermedia, C. micrantha var. lepida, Descourainia sophia, Gilia capitata,
G. latiflora var. cana, Hackelia patens, Lupinus, Mentzelia, Nemophila maculata,
Phacelia linearis, Plagiobothrys canescens, P. nothofulvus, Prunus andersonii, Salix,
Senecio canus.

Andrena (Andrena) chapmanae Viereck, 1904. Canad. Ent. 36: 191. °.

Andrena yosemitensis Cockerell, 1924. Pan-Pacifie Ent. 1: 61. °.

1824 Hymenoptera in America North of Mexico

Taxonomy: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 166-170, figs. 1, 8, 90-94 (redescription).

Biology: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 168, table 5 (floral relationships).
cruciferarum Ribble. Calif. Pollen: Unknown, but visits flowers of Amsinckia, Brassica,
Capsella bursa-pastoris, Phacelia, Sisymbrium irio.
Andrena (Scaphandrena) cruciferarum Ribble, 1974. Amer. Ent. Soc., Trans. 100: 128, figs.
1-2, 29-33. 2, 3.
gordoni Ribble. Northeast. Calif., Oreg., Wash., Wyo.
Andrena (Scaphandrena) gordoni Ribble, 1974. Amer. Ent. Soce., Trans. 100: 134, figs. 1-2,
39-43. 2, 3.
hicksi Cockerell. Colo., Wyo., Utah and Nev. (Ruby Mts.). Parasite: Stylops sp.
Andrena hicksi Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 628. °.
Andrena unicula Cockerell, 1934. Pan-Pacific Ent. 9: 155. 6.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 265. 2 (key). —Lanham, 1941. Ent.
Soc. Amer., Ann. 34: 705, 707. 2, ¢ (key). —Lanham, 1941. Ent. Soc., Amer., Ann. 34: 707.
3 (key, as unicula). —Ribble, 1974. Amer. Ent. Soc., Trans. 100: 164-166, figs. 1, 6, 85-89
(redescription, synonymy).
Biology: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 165 (stylopization).
kaibabensis Ribble. Ariz., Utah.
Andrena (Scaphandrena) kaibabensis Ribble, 1974. Amer. Ent. Soc., Trans. 100: 160, figs.
1a: oe
lomatii Ribble. Calif. Pollen: Unknown, but visits flowers of Lomatiwm dasycarpum, Rhus
trilobata, Salvia sonomensis, Sanicula nevadensis.
Andrena (Scaphandrena) lomatii Ribble, 1974. Amer. Ent. Soc., Trans. 100: 131, figs. 1, 5,
34-38. 2, 3.
mackieae Cockerell. South. Oreg., Calif., Nev., Utah, Ariz.; Mexico (Baja California). Pollen:
Unknown, but visits flowers of Amsinckia, Arctostaphylos, Brassica, Ceanothus
cordulatus, C. crassifolius, C. cuneatus, C. integerrimus, C. verrucosus, Cryptantha,
Phacelia distans, Plagiobothrys, Prunus virginiana, Rhamnus crocea, Rhus diversiloba,
Salix lasiolepis.
Andrena mackieae Cockerell, 1937. Amer. Mus. Novitates 899: 3. 9.
Andrena (?Scaphandrena) mackiae(!) Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 201.

Taxonomy: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 144-149, figs. 1, 6, 59-63 (redescription).

Biology: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 147, table 3 (floral relationships).
merriami Cockerell. B. C. to Calif., east to Mont., Nebr. Colo. and N. Mex. Pollen: Unknown,
but visits flowers of Acer, Astragalus, Cardaria draba, Claytonia, Daucus carota,
Fraxinus, Gilia capitata, Hackelia patens, Lappula floribunda, Leptotaenia,
Lithophragma affinis, Lomatium ambiguum var. leptocarpum, L. dissectum, L.
orientale, L. triternatum, Phacelia, Prunus demissa, P. virginiana, var. melanocarpa,
P. subcordata, Pyrus, Ranunculus, Salix, Taraxacum officinale, Thelypodium.
Andrena merriami Cockerell, 1901. Psyche 9: 284. 2.
Andrena pullmani Viereck, 1904. Canad. Ent. 36: 191, 195. 2, o.
Andrena amplificata Cockerell, 1910. Canad. Ent. 42: 368. 2, d.
Andrena fulvinigra Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 42. 9.

Taxonomy: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 150-155, figs. 1, 3, 64-69 (redescription,
synonymy).
Biology: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 153, table 4 (floral relationships).
mohavensis Ribble. South. Calif. (Mohave Desert). Pollen: Unknown, but visits flowers of

Lepidium flavum, Phacelia, Salix exigua.
Andrena (Scaphandrena) mohavensis Ribble, 1974. Amer. Ent. Soc., Trans. 100: 126, figs.
1, 2, 24-28. 9, 6.
plana Viereck. Oreg., Calif. Pollen: Apparently oligolectie on Trifoliwm including T.
involucratum, T. microcephalum, T. pratense, T. repens, T. tridentatum, T. variegatum,
but visits other flowers presumably for nectar including Brassica, Chamaebatia

Superfamily APOIDEA = 1825

foliolosa, Cryptantha, Gilia, Lasthenia chrysostoma, Layia platyglossa var. breviseta,
Limnanthes douglasii, Nemophila, Orthocarpus faucibarbatus, Plagiobothrys
nothofulvus, Ranunculus, Rhus trilobata, Sisyrinchium bellum, Vicia.

Andrena plana Viereck, 1904. Canad. Ent. 36: 193. °.

Taxonomy: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 141-144, figs. 1, 7, 54-58 (redescription).
primulifrons Casad. Tex. to Ariz. and south. Nev. (Hoover Dam). Pollen: Possibly an oligolege
of Lesquerella including L. densiflora, L. gordonii, L. ludoviciana, but visits other
flowers including Descourainia pinnata, Dithyraea wislizenii, Larrea tridentata,

Lobularia, Machaeranthera tanacetifolia, Oreocarya, Phacelia crenulata, Physaria,
Prunus, Sisymbrium.
Andrena primulifrons Casad, 1896. Canad. Ent. 28: 183. ¢, 2.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48, 49. 3d, 2 (key). —Ribble,
1968. Nebr. Univ. State Mus., Bul. 8: 328-332, figs. 85-87, map 10, table 11 (redescription).
—Ribble, 1974. Amer. Ent. Soc., Trans. 100: 104, fig. 1 (subgeneric position).

santaclarae Ribble. Calif. Pollen: Unknown, but males visit flowers of Cryptantha, Lasthenia,
Nemophila, Prunus subcordata.
Andrena (Scaphandrena) santaclarae Ribble, 1974. Amer. Ent. Soc., Trans. 100: 136, figs.
1, 4, 44-48. 2, d.
scurra Viereck. B. C. to Calif., east to Idaho and north. Utah. Pollen: Unknown, but visits
flowers of Achillea lanulosa, Baccharis, Brassica campestris, Capsella bursa-pastoris,
Cardaria draba, Ceanothus greggii, C. palmeri, Cosmos, Cotoneaster, Descourainia
sophia, Lepidium, Lomatium dissectum, L. triternatum, Medicago sativa, Prunus
subcordata, Salix, Scrophularia californica, Sisymbrium altissimum, Spiraea,
Taraxacum, Thlaspi arvense. Hybridizes with A. arabis Robertson and A. capricornis
Casad and Cockerell in the Great Basin area of the western United States and in the
Rocky Mountain area.
Andrena scurra Viereck, 1904. Canad. Ent. 36: 193, 195. 2, ¢.

Taxonomy: Ribble, 1973. Ent. Soc. Amer., Ann. 66: 1281-1286 (hybridization). —Ribble, 1974.
Amer. Ent. Soc., Trans. 100: 110-115, figs. 1, 9-13 (redescription).
scurra x arabis x capricornis Hybrids. Idaho, Wyo., Colo., Utah, Nev. and along the east slope
of the Rocky Mountains from north. N. Mex. to south. Alta; Calif. (Seven Oaks) and
Ariz. (Fredonia). Pollen: Apparently polylectic, exhibits a preference for the flowers of
the family Cruciferae, but visits a variety of flowers including Agoseris, Astragalus,
Brassica, Cardaria draba, Cleome serrulata, Euphorbia, Lappula floribunda, Lepidium
virginicum var. medium, Lesquerella ovalifolia, Oxytropis, Philadelphus, Physaria
didymocarpa, Prunus, Ribes, Senecio canus, Sisymbrium altissimum, S. pinnatum,
Taraxacum officinale.
Andrena sieverti Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 436. 2.
Andrena lappulae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 437. ¢.
Andrena ellisiae Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 9. 2.
Andrena montrosensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 48. 9.
Andrena speculifera Cockerell, 1930 (1929). N. Y. Ent. Soc., Jour. 37: 444. 9.
Andrena physariae Cockerell, 1934. Pan-Pacifie Ent. 9: 154. 9.
Andrena sieverti var. opacicauda Cockerell, 1936. Amer. Mus. Novitates 831: 1. 9.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 13. 2 (key, as sieverti). —Lanham,
1941. Ent. Soc. Amer., Ann. 34: 706. 2 (key, as ellisiae). —Lanham, 1941. Ent. Soc. Amer.,
Ann. 34: 706. 2 (key, as physariae). —Ribble, 1973. Ent. Soc. Amer., Ann. 66: 1281-1286
(hybridization). —Ribble, 1974. Amer. Ent. Soc., Trans. 100: 122-126 (synonymy).

Biology: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 124, table 2 (floral relationships,
stylopization).

shoshoni Ribble. Wyo. (Big Horn and Foxpark).
Andrena (Scaphandrena) shoshoni Ribble, 1974. Amer. Ent. Soc., Trans. 100: 172, figs. 1,

ore.

1826 Hymenoptera in America North of Mexico

sladeni Viereck. B. C. to Wyo. and Nev. Pollen: Unknown, but visits flowers of Taraxacum
officinale.
Andrena (Andrena) sladeni Viereck, 1924. Canad. Ent. 56: 239. @.
Taxonomy: Cockerell, 1932. Canad. Ent. 64: 286 (key). —Ribble, 1974. Amer. Ent. Soc., Trans.
100: 155-158, figs. 1, 4, 70-74 (redescription).
tildeni Ribble. Calif., Ariz. Pollen: Unknown, but visits flowers of Cryptantha intermedia,
Salix argophylla, S. exigua.
Andrena (Scaphandrena) tildeni Ribble, 1974. Amer. Ent. Soc., Trans. 100: 139, figs. 1, 3,
49-53. do.
trapezoidea Viereck. Tex., Okla. Pollen: Unknown, but visits flowers of Descourainia pinnata,
Lesquerella densiflora, L. gordonii, L. gracilis, L. recurvata, Rhus microphylla.
Andrena (Scrapter) trapezoidea Viereck, 1917. Amer. Ent. Soc., Trans. 43: 402. °.
Taxonomy: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 328-332, figs. 85-87, map 10, table 11
(redescription). —Ribble, 1974. Amer. Ent. Soc., Trans. 100: 104, fig. 1 (subgeneric
position).
vestali Cockerell. Colo., Nebr. Pollen: Unknown, but visits flowers of Viola.
Andrena vestali Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 64. o.
Andrena vestali var. dolichocera Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48:
195G-
Taxonomy: Lanham, 1941. Ent. Soc. Amer., Ann. 34: 707. ¢ (key). —Ribble, 1974. Amer. Ent.
Soc., Trans. 100: 170-172, figs. 1, 8, 95-99 (redescription).
walleyi Cockerell. B. C. to Calif. (Alpine Co.), east to Idaho and Utah. Pollen: Unknown, but
visits flowers of Cardaria draba.
Andrena walleyi Cockerell, 1932. Canad. Ent. 64: 286. 2.

Taxonomy: Ribble, 1974. Amer. Ent. Soc., Trans. 100: 158, figs. 1, 5, 75-79 (redescription).

Genus ANDRENA Subgenus SCOLIANDRENA Lanham

Andrena subg. Scoliandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 223.
Type-species: Andrena osmioides Cockerell. Monotypic and orig. desig.

The species of this subgenus chiefly visit the flowers of Cryptantha upon which the females
are apparently solely dependent for pollen.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 283, 291, 313 (tax. characters).

cryptanthae Timberlake. South. Calif. Pollen: Oligolege of Cryptantha including C. barbigera,
C. intermedia, but visits other flowers for nectar.
Andrena (Scoliandrena) cryptanthae Timberlake, 1951. U. S. Natl. Mus. Proc. 101: 401. 2,
é.
osmioides benitonis Lanham. North. Calif. Pollen: Oligolege of Cryptantha.
Andrena osmioides benitonis Lanham, 1949. Pan-Pacifie Ent. 25: 148. 9, do.
osmioides osmioides Cockerell. South. Calif. Pollen: Oligolege of Cryptantha including C.
intermedia, but visits other flowers for nectar including Plagiobothrys californica, P.
nothofulvus.
Andrena osmioides Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 45. 6.

Genus ANDRENA Subgenus SCRAPTEROPSIS Viereck

Andrena subg. Scrapteropsis Viereck, 1922. Boston Soc. Nat. Hist., Occas. Papers 5: 42.
Type-species: Andrena (Scrapteropsis) fenningeri Viereck. Monotypic.
Andrena subg. Mimandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 217.
Type-species: Andrena imitatrix Cresson. Orig. desig.
Revision: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 441-504, figs. 1-53, 1 table (N. Amer.
Spp.).
Taxonomy: Warncke, 1968. Coimbra Univ. Est. Mus. Zool., Mem. 307: 68-69 (synonymy).
—LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 235, 237-240 (phylogeny, tax. status).

Superfamily APOIDEA 1827

alamonis Viereck. West. Tex., N. Mex., Ariz. Pollen: Unknown, but visits flowers of
Lesquerella.

Andrena (Scrapter) alamonis Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 599. 2.
alleghaniensis Viereck. Ont. and Maine to S. C. west to Sask. and Colo. Pollen: Apparently
polylectic, visits flowers of Amelanchier, Barbarea, Brassica, Chrysanthemum
leucanthemum, Cornus, Crataegus, Euphorbia cyparissias, Fragaria vesca, F.
virginiana, Ledum, Melilotus alba, Prunus, Ranunculus, Rubus idaeus, Salix,

Viburnum opulus.
Andrena (Trachandrena) alleghaniensis Viereck, 1907. Ent. News 18: 280, 286. 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 177 (redescription). —Knerer
and Atwood, 1963. Canad. Ent. 95: 583, figs. 1, 3. d. —Knerer and Atwood, 1964. Ent. Soc.
Ontario, Proc. 94: 47-48. —LaBerge, 1971. Ent. Soc. Amer., Trans. 97: 474-479, figs. 1, 5,
27-31 (redescription).

angusticrus LaBerge. Calif. Pollen: Unknown, but visits flowers of Lasthenia chrysostoma,
Platystemon californicus.
Andrena (Scrapteropsis) angusticrus LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 499, figs.
Grae.
aquila LaBerge Calif. (Fresno County). Pollen: Unknown, but visits flowers of Platystemon
californicus.
Andrena (Scrapteropsis) aquila LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 502, figs. 1, 6.
2

atlantica Mitchell. N. J. to Fla., west to Miss. Pollen: Apparently polylectic, visits flowers of
Crataegus, Diospyros virginiana, Fragaria, Ilex glabra, Ligustrum, Linaria, Lyssa
sylvatica, Polycodium, Pyracantha, Rubus, Salix nigra, Senecio, Spiraea,
Toxicodendron.
Andrena (Trachandrena) atlantica Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
178, figs. 35-37. 2, ¢.

Taxonomy: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 479-481, figs. 1, 5 (redescription).
biareola LaBerge. Calif. Pollen: Unknown, but visits flowers of Cryptantha, Meconella,
Platystemon californicus.
Andrena (Scrapteropsis) biareola LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 494, figs. 1, 3.
Oy Ore
buccata LaBerge. Calif. Pollen: Unknown, but visits flowers of Blennosperma nana,
Lasthenia, Platystemon.
Andrena (Scrapteropsis) buccata LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 497, figs. 1, 6,
49-53. 2, d.
daeckei Viereck. N. J., Pa.
Andrena (Trachandrena) daeckei Viereck, 1907. Ent. News 18: 280, 285, 286. °, d.

Taxonomy: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 486-489, figs. 1, 4, 37-41 (redescription,
distribution).
fenningeri Viereck. Mass. to Ga., west to IIl., Tenn. and Tex. Pollen: Polylectic, with a strong
preference for Salix including S. humilis, S. sericea, S. tristis and a secondary
preference for Prunus including P. americanus, P. persica, but visits other flowers
including Acer rubrum, Alsine media, Amelanchier, Aronia, Pyrus malus.
Andrena (Scrapteropsis) fenningeri Viereck, 1922. Boston Soc. Nat. Hist., Occas. Papers 5:
42, figs. 2, 3.
Andrena (Trachandrena) indianensis Cockerell, 1929. Ent. Soc. Amer., Ann. 22: 757. 2.
Andrena (?Mimandrena) verna Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 206,
figs. 37, 38. 6.
Taxonomy: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 465-470, figs. 1, 6, 17-21 (redescription,
synonymy).
flaminea LaBerge. Tex. (Beeville and Edinburg).
Andrena (Scrapteropsis) flaminea LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 472, fig. 1.
OF

1828 Hymenoptera in America North of Mexico

ilicis Mitchell. N. Y. to north. Fla., west to Kans. and Tex. Pollen: Unknown, but visits flowers
of Batodendron, Castanea pumila, Celastrus, Crataegus, Diospyros virginiana, Ilex,
Prunus, Rhus glabra, Salix.
Andrena (Trachandrena) ilicis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 186,
fig. 35. 2.

Taxonomy: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 459-463, figs. 1, 3, 12-16
(redescription).
imitatrix Cresson. Que. to Fla. west to Alta., Mont., Utah and Ariz. Pollen: Polylectic, although

exhibits a strong preference for the flowers of the Rosaceae and Salicaceae; visitation
records include Acer negundo, Alsine media, Amelanchier arborea, A. canadensis,
Amorpha fruticosa, Angelica atropurpurea, Antennaria dioicum, Astragalus, Barbarea
vulgaris, Benzoin aestivale, Cassia fasciculata, Ceanothus ovatus, Claytonia, Cornus,
Crataegus mollis, C. occidentalis, Dentaria laciniata, Descourainia intermedia,
Diospyros virginiana, Erythronium, Euphorbia, Exochoroa racemosa, Forsythia,
Grossularia, Heracleum, Ilex, Laurocerasus caroliniana, Lepargyraea canadensis,
Lomatium daucifolium, L. foeniculaceum, Lonicera morrowi, Medicago sativa,
Pastinaca sativa, Paeonia, Phacelia, Physocarpus opulifolius, Populus, Prunus
americana, P. gracile, P. pennsylvanica, P. serotina, P. virginiana, Ptelea, Pyracantha,
Pyrus malus, Rhamnus utilis, Rhus aromatica, R. canadensis, Ribes gracile, Rosa,
Rubus, Salix amygdaloides, S. babylonica, S. bebbiana, S. discolor, S. longifolia, S.
nigra, Senecio, Spiraea aruncus, S. thunbergi, S. vanhouttei, Tamarix gallica,
Taraxacum officinale, Thlaspi arvense, Vaccinium, Viburnum dentatum, V.
prunifolium, Zygadenia, Zizia.

Andrena imitatrix Cresson, 1872. Amer. Ent. Soc., Trans. 4: 258. 2.

Andrena claytoniae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 59. 2, 6.

Trachandrena albofoveata Graenicher, 1903. Canad. Ent. 35: 166. 2.

Andrena johnsoniana Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 224. 9.

Andrena (Trachandrena) crataegiphila Viereck and Cockerell, 1914. U. S. Natl. Mus.,
Proc. 48: 7. ¢.

Andrena (Trachandrena) titusi Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 12.
On

Andrena (Scrapter) imitatrix var. profunda Viereck, 1917. Amer. Ent. Soc., Trans. 43: 398.

oO.
Andrena (Trachandrena) tardula Cockerell, 1930. N. Y. Ent. Soc., Jour. 37: 447. 2.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190. 2, 6 (key). —Cockerell, 1929.
Ent. Soc. Amer., Ann. 22: 755 (tax. characters). —Cockerell, 1931. Canad. Ent. 68: 22. 2 (as
johnsoniana). —Cockerell, 1931. Amer. Mus. Novitates 458: 11. 2 (as johnsoniana).
—Lanham and Timberlake, 1951. Jn Muesebeck, Krombein and Townes, U. S. Dept. Agr.,
Agr. Monog. 2: 1068 (synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
205-206, fig. 37-39, table 5 (redescription, synonymy). —Yager and Rozen, 1966. Amer.
Mus. Novitates 2265: 6 (pupa). —LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 451-459, figs.
1-2, 7-11 (redescription, synonymy).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 34 (nesting habits). —Clements and Long,
1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology). —LaBerge, 1971. Amer. Ent. Soc.,
Trans. 97: 457-459, table 1 (floral relationships).

kalmiae Atwood. N. S., Que. and Ont. south to Mass. and Conn. Pollen: Apparently an
oligolege of Kalmia including K. angustifolia, but visits other flowers presumably for
nectar including Ledum, Lyonia ligustrina, Vaccinium.
Andrena kalmiae Atwood, 1934. Canad. Jour. Res. 10: 210, fig. 2, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 186, fig. 35, table 5
(redescription). —LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 489-491, figs. 1, 4
(redescription, distribution).

morrisonella Viereck. Ont. to Fla., west to Ill. and Ala. Pollen: Unknown, but visits flowers of
Amelanchier, Crataegus, Gypsophila, Ilex, Malus, Populus, Prunus, Ranunculus acris,
Rubus, Salix.
Andrena (Scrapter) morrisonella Viereck, 1917. Amer. Ent. Soc., Trans. 43: 399. 9.

Superfamily APOIDEA = 1829

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 192, fig. 35, table 5
(redescription). —LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 463-465, fig. 1 (redescription).

phenax Cockerell. South. Calif.
Andrena phenax Cockerell, 1898. Amer. Ent. Soc. Trans. 25: 188. °.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 129 (tax. status). —Thorp, 1969.
Calif. Univ. Pubs. Ent. 52: 121-123 (tax. status). —LaBerge, 1971. Amer. Ent. Soc., Trans.
97: 503-504, fig. 1 (redescription).
rubi Mitchell. N. C. to Ga., west to Tex., Okla., Kans. and Nebr. Pollen: Unknown, but visits
flowers of Aronia, Crataegus, Descourainia intermedia, D. pinnata, Prunus americana,
P. gracilis, Rubus, Salix nigra, Spiraea.
Andrena (Trachandrena) rubi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 198, fig.
35. 9.
stipator LaBerge. Calif. Pollen: Unknown, but visits flowers of Cryptantha, Meconella,
Platystemon californicus.
Andrena (Scrapteropsis) stipator LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 491, figs. 1, 3,
44-48. 2, 6.
unicostata LaBerge. Tex., Okla. Pollen: Unknown, but visits flowers of Salix.
Andrena (Scrapteropsis) unicostata LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 481, figs. 1,
5, 32-36: 9, d-

Genus ANDRENA Subgenus SIMANDRENA Perez

Simandrena Perez, 1890. Soc. Linn. Bordeaux, Actes. 44: 174.
Type-species: Andrena propinqua Schenck. Desig. by Hedicke, 1933.
Andrena subg. Platandrena Viereck, 1924. Canad. Ent. 56: 21.
Type-species: Andrena vestita Provancher. Monotypic and orig. desig. (=Andrena
nasonii Robertson)
Andrena subg. Stenandrena Timberlake, 1949. In Lanham, Calif. Univ. Pubs. Ent. 8: 213.
Type-species: Andrena pallidifovea (Viereck). Orig. desig.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 285, 286, 293, 304 (tax. characters,
synonymy).
angustitarsata Viereck. Wash. to Calif. Pollen: Apparently polylectic, visits a wide variety of
flowers including Achillea lanulosa, A. millefolium, Arenaria douglasti, Ceanothus
cuneatus, C. greggii, C. integerrimus, C. leucodermis, C. orcuttu, C. palmeri, C.
soredatus, Chamaebatia foliolosa, Claytonia spatulata, Eriodictyon californicum,
Eriophyllum confertiflorum, Gilia capitata, Grossularia californica, Horkelia bolanderi,
Lasthenia chrysostoma, Linanthus aureus, Lomatium utriculatum, Paeonia californica,
Plagiobothrys nothofulvus, Potentilla glandulosa, Prunus ilicifolia, Ranunculus
californicus, Rhamnus californicus, R. crocea, Rhus diversiloba, Salix laevigata, S.
lasiandra, S. lasiolepis, Sanicula bipinnatifida, S. nevadensis, S. tuberosa, Saxifraga
integrifolia, Swertia parryi, Thysanocarpus curvipes.
Andrena angustitarsata Viereck, 1904. Canad. Ent. 36: 189, 196. 2.
Pterandrena nudiscopa Viereck, 1904. Canad. Ent. 36: 227, 228. °.
essigi Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Cryptantha intermedia,
Lasthenia aristata, L. gracilis, Layia elegans, Rhamnus crocea, Rhus trilobata,
Sisymbrium irio.
Andrena (Stenandrena) essigi Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 384. 9, ¢.
friesei Viereck. Calif. Ecology: Unknown, but visits flowers of Amsinckia douglasiana,
Brassica, Calandrinia menziesii, Eschscholzia californica, Lasthenia, Raphanus
sativus.
Andrena (Andrena) friesei Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 558. 9°.
huardi Viereck. Calif. Pollen: Possibly an oligolege of Salix including S. argophylla, S.
gooddingii, S. hindsiana, S. laevigata, S. lasiolepis, S. nigra, but visits other flowers
possibly for nectar only including Alyssum maritimum, Sambucus.
Andrena (Andrena) angustitarsata huardi Viereck, 1917. Amer. Ent. Soc., Trans. 43: 368.
2.

1830 | Hymenoptera in America North of Mexico

hypoleuca Cockerell. South. Calif. (Catalina Island).
Andrena hypoleuca Cockerell, 1939. Pomona Jour. Ent. Zool. 31: 25. °.
nasonii Robertson. Eastern Canada and Minn., south to Ga. and Tex., Colo. Pollen: Apparently
polylectic, visits a wide variety of flowers including Acer, Amelanchier, Apocynum,
Arabis, Capsella, Chaerophyllum, Claytonia, Crataegus, Forsythia, Fragaria, Ilex,
Ligustrum, Polemonium, Polytaenia, Potentilla, Prunus, Pyracantha, Ranunculus,
Rhamnus, Rubus, Senecio, Sisymbrium, Spiraea, Taenidia, Toxicodendron, Viburnum,
Viola, Zizia.
Andrena vestita Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 310. ¢.
Preoce.
Andrena nasonii Robertson, 1895. Amer. Ent. Soc., Trans. 22: 120. 2.
Andrena hartfordensis Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 103. 2.
Andrena (Andrena) nasonii var. fulvodorsata Viereck, 1917. Amer. Ent. Soc., Trans. 43:
385. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 192. 2, 6 (key). —Viereck,
1907. Ent. News 18: 283, 285. 2, 3 (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
141: 244-245, figs. 51-52, 54 (redescription).

opacella Timberlake. Calif. (Mt. Hamilton Range). Pollen: Unknown, but visits flowers of
Salix.
Andrena (Simandrena) opacella Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 406. °.
opacibasis Cockerell. Calif.
Andrena (Platandrena) opacibasis Cockerell, 1936. Pan-Pacific Ent. 12: 146. 9.
opacissima Cockerell. Idaho, Wash. to Utah and Calif. Pollen: Unknown, but visits flowers of
Ceanothus cordulatus, Cryptantha intermedia, Lomatium, Salix lasiolepis.
Andrena opacissima Cockerell, 1918. Ann. and Mag. Nat. Hist. (9) 1: 165. @.

Taxonomy: Cockerell, 1933. Pan-Pacific Ent. 9: 158. 2. —Cockerell, 1936. Ann. and Mag. Nat.
Hist. (10) 18: 632.
opaciventris Cockerell. Calif. Pollen: Unknown, but visits flowers of Ceanothus crassifolius, C.
integerrimus, C. vestitus, Cryptantha intermedia, Lomatium dasycarpum, Prunus,
Ranunculus occidentalis, Rhamnus crocea, Salix lasiolepis, Sanicula tuberosa, S.
bipinnatifida.
Andrena opaciventris Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 47. 9.
orthocarpi Cockerell. Calif. Pollen: Unknown, but visits flowers of Achillea millefolium,
Calycadenia multiglandulosa, Ceanothus cuneatus, C. soredatus, Limnanthes douglasii,
Lomatium utriculatum, Orthocarpus densiflorus, O. lithospermoides, Ranunculus
californicus, Salix lasiolepis, Phoradendron, Thysanocarpus curvipes.
Andrena (Platandrena) orthocarpi Cockerell, 1936. Pan-Pacific Ent. 12: 147. @.

Taxonomy: Lanham, 1947. Pan-Pacifie Ent. 23: 72. ¢.
pallidifovea (Viereck). B. C. to Calif., Utah. Pollen: Collects pollen from the flowers of
Compositae including Chaenactis, Ericameria cooperi, Eriophyllum, Helianthus
gracilentus, Layia, but visits other flowers for nectar including Gilia capitata,
Lasthenia chrysostoma.
Pterandrena pallidifovea Viereck, 1904. Canad. Ent. 36: 195, 227, 228. 6, 9.
Andrena plumifera Cockerell, 1916. Canad. Ent. 48: 393. 2.

Taxonomy: Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1075 (synonymy). —Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 383.
é.

pensilis Timberlake. Calif. Pollen: Unknown, but visits flowers of Eriodictyon californicum,
Eschscholzia californica, Lasthenia chrysostoma, Limnanthes douglasii, Layia
platyglossa, Orthocarpus densiflorus, O. lithospermoides, Ranunculus californicus,
Sisyrinchium bellum.
Andrena (Platandrena) pensilis Timberlake, 1938. Pan-Pacifie Ent. 14: 27. 9, 3.

runcinatae Cockerell. B. C. and Alta. to Colo., N. Dak.
Andrena runcinatae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 434. 9, é.

Superfamily APOIDEA 1831

uvulariae Mitchell. Md. (Beltsville). Pollen: Unknown, but visits flowers of Uvularia
sessilifolia.
Andrena (?Simandrena) uvulariae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
245, fig. 54. 9.
vexabilis Timberlake. North. Calif. Pollen: Unknown, but visits flowers of Achillea lanulosa,
Agoseris heterophylla, Chamaebatia foliolosa, Coreopsis maritima, Cryptantha
micrantha, Encelia californica, Eriophyllum confertiflorum, Lasthenia chrysostoma, L.
gracilis, Layia platyglossa, Helianthella californica, Horkelia fusca, Senecio
californicus, Viguiera, Wyethia angustifolia.
Andrena (Stenandrena) vexabilis Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 385. 2, ¢.
wheeleri Graenicher. N.S. and Maine to Alta., south to Colo., Minn., Wis. and N. C. Pollen:
Unknown, but visits flowers of Centaurea, Chrysanthemum leucanthemum, Cornus,
Rhododendron, Rubus, Sedum acre.
Andrena wheeleri Graenicher, 1904. Ent. News 15: 65. ?, 6d.
Andrena wheeleri pallidior Cockerell, 1938. Canad. Ent. 70: 7. °.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10: 208, 209. 2, ¢ (key). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 246-247, figs. 51-52 (redescription, synonymy).

Genus ANDRENA Subgenus TAENIANDRENA Hedicke

Andrena subg. Taeniandrena Hedicke, 1933. Berlin Zool. Mus., Mitt. 19: 219.
Type-species: Melitta ovatula Kirby. Orig. desig.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 286, 292, 306-307 (tax. characters).
wilkella (Kirby). Holarctic; N.S. to Wis., south to Va. and Ohio. Presumably introduced.

Pollen: Apparently polylectic, visits flowers of Achillea, Amorpha, Brassica, Cirsium,
Crataegus, Daucus, Epilobium, Fragaria, Leontodon, Malus, Philadelphus, Pinus,
Prunus, Raphanus, Rhododendron, Rhodura, Rubus, Solidago, Stellaria, Taraxacum,
Trifolium, Vaccinium.

Melitta wilkella Kirby, 1802. Monog. Apum Angliae, v. 2, p. 145. 2, d.

Andrena winkleyi Viereck, 1907. Ent. News 18: 288, 285. 2, 6.

Taxonomy: Malloch, 1918. Biol. Soc. Wash., Proc. 31: 61. —Atwood, 1934. Canad. Jour. Res.
10: 207, 210. 2, d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 250-251,
figs. 52, 54-55 (redescription).

Biology: Brittain, 1933. Canada Dept. Agr., Bul. 162: 94 (floral relationships). —Atwood, 1933.
Canad. Jour. Res. 9: 456 (floral relationships). —Brittain and Newton, 1933. Canad. Jour.
Res. 9: 341 (flower records). —Brittain and Newton, 1934. Canad. Jour. Res. 10: 261
(flower records).

Genus ANDRENA Subgenus THYSANDRENA Lanham

Andrena subg. Thysandrena Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 213.
Type-species: Andrena candida Smith. Orig. desig.

Taxonomy: LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 287, 2938, 294, 306 (tax. characters).
argentiscopa Viereck. Calif.
Andrena (Andrena) argentiscopa Viereck, 1917. Amer. Ent. Soc., Trans. 43: 368. 2.
beckeri Cockerell. Colo.
Andrena beckeri Cockerell, 1921. Ann. and Mag. Nat. Hist. (9) 7: 211. 2.

Taxonomy: Cockerell, 1932. Pan-Pacific Ent. 8: 175. —Lanham, 1941. Ent. Soc. Amer., Ann.
34: 706. @.
bisalicis Viereck. N. Dak. to New England states, south to Ga. Parasite: Nomada sp. Pollen:
Unknown, but visits flowers of Amelanchier, Aronia, Brassica, Crataegus, Ilex, Prunus,
Salix.
Andrena salicis Robertson, 1891. Amer. Ent. Soc., Trans. 18: 53. 2, ¢. Preoee.
Andrena bisalicis Viereck, 1908. Ent. News 19: 42. N. name.
Andrena (Andrena) adelae Viereck, 1922. Boston Soc. Nat. Hist., Occas. Papers 5: 37. °.

1832 Hymenoptera in America North of Mexico

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191, 192. 2, ¢ (key). —Mitchell,
1958. In Krombein, U. S. Dept. Agr., Agr. Monog 2, First Suppl. p. 212 (synonymy).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 219-220, figs. 42-44, table 6
(redescription). —Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 5, figs. 1-2 (pupa).

Biology: Rozen, 1966. Amer. Mus. Novitates 2244: 26 (parasite).

brevipalpis Cockerell. Colo.
Andrena brevipalpis Cockerell, 1930. Ann. and Mag. Nat. Hist. (10) 5: 109. °.

Taxonomy: Cockerell, 1932. Pan-Pacifie Ent. 8: 175. 2. —Cockerell, 1937. Amer. Mus.
Novitates 899: 4. 9.

campanulae Viereck and Cockerell. Colo., Nebr.
Andrena campanulae Viereck and Cockerell, 1914. U.S. Natl. Mus., Proce. 43: 38, 43. 6, 9.

Taxonomy: Cockerell, 1932. Pan-Pacific Ent. 8: 177. 2, 6.
candida Smith. B. C. to south. Calif., Colo. and Utah; Mexico (Baja California). Pollen:
Polylectic, visits a wide variety of flowers during both the first and second broods
including Acer grandidentatum, A. platynoides, Adenostoma fasciculatum, Alyssum
maritimum, Amelanchier, Arctostaphylos glauca, Baccharis glutinosa, B. viminea,
Brassica alba, B. adpressa, B. arvensis, B. campestris, B. nigra, Calandrinia menziesii,
Ceanothus cordulatus, C. crassifolius, C. cuneatus, C. integerrimus, C. orcuttii, C.

_ palmer, C. verrucosus, Chamaebatia foliolosa, Conium maculatum, Cryptantha
flaccida, C. lepida, Cymopterus longipes, Daucus carota, Eriogonum fasciculatum, E. f.
polifolui, Eriophyllum tridactyllum, Gnaphalium bicolor, Isomeris arborea, Lomatiwm
dasycarpum, L. grayi, Ligusticum vulgare, Lupinus paynei, Malus pumila, Medicago
sativa, Melilotus alba, Nemophila menziesii, Phacelia affinis, P. distans, P. minor, P.
ramosissima, Plagiobothrys tenellus, Prunus fremontii, P. ilicifolia, P. lyonii, P.
subcordata, Ranunculus californicus, Rhamnus californica, R. crocea, R. ilicifolia,
Salix argophylla, S. discolor, S. exigua, S. hindsiana, S. laevigata, S. lasiolepis, S. nigra,
Salvia mellifera, Sambucus caerulea, Sisymbrium irio, S. officinale.

Andrena candida Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 56. @.

?Andrena (Andrena) candida tramoserica Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68:
533. 2.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 155 (type). —Cockerell, 1906. Psyche
13: 10 (type). —Lanham, 1941. Ent. Soc. Amer., Ann. 34: 706, 708. 2, ¢ (key). —Youssef
and Bohart, 1968. Kans. Ent. Soc., Jour. 41: 450-455, figs. 2-7, 11-22 (immature stages).

Biology: Linsley, 1937. Brooklyn Ent. Soc., Bul. 32: 127 (floral relationships, spring and early
summer broods). — Youssef and Bohart, 1968. Kans. Ent. Soc., Jour. 41: 442-455, 22 figs.
(nest architecture, life history, floral relationships).

ceanothina Cockerell. Calif. Pollen: Unknown, but visits flowers of Arctostaphylos, Ceanothus
crassifolius, Ribes indecorum.

Andrena ceanothina Cockerell, 1936. Pan-Pacifie Ent. 12: 138. 9.

cercocarpi Cockerell. Calif. Pollen: Unknown, but visits flowers of Cercocarpus.

Andrena (Platanarena) cercocarpi Cockerell, 1986. Pan-Pacific Ent. 12: 140. °.

Taxonomy: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 332 (subgeneric position).

chlorinella Viereck. Oreg.

Andrena chlorinella Viereck, 1904. Canad. Ent. 35: 189. 9.

Andrena xanthostigma Viereck, 1904. Canad. Ent. 35: 193. 9.

Taxonomy: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 333 (subgeneric position).

claremonti Viereck. Calif.
Andrena claremonti Viereck, 1926. Pomona Jour. Ent. Zool. 18: 2. ¢.

coloradina Viereck and Cockerell. Colo.
Andrena coloradina Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 53, 56. 2.
Taxonomy: Cockerell, 1933. Pan-Pacifie Ent. 9: 154. 2 (key).

fulvihirta Viereck and Cockerell. B. C. to Colo. and Calif.
Andrena fulvihirta Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 54. 2.

Superfamily APOIDEA 1833

Andrena washingtoni manitouensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce.
48: 31. 6.

Taxonomy: Timberlake, 1951. Jn Linsley, Jn Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1066 (synonymy).
geranii Robertson. New England states and Va., west to Minn. and Colo. Pollen: Unknown,
but visits flowers of Arabis, Blephilia, Geranium, Hydrophyllum, Osmorrhiza,
Polemonium.
Andrena geranii Robertson, 1891. Amer. Ent. Soc., Trans. 18: 54. 2, 6.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 192. 2, d (key). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 221-222, figs. 44-45, table 6 (redescription).

haroldi Timberlake. Calif. Pollen: Unknown, but visits flowers of Lasthenia.
Andrena (Micrandrena) haroldi Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 398. 2, é.

Taxonomy: Ribble, 1968. Nebr. Univ. State Mus., Bul. 8: 333 (subgeneric position).
knuthiana Cockerell. Oreg., Calif. Pollen: Unknown, but visits flowers of Ceanothus

cordulatus, Cryptantha lepida, Rhamnus californica, Salix, Sambucus velutina.
Andrena knuthiana Cockerell, 1906. Ztschr. System. Hym. Dipt. 1: 80. 2, 6.

Taxonomy: Cockerell, 1901. Ent. News 12: 74. °.
lata Viereck. N.S. to Minn., south to Ga. Pollen: Unknown, but visits flowers of Brassica,
Hydrangea, Potentilla, Prunus, Pyrus, Rubus.
Andrena (Andrena) lata Viereck, 1922. Boston Soc. Nat. Hist., Occas. Papers 5: 39. 2.
Andrena (Andrena) vulgaris Viereck, 1922. Boston Soe. Nat. Hist., Occas. Papers 5: 40. d.

Taxonomy: Atwood, 1934. Canad. Jour. Res. 10 (2): 208, 210. 2, d (key). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 222-224, figs. 44-45, table 6 (redescription, synonymy).
medionitens Cockerell. B. C. and Alta. to Calif. and Colo. Pollen: Unknown, but visits flowers
of Cymopterus terebinthus, Eriogonum latifolium var. nudum, E. subscaposum,
Erysimum asperum, Gilia tenuiflora, Hackelia jessicae, Holodiscus discolor, Horkelia
bernardina, H. fusca, Oxytropis, Potentilla glandulosa, P. wheeleri, Sisymbrium
incisum, Spraguea umbellata, Streptanthus tortwosus.
Andrena medionitens Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 101. °.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 192, 194. 2, 6. —Viereck and Cockerell, 1914. U.
S. Natl. Mus., Proc. 48: 48. 2 (variation).

Biology: Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 249 (ecology, as
madronitens).
phaceliae Mitchell. Md., Va., Ill. Pollen: Unknown, but visits flowers of Alsine pubera,
Phacelia dubia, P. purshii.
Andrena (Thysandrena) phaceliae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 224,
figs. 44, 45. 2, o.
phocata Cockerell. Colo.
Andrena phocata Cockerell, 1910. Canad. Ent. 42: 369. °.
subcandida Viereck. B. C. to Calif.
Andrena subcandida Viereck, 1904. Canad. Ent. 36: 198. °.
subdistans Viereck. Wash., Oreg.
Andrena subdistans Viereck, 1904. Canad. Ent. 36: 193. 9.
subtrita Cockerell. Nev.
Andrena subtrita Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 268. 2.
taeniata Viereck. Calif.
Andrena (Andrena) taeniata Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 538. 9°.
trizonata (Ashmead). Colo.
Cilissa trizonata Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 6. °.
vierecki Cockerell. Utah, Colo., Calif. Parasite: Myopa rubida (Bigot).
Andrena vierecki Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 26. @.
Andrena boharti Linsley, 1939. Pan-Pacific Ent. 15: 159. 9, 3.

1834 Hymenoptera in America North of Mexico

Taxonomy: Lanham, 1941. Ent. Soc. Amer., Ann. 34: 706, 707. 2, 5 (key). —Timberlake,
1951. In Linsley, In Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2:
1085 (synonymy).

Biology: MacSwain and Bohart, 1947. Pan-Pacific Ent. 23: 30 (parasite).

w-scripta Viereck. B. C. to Calif. Pollen: Unknown, but visits flowers of Acacia greggii,
Adenostoma fasciculatum, Arctostaphylos glandulosa, A. mariposa, A. patula, A.
pungens, Ceanothus cordulatus, C. cuneatus, C. greggit, C. soredatus, Chamaebatia
foliolosa, Claytonia spathulata, Cryptantha intermedia, C. lepida, Daucus carota,
Eriogonum fasciculatum, Eriophyllum confertiflorum, Fritallaria, Heteromeles
arbutifolia, Holodiscus discolor, Horkelia, Lasthenia chrysostoma, Lomatium, Lotus
scoparius, Phacelia distans, Prunus ilicifolia, Ranunculus californicus, Rhamnus

californicus, Salix laevigata.
Andrena w-scripta Viereck, 1904. Canad. Ent. 36: 193, 194. 2, 6.

Genus ANDRENA Subgenus TRACHANDRENA Robertson

Trachandrena Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189.
Type-species: Andrena rugosa Robertson. Orig. desig.

All the species of this subgenus, except the Eurasian Andrena lagopus Latreille, are centered
in America north of Mexico and are not known to occur south of the United States.

Revision: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 235-371, 69 figs., 2 tables, 15 maps (N.
Amer. spp.).

Taxonomy: Lanham, 1949. Calif. Univ. Pubs. Ent. 8: 216 (tax. characters). —LaBerge, 1964.
Nebr. Univ. State Mus., Bul. 4: 282, 292, 308 (tax. characters). —Warncke, 1968. Coimbra
Univ. Est. Mus. Zool., Mem. 307: 68-69 (tax. status).

amphibola (Viereck). B. C. south to Calif., east to Alta., Mont. Wyo., Nebr. and Colo. Pollen:

Apparently polylectic, visits flowers of Astragalus, Daucus pusilla, Drymocallis,
Eriogonum flavum, Geum triflorum, Holodiscus discolor, Iris, Lappula floribunda,
Lonicera, Lupinus, Medicago sativa, Penstemon, Phacelia, Potentilla, Salix, Senecio,
Symphoricarpos, Taraxacum officinale, Thermopsis divaricata.

Trachandrena amphibola Viereck, 1904. Canad. Ent. 35: 158, 159. 9, 6.

Trachandrena crassihirta Viereck, 1904. Canad. Ent. 35: 158. 9.

Trachandrena perdensa Viereck, 1904. Canad. Ent. 35: 158. 9.

Trachandrena hadra Viereck, 1904. Canad. Ent. 35: 158. °.

Trachandrena limarea Viereck, 1904. Canad. Ent. 36: 158, 159. 2, 6.

Andrena (Trachandrena) eriogoni Cockerell, 1927. Ent. Soc. Amer., Ann. 20: 397. °.

Andrena (Trachandrena) seneciophila Cockerell, 1928. Psyche 35: 62. °.

Taxonomy: Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 10. 2 (as perdensa).
—Lanham, 1914. Ent. Soc. Amer. Ann. 34: 704. 2 (key, as seneciophila). —LaBerge, 1973.
Amer. Ent. Soc., Trans. 99: 284-292, figs. 1, 7, 34-35 (redescription, synonymy, variation in
color of vestiture).

ceanothi Viereck. N. S. and Que., south to Ga., west to Minn. and Colo. Pollen: Polylectic, visits
flowers of Amelanchier, Amorpha fruticosa, Barbarea vulgaris, Ceanothus americanus,
Cerasus serotina, Chamaedaphne, Chrysanthemum leucanthemum, Cornus
candidissimus, Crataegus, Fragaria, Ledum; Prunus, Ptelea, Rhus, Rubus argutus, R.
villosus, Salix, Taraxacum officinale, Vaccinium angustifolium, Viburnum,

Waldsteinia.

Andrena (Trachandrena) ceanothi Viereck, 1917. Amer. Ent. Soc., Trans. 48: 404. 2 (6
misdet.).

Andrena (Trachandrena) compacta Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
181. 3.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 17. 2 (key). —Atwood, 1934. Canad.
Jour. Res. 10: 212. 9, d. —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 179-180,
figs. 35-36, table 5 (redescription, synonymy). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99:
346-349, figs. 1, 12, 58, 59 (redescription, synonymy).

Superfamily APOIDEA 1835

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 349 (floral relationships).
cleodora cleodora (Viereck). B. C. to north. Calif., east to Mont., Wyo. and Colo., ?Ariz. (Santa

Catalina Mts.). Pollen: Oligolege of Ceanothus including C. cuneatus, C. integerrimus, C.
laevigatus, but visits other flowers presumably for nectar including Arctostaphylos,
Aster, Chamaebatia foliolosa, Conium maculatum, Eriogonum nudum, Lupinus,
Phacelia, Prunus, Rhamnus, Ribes, Salix, Sorbus.

Trachandrena cleodora Viereck, 1904. Canad. Ent. 35: 158. @.

Andrena (Trachandrena) lutzi Cockerell, 1931. Amer. Mus. Novitates 458: 12. °.

Taxonomy: Cockerell, 1932. Pan-Pacifie Ent. 8: 173. ¢. —LaBerge, 1973. Amer. Ent. Soc.,
Trans. 99: 323-326, figs. 1, 12, 48-49 (redescription).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 327 (floral relationships).
cleodora melanodora Cockerell. South. Calif. Pollen: Oligolege of Ceanothus including C.
cordulatus, C. integerrimus, C. palmeri, but visits other flowers presumably for nectar
including Arctostaphylos patula, Barbarea orthoceras, Cryptantha lepida, Eriodictyon
californicum, Prunus emarginata, P. ilicifolia, Pyrus malus, Rhamnus californica.
Andrena (Trachandrena) melanodora Cockerell, 1932. Pan-Pacifie Ent. 8: 173. °.

Taxonomy: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 326-327, figs. 48-49 (redescription).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 327 (floral relationships).
cupreotincta Cockerell. B. C., Alta. and Sask., south to Calif., Nev., Utah and N. Mex. Pollen:

Apparently polylectic, visits flowers of Abies concolor, Arctostaphylos nevadensis,
Ceanothus integerrimus, Eriogonum, Hackelia, Lesquerella, Penstemon, Prunus
virginiana demissa, Pyrus malus, Rhamnus californica, R. aul Ribes inerme,
Rubus, Salix, Taraxacum officinale.

Andrena cupreotincta Cockerell, 1901. Canad. Ent. 33: 153. 9.

Trachandrena ochreopleura Viereck, 1904. Canad. Ent. 35: 158. °.

Andrena (Trachandrena) swenki Viereck and Cockerell, 1914. U. S. Natl. Mus., Proe. 48: 3.
OF

Andrena (Trachandrena) jockorum Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48:
3. d.

Trachandrena abjuncta Cockerell, 1929. N. Y. Ent. Soc., Jour. 37: 445. 2.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 158. 2 (key). —Cockerell, 1937. Ent. News 49: 257.
2 (as abjuncta). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 315-321, figs. 1, 11, 46-47
(redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 321 (floral relationships).

cyanophila Cockerell. B. C. and Alta., south to Calif., Nev. and Ariz. Pollen: Apparently
polylectic, visits flowers of Argentina anserina, Ceanothus fendleri, Euphorbia,
Geranium fremontii, Oreocarya nana, Pinus ponderosa, Polemonium, Potentilla
arguta, P. glandulosa, Prunus melanocarpa, Ranunculus Montensis, Ribes
longiflorum, Senecio perplexis, Taxaxacum officinale.
Andrena cyanophila Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 431. 2.
Andrena tacitula Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 262. 6.

Taxonomy: Cockerell, 1916. Canad. Ent. 48: 253. —Lanham, 1941. Ent. Soc. Amer., Ann. 34:
704, 707. 2, d (key). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 353-357, figs. 1, 10, 64-65
(redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 357 (flora! relationships).

forbesii Robertson. N.S. to B. C., south to Ga., west to Wash., Oreg. and Calif. Pollen:
Polylectic, apparently prefers pollens of Rosaceae and Salicaceae over other sources;
visitation records include Acer negundo, Alliaria officinalis, Amelanchier arborea, A.
canadensis, Aronia melanocarpa, Astragalus, Barbarea vulgaris, Berberis vulgaris,
Brauneria purpurea, Callirhoe digitata, Cardamine bulbosa, Ceanothus ovatus, Cercis
canadensis, Claytonia caroliniana, C. virginica, Cornus florida, C. mas, Crataegus,
Dentaria laciniata, Erythronium americanum, Euphorbia commutata, Forsythia
suspensa, Fragaria virginiana, Heracleum lanatum, Hieracium, Ilex, Lappula

1836 Hymenoptera in America North of Mexico

heterosperma, Lepargyrea canadensis, Lomatium daucifolium, L. foeniculaceum,
Lonicera, Melilotus alba, M. officinalis, Paeonia, Pastinaca sativa, Physocarpus
opulifolius, Populus deltoides, Prunus americanus, P. demissa, P. melanocarpa, P.
serotina, P. virginiana, Ptelea, Pyracantha, Pyrus leonis, P. malus, Ranunculus
abortivus, Rhamnus lanceolatus, Rhus aromatica, R. canadensis, R. toxicodendron,
Ribes gracile, Rosa, Rubus argenteus, Salix discolor, S. nigra, Sedum ternatum,
Shepherdia argentea, Spiraea aruncus, S. latifolia, Symphoricarpos, Tamarix gallica,
Taraxacum officinale, Thlaspi arvense, Vaccinium, Viburnum opulus, V. prunifolium,
Zizia aurea.

Andrena forbesti Robertson, 1891. Amer. Ent. Soce., Trans. 18: 59. @.

Trachandrena indotata Viereck, 1904. Canad. Ent. 36: 158, 159. 2, 6.

Andrena (Trachandrena) lincolni Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48:
130%

Andrena (Trachandrena) rodecki Cockerell, 1929. Ent. Soe. Amer., Ann. 22: 755. °.

Andrena (Trachandrena) pyracanthae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
194, fig. 35. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189, 190. 2, 6. —Cockerell, 1931.
Amer. Mus. Novitates 458: 16. 2. (as rodecki). —Cockerell, 1931. Amer. Mus. Novitates
458: 17 (key, as lincolni). —Atwood, 1934. Canad. Jour. Res. 10: 208, 209, figs. 2, ¢d (key).
—Lanham, 1941. Ent. Soc. Amer., Ann. 34: 704, 707. 2, ¢ (key). —Lanham, 1941. Ent. Soe.
Amer., Ann. 34: 704. ? (key, as lincolni). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 183-184, figs. 35-36, table 5 (redescription). —LaBerge, 1973. Amer. Ent. Soc.,
Trans. 99: 261-267, figs. 1, 4, 24-25 (redescription, synonymy, geogr. variation).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 266-267, table 1 (floral relationships).
fuscicauda (Viereck). B. C. to Calif. and west. Nev. and Ariz (Yuma). Pollen: Apparently

polylectic, visits flowers of Arctostaphylos patula, Berberis, Ceanothus cordulatus, C.
mtegerrimus, C. leucodermis, C. palmeri, C. soredatus, Chamaebatia foliolosa,
Cryptantha intermedia, Eriodictyon californicum, Eriogonum, Gilia, Horkelia parryi,
Malus, Nemophila menziesii, Pinus ponderosa, Potentilla, Prunus demissa, P.
emarginata, P. ilicifolia, P. subcordata, Quercus dumosa, Rhamnus californica, R.
crocea, Salix laevigata, Taraxacum denselionis.

Trachandrena fuscicauda Viereck, 1904. Canad. Ent. 35: 159. 9.

Andrena (Trachandrena) californica Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 373. 2,
é.

Andrena (Trachandrena) californica wickhami Timberlake, 1951. U. S. Natl. Mus., Proc.
101: 375. 2.

Taxonomy: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 357-361, figs. 1, 14, 66-67
(redescription, synonymy).
Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 361 (floral relationships).
heraclei Robertson. N. Y. to N. C., west to Kans. Pollen: Unknown, but visits flowers of
Camassia esculenta, Celastrus scandens, Cercis canadensis, Crataegus, Cornus,
Heracleum, Pastinaca sativa, Prunus, Ptelea, Salix, Taenidia, Washingtonia

longistylus.
Andrena heraclei Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 336. 2.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189. 2. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 184, fig. 35, table 5 (redescription). —LaBerge, 1973. Amer. Ent.
Soc., Trans. 99: 343-346, figs. 1, 3, 56-57 (redescription).

hippotes Robertson. N.S. to B. C., south to Ga., west to Wash., Oreg. and north. Calif. Pollen:
Polylectic, visits a wide variety of flowers including Acer, Aesculus glabra sargentii,
Amelanchier arborea, Amorpha canescens, Angelica atropurpurea, Barbarea vulgaris,
Berberis, Caltha palustris, Camassia esculenta, Ceanothus ovatus, Celastrus, Cercis
canadensis, Chamaedaphne, Claytonia, Cornus mas, Crataegus mollis, Daucus carota,
Dentaria laciniata, Descourainia intermedia, Echinacea angustifolia, E. purpurea,
Encelia, Euphorbia, Fragaria, Fraxinus, Hieracium, Ligustrum vulgare, Lomatium
triternatum, Lonicera fragrantissima, Melilotus alba, M. officinalis, Nyssa sylvatica,

Superfamily APOIDEA 1837

Paeonia, Pastinaca sativa, Phacelia, Physocarpus opulifolius, Potentilla anserina,
Prunus americana, P. demissa, P. serotina, P. virginiana, Ptelea, Pyracantha
yunannensis, Pyrus malus, Ranunculus abortivus, Rhamnus, Rhus glabra, Ribes
aureum, Rosa carolina, Rubus, Salix nigra, Shepherdia argentea, Spiraea discolor, S.
sorbifolia, Sphaeralcea coccinea, Symphoricarpos, Taraxacum officinale, Thlaspi
arvense, Trollius laxa, Vaccinium, Viburnum dentatum, V. opulus, Zizia aurea.

Andrena hippotes Robertson, 1895. Amer. Ent. Soe., Trans. 22: 120. 2, 3.

Ae (Trachandrena) perforatella Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17:

we

Andrena (Trachandrena) arenakensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:

Migtiga sos or

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 184-185, figs. 35, 37, table 5
(redescription, synonymy). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 292-299, figs. 1, 8,
36-37 (redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 298, table 2 (floral relationships).

mariae Robertson. N.S. to N. W. T. and B. C., south to Ga., west to N. Mex., Utah, Idaho and

Wash. Pollen: Oligolege of Salix including S. babylonica, S. bebbiana, S. nigra, S. tristis,
but visits other flowers including Amelanchier, Astragalus, Barbarea vulgaris,
Cogswellia, Comandra, Crataegus mollis, Erigenia, Erigeron philadelphicus,
Erythronium, Fragaria, Hackelia floribunda, Kalmia, Lomatium daucifolium, L.
foeniculaceum, Pastinaca sativa, Prunus americanus, Pulsatilla, Pyrus malus, Ribes
gracile, Rubus, Sisymbrium, Spiraea, Taraxacum officinale.

Andrena mariae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 58. 2, 6.

Andrena sphecodina Cockerell and Casad, 1896. Ann. and Mag. Nat. Hist. (6) 18: 78. 2, 6.

Andrena mariae var. concolor Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 46. ?, 3.

Andrena (Trachandrena) submariae Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce.
48: 4. 9.

Andrena (Trachandrena) martialis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48:
Sor

Andrena (Trachandrena) sphecodiniformis Viereck and Cockerell, 1914. U. S. Natl. Mus.,
Proce. 48: 6. d.

Andrena (Trachandrena) profundiformis Viereck and Cockerell, 1914. U. S. Natl. Mus.,
Proc. 48: 8. d.

Andrena (Trachandrena) stricklandi Cockerell, 1936. Canad. Ent. 68: 275. °.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189, 190. 2, 6. —Viereck, 1907. Ent.
News 18: 284, 286. 2, ¢ (key). —Cockerell, 1929. Ent. Soc. Amer., Ann. 22: 754 (as
martialis). —Cockerell, 1929. Ent. Soc. Amer., Ann. 22: 754 (as sphecodina). —Atwood,
1934. Canad. Jour. Res. 10: 208, 210. 2, ¢ (key). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 187-188, figs. 35, 38, table 5 (redescription). —LaBerge, 1973. Amer. Ent.
Soc., Trans. 99: 299-305, figs. 1, 9, 39-41 (redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 305 (floral relationships).
miranda Smith. N.S. to N. J. and Pa., west to Alaska, south to N. Mex., Ariz., Nev. and Calif.

(Alta and Carrville). Parasite: Stylops sp. Pollen: Polylectic, visits a wide variety of
flowers including Achillea millefolia, Angelica, Aralia latifolia, Argemone, Astragalus,
Barbarea vulgaris, Blepharipappus, Brassica campestris, Ceanothus, Cirsium
undulatum, Cornus, Crataegus, Drymocallis arguta, Euvhorbia, Geranium richardsoni,
Heracleum lanatum, Lappula heterosperma, Lotus corniculatus, Malvastrum, Melilotus
alba, M. officinalis, Pastinaca sativa, Potentilla fruticosa, P. recta, Prunus virginiana
demissa, Ranunculus septentrionalis, Ribes, Rosa arkansana, R. carolina, R.
multiflora, Rubus adoratus, Rudbeckia hirta, Sanicula marilandica, Spiraea latifolia,
S. sorbifolia, Symphoricarpos occidentalis, Taraxacum officinale, Vaccinium,
Viburnum opulus, Waldsteinia fragarioides, Zizia aptera, Z. aurea.

Andrena miranda Smith, 1879. Desc. New Species Hym. Brit. Mus., p. 54. 2, d.

Andrena grandior Cockerell, 1897. Entomologist 30: 307. 2.

Andrena multiplicata Cockerell, 1902. Canad. Ent. 34: 46. °.

Andrena multiplicatiformis Viereck, 1907. Ent. News 18: 284. °.

1838 Hymenoptera in America North of Mexico

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 153. 2, d (types). —Cockerell, 1906.
Psyche 13: 34 (type). —Atwood, 1934. Canad. Jour. Res. 10: 208, 210. °, 3 (key).
— Atwood, 1934. Canad. Jour. Res. 10 (2): 208, 210 (as grandior). —Timberlake, 1951. In
Linsley, In Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2: 1072
(synonymy). —Mitchell, 1958. Jn Krombein, U. S. Dept. Agr., Agr. Monog. 2, First Suppl.
p. 215 (synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 190-191, figs. 35,
38, table 5 (redescription, synonymy). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99:
267-273, figs. 1, 5, 26-27 (redescription).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 272-273 (floral relationships,
stylopization).
nuda Robertson. Maine to Ga., west to Tex., N. Mex., Utah and Wyo. Pollen: Polylectic,

although with a preference for flowers of the Rosaceae: visitation records include
Aruncus, Benzoin aestivale, Cercocarpus, Claytonia, Conium, Crataegus mollis,
Erigenia, Euphorbia, Heracleum, Ilex, Lomatium dissectum, Melilotus officinalis,
Myosotis, Pastinaca sativa, Penstemon, Prunus americana, P. serotina, P. virginica,
Ptelea, Pyracantha, Pyrus leonis, P. malus, P. pyrus, Rhamnus lanceolata, Rhus,
Rubus argutus, Salix, Sanicula, Sassafras, Staphylea, Symplocus, Taenidia, Viburnum
prunifolium, Zizia.

Andrena nuda Robertson, 1891. Amer. Ent. Soc., Trans. 18: 57. 9.

Andrena (Trachandrena) davisiana Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce.
48: 6. d.

Andrena (Trachandrena) pseudobscura Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul.
141: 198, fig. 36. 6.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189. ? (key). —Lanham, 1941. Ent.
Soc. Amer., Ann. 34: 704. 2? (key). —Lanham, 1941. Ent. Soc. Amer., Ann. 34: 707. 3 (key,
as davisiana). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 192-193, fig. 35
(redescription). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 333-338, figs. 1, 14, 52-53
(redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 337-338 (floral relationships).

quintiliformis Viereck. B. C. to Calif., east to Idaho, Nev. and Utah. Pollen: Apparently
polylectic, visits flowers of Achillea millefolia, Allium, Arctostaphylos, Ceanothus
cuneatus, C. laevigatus, Chamaebatia foliolosa, Cirsium, Eriogonum, Lappula,
Lomatium triternatum, Phacelia humilis, Ranunculus, Salix, Solidago californica.
Andrena (Trachandrena) quintiliformis Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 606.

On
Andrena (Trachandrena) coactifera Viereck, 1926. Calif. Acad. Sci., Proc. (4) 15: 399. °.

Taxonomy: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 278-281, figs. 1, 6, 30-31 (redescription,
synonymy).
Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 281 (floral relationships).
quintilis Robertson. Wis. and IIl., west to N. Dak., S. Dak., Nebr., Colo. and Kans. Pollen:
Polylectic, although exhibits a decided preference for the pollen and nectar of Amorpha
including A. canescens, A. fruticosa, but visits other flowers including Apocynum
cannabinum, Chrysanthemum leucanthemum, Crataegus, Drymocallis arguta,
Echinacea angustifolia, Erigeron ramosus, Pastinaca sativa, Ptelea, Pycnanthemum
flexuosum, P. linifolium, Salix, Symphoricarpos, Zizia aurea.
Andrena quintilis Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 46. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189. 2 (key). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 14: 195-196, fig. 35, table 5 (redescription). —LaBerge, 19783.
Amer. Ent. Soc., Trans. 99: 281-284, figs. 1, 6, 32-33 (redescription).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 284 (floral relationships).
rehni Viereck. Que. and Ont., south to G., west to Wis. Pollen: Apparently polylectic, visits
flowers of Aruncus dioecium, Castanea pumila, Ceanothus, Chrysanthemum
leucanthemum, Cicuta maniculata, Hydrangea, Ilex opaca, Quercus prinoides, Salix.
Andrena rehni Viereck, 1907. Ent. News 18: 284. °.

Superfamily APOIDEA 1889

Andrena (Trachandrena) votula Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 204.
Of

Taxonomy: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 349-353, figs. 1, 15, 60-63
(redescription, synonymy).
rugosa Robertson. Que. and New England, south to Ga., west to Minn. and Nebr. Pollen:

Polylectic, visits flowers of Acer, Amelanchier arborea, A. canadensis, Angelica,
Astragalus, Barbarea vulgaris, Brassica arvensis, Claytonia virginica, Cornus mas,
Crataegus punctata, Dentaria laciniata, Dirca, Erigenia bulbosa, Fragaria, Geranium,
Hepatica, Heracleum, Isopyrum, Macrocalyx nyctelea, Melilotus officinalis, Paeonia,
Pastinaca sativa, Physocarpus opulifolius, Prunus americana, P. virginiana, Ptelea,
Pyracantha, Pyrus malus, Rhus, Ribes, Rosa multiflora, Rubus argutus, R.
occidentalis, Salix, Sanicula, Spiraea aruncus, Symphoricarpos, Taraxacum officinale,
Viburnum, Zanthoxylum, Zizia aurea.

Andrena rugosa Robertson, 1891. Amer. Ent. Soc., Trans. 18: 58. 2, 6.

Andrena paenerugosa Viereck, 1907. Ent. News 18: 285. °.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189, 190. 2, ¢ (key). —Atwood,
1934. Canad. Jour. Res. 10: 208, 210. 2, o (key). —Mitchell, 1960. N. C. Expt. Sta. Tech.
Bul. 141: 198-199, figs. 35-36, table 5 (redescription, synonymy). —LaBerge, 1973. Amer.
Ent. Soc., Trans. 99: 327-332, figs. 1, 18, 50-51 (redescription).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 332 (floral relationships).

salicifloris Cockerell. B. C. and Alta., south to Calif., Nev., Utah and Colo. Pollen: Polylectic,

but with some predilection for the flowers of Salix; visitation records include Acacia,
Acer, Amelanchier, Arctostaphylos, Berberis, Brassica, Ceanothus, Cerastium
oreophyllum, Chamaebatia foliolosa, Cicuta, Cistus, Epilobium angustifolium,
Eriogonum, Geranium, Grossularia, Hackelia patens, Hieracium lanatum, Lasthenia,
Lepidium, Lomatium dissectum, L. triternatum, Pastinaca sativa, Prunus demissa, P.
ilicifolia, P. virginiana, Pulsatilla patens, Pyracantha yunannensis, Pyrus malus,
Ranunculus californicus, Ribes lacustre, Rubus nutkana, R. ursinus, R. vitifolius,
Taraxacum densileonis, T. officinalis.

Andrena salicifloris Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 351. °.

Trachandrena auricauda Viereck, 1904. Canad. Ent. 35: 159, 161. 9.

Andrena (Trachandrena) tacitula var. grossulariae Viereck and Cockerell, 1914. U.S.
Natl. Mus., Proc. 48: 9. 6.

Andrena (Trachandrena) nortoni Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 602. 2, d.

Andrena (Trachandrena) veris Cockerell, 1934. Pan-Pacific Ent. 9: 156. 2, d.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 158, 159. 2, d (variation). —Lanham, 1941. Ent.
Soc. Amer., Ann. 34: 704, 707. 2°, d (key, as veris). —LaBerge, 1973. Amer. Ent. Soc.,
Trans. 99: 255-261, figs. 1, 3, 22-23 (redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 260 (floral relationships).
semipunctata Cockerell. Wash. to south. Calif., east to southeast. Utah and southwest. N. Mex.
Pollen: Oligolege of Salix including S. exigua, S. hindsiana, S. laevigata, S. lasiolepis, S.
nigra, but visits other flowers presumably for nectar including Baccharis, Ceanothus
soredatus, Prunus, Pyrus malus, Rhus trilobata, Sambucus, Senecio salignifolia,
Sisymbrium irio, Tamarix gallica, Taraxacum officinale.
Andrena semipunctata Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 102. 2.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 159. 2, d (key). —LaBerge, 1973. Amer. Ent. Soc.,
Trans. 99: 311-315, figs. 1, 9, 44-45 (redescription).

Biology: Linsley, 1937. Brooklyn Ent. Soc., Bul. 32: 127 (floral relationships, adult behavior).
sigmundi Cockerell. N. B. south to Ga., west to B. C., south to Calif., Utah and N. Mex. Pollen:
Oligolege of Salix including S. bebbiana, S. discolor, but visits other flowers including
Acer saccharinum, Amelanchier canadensis, Barbarea vulgaris, Ceanothus fendleri,
Crataegus, Fragaria virginiana, Fraxinus, Kalmia latifolia, Lepargyrea argentea, L.
canadensis, Melilotus alba, Populus, Potentilla, Prunus demissa, P. virginiana,

1840 Hymenoptera in America North of Mexico

Pulsatilla ludoviciana, Spiraea, Taraxacum officinale, Vaccinium angustifolium,
Waldsteinia fragarioides.

Andrena sigmundi Cockerell, 1902. Canad. Ent. 34: 45. 9.

Andrena radiatula Cockerell, 1902. Canad. Ent. 34: 46. °.

Andrena weedi Viereck, 1907. Ent. News 18: 284, 285. 2, 6.

Andrena (Trachandrena) prunicola Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 12: 375.
oe

Andrena (Trachandrena) moscovensis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc.
48: 13. 2.

Andrena (Trachandrena) corrugata Cockerell, 1931. Amer. Mus. Novitates 458: 9. 2.

Taxonomy: Cockerell, 1931. Amer. Mus. Novitates 458: 17 (key, as moscovenis). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 200-202, figs. 36, table 5 (redescription,
synonymy). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 249-255, figs. 1, 2, 16-21
(redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 255 (floral relationships).

spiraeana Robertson. Ont. and Maine to Ga., west to Minn., Nebr. and Kans. Pollen: Polylectic,

visits flowers of Alliaria officinalis, Angelica, Aruncus, Azalea, Barbarea vulgare,
Castanea pumila, Ceanothus americanus, Cornus, Crataegus, Cryptotaenia, Erigeron
philadelphicus, Euphorbia commutata, Heracleum, Hydrangea, Melilotus alba,
Pastinaca sativa, Physocarpus opulifolius, Ptelea, Rhus glabra, R. typhine, Rosa,
Spiraea aruncus, S. latifolia, Sorbaria sorbifolia, Viburnum nudum, Zizia.

Andrena spiraeana Robertson, 1895. Amer. Ent. Soc., Trans. 22: 120. 2, d.

Andrena (Trachandrena) montensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
191, fig. 35. 9.

Andrena (Trachandrena) unica Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 208,
fig. 36. d.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 189, 190. 2, ¢ (key). —Viereck,
1907. Ent. News 238: 107 (key). —Lanham, 1941. Ent. Soc. Amer., Ann. 34: 704, 707. 2, 3
(key). —-Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 202-2038, figs. 35, 38, table 5
(redescription). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 338-342, figs. 1, 11, 58-54
(redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 342 (floral relationships).

striatifrons Cockerell. B. C. and Alta., south to Ariz. and Calif. Pollen: Oligolege of Salix, but

visits other flowers including Arctostaphylos, Eriogonum, Hackelia floribunda,
Lomatium, Prunus, Pyrus malus, Ranunculus, Rubus parviflorus, Symphoricarpos,
Taraxacum officinale, Vaccinium.

Andrena striatifrons Cockerell, 1897. Entomologist 30: 308. 2.

Trachandrena pernuda Viereck, 1904. Canad. Ent. 35: 159. °.

Andrena trachandrenoides Viereck, 1904. Canad. Ent. 36: 190, 195. 2, ¢.

Andrena (Trachandrena) marioides Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 601. °.

Andrena (Trachandrena) politissima Cockerell, 1918. Ann. and Mag. Nat. Hist. (9) 166. 9.

Andrena dolichotricha Cockerell, 1924. Ent. News 35: 348. 6.

Andrena (Trachandrena) brevibasis Cockerell, 1931. Amer. Mus. Novitates 458: 8. 9.

Andrena (Trachandrena) postnitens Cockerell, 1931. Amer. Mus. Novitates 458: 15. 2.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 159. 2 (key). —LaBerge, 1973. Amer. Ent. Soc.,
Trans. 99: 305-311, figs. 1, 10, 42-43 (redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 311 (floral relationships).
virginiana Mitchell. P. E. I., Que, and Ont., south to Ga., west to Minn., Il. and Ark. Pollen:

Apparently polylectic, visits flowers of Ceanothus americanus, Cerasus serotina,
Daucus carota, Ilex verticillata, Kalmia, Melilotus alba, Pycnanthemum linifolium,
Spiraea latifolia, Solidago.

Trachandrena obscura Robertson, 1902. Amer. Ent. Soc., Trans. 35: 189. 2. Preoce.

Andrena (Trachandrena) virginiana Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
203522

Andrena (Biareolina) obscurida Warncke, 1970. Bayer. Ent., Nachr. 19: 30. N. name.

Superfamily APOIDEA 1841

Taxonomy: Viereck, 1907. Ent. News 18: 284. 2 (key, as obscura). —Cockerell, 1929. Ent.
Soe. Amer., Ann. 22: 757. 2 (as obscura). —LaBerge, 1973. Amer. Ent. Soc., Trans. 99:
273-277, figs. 1, 6, 28-29 (redescription, synonymy).

Biology: LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 277 (floral relationships).

winnemuccana LaBerge. Nev., Oreg. Pollen: Unknown, but visits flowers of Thelypodium
laciniatum.
Andrena (Trachandrena) winnemuccana LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 366,
fig lee
zionensis LaBerge. South. Utah. Pollen: Unknown, but visits flowers of Amelanchier, Saliz.
Andrena (Trachandrena) zionensis LaBerge, 1973. Amer. Ent. Soc., Trans. 99: 361, figs. 1,
68-69. 2, 3.

Genus ANDRENA Subgenus TYLANDRENA LaBerge

Andrena subg. Tylandrena LaBerge, 1964. Nebr. Univ. State Mus., Bul. 4: 312.
Type-species: Cilissa erythrogaster Ashmead. Orig. desig.

Revision: LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 543-605, 75 figs., 5 maps
(N. Amer. spp.).
coracina LaBerge and Bouseman. South. Calif. (Kramer Hills).
Andrena (Tylandrena) coracina LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96:
591, figs. 1, 61-65. 2, d.
erythrogaster (Ashmead). Que. to B. C., south to N. C., Ga., Ark., Okla., N. Mex. and Utah.
Pollen: Apparently an oligolege of Salix including S. amygdaloides, S. bebbiana, S.
cordata, S. humilis, S. interior, S. longifolia, S. nigra, S. sericea, but visits other flowers
presumably for nectar including Amelanchier canadensis, Anemone canadensis,
Antennaria plantaginifolia, Aronia, Claytonia virginica, Crataegus coccinea, C.
crus-galli, Descourainia intermedia, Heracleum lanatum, Lomatium foeniculaceum,
Prunus americana, P. serotina, P. virginiana, Pyrus, Radicula obtusa, Rhamnus
lanceolata, Rhus aromatica, R. canadensis, R. trilobata, Ribes, Rubus, Shepherdia
argentea, Spiraea, Taraxacum officinale, Viburnum prunifolium, Zizia aurea.
Cilissa erythrogaster Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 6. d.
Andrena perezi Robertson, 1891. Amer. Ent. Soc., Trans. 18: 51. 2, d.
Andrena rhodura Cockerell, 1898. Ent. News 9: 171. °.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 190, 192. 2, 6 (key). —Lanham,
1941. Ent. Soc. Amer., Ann. 34: 704, 707. 2°, 6 (key). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 114-115, figs. 18-19 21, table 3 (redescription). —LaBerge and Bouseman,
1970. Amer. Ent. Soc., Trans. 96: 550-556, figs. 1-2, 6-10 (redescription).

Biology: Rau, 1935. Ent. News 45: 35 (nesting habits, floral relationships).
fuscipennis LaBerge and Bouseman. Tex. (Kerrville). Pollen: Unknown, but visits flowers of
Prunus.
Andrena (Tylandrena) fuscipennis LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans.
96: 564, figs. 1, 21-25. 9, d.
hallii Dunning. Nebr. and Kans., west to Wash. and Nev. Pollen: Apparently an oligolege of
cruciferous flowers, especially Stanleya pinnata, but also visits flowers of Brassica
nigra, Malvastrum coccineum, Melilotus, Opuntia, Penstemon, Prunus, Romneya,
Sisymbrium linifolium.
Andrena Hallii Dunning, 1898. Canad. Ent. 30: 268. 2.
Andrena heterura Cockerell, 1930. N. Y. Ent. Soc., Jour. 37: 445. °.

Taxonomy: Viereck, 1904. Canad. Ent. 36: 191. ° (key). —Cockerell, 1931. Amer. Mus.
Novitates 458: 10. 9. —LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 588-591,
figs. 1, 4, 56-60 (redescription, synonymy).

hurdi Lanham. Calif. Pollen: Unknown, but visits flowers of Brassica, Layia. A. waldmerei
LaBerge and Bouseman is possibly a subspecies or synonym of this species which is
perhaps the most elegant N. Amer. species of the genus Andrena.
Andrena hurdi Lanham, 1949. Pan-Pacific Ent. 25: 33. 9, ¢.

1842 Hymenoptera in America North of Mexico

Taxonomy: LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 583-584, figs. 1, 46-50
(redescription).
jessicae Cockerell. N. Mex., Ariz. Pollen: Unknown, but visits flowers of Lesquerella gordonii.
Andrena jessicae Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 79. 6.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48 (key). —LaBerge and
Bouseman, 1970. Amer. Ent. Soce., Trans. 96: 571-574, figs. 1, 3, 31-35 (redescription).
layiae Timberlake. Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of

Brassica, Lasthenia chrysostoma, L. minor, Layia glandulosa, Senecio californicus.
Andrena (Ptilandrena) layiae Timberlake, 1951. U.S. Natl. Mus., Proc. 101: 381. 9.

Taxonomy: LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 594-597, figs. 1, 3,
66-70 (redescription).
mesillae Cockerell. Tex. to Ariz. Pollen: Apparently an oligolege of Lesquerella including L.
gordonii, L. gracilis, but visits other flowers including Descourainia sophia, Phacelia,
Physaria, Senecio salignus.
Andrena mesillae Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 90. 6.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 49. ¢. (key). —LaBerge and
Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 584-588, figs. 1, 4, 51-55 (redescription).

perplexa Smith. Occupies a disjunct range; in the east occurs from Ont. and Conn., south to

Ga., west to Minn. Iowa, Okla. and Tex.; in the west occurs from Colo. and Mont., west
to B. C., Wash., Oreg. and north. Calif. (Shasta County). Pollen: Apparently polylectic,
visits flowers of Acer, Amelanchier, Aronia, Aruncus, Barbarea, Brassica, Crataegus,
Crepis runcinata, Euonymus japonica, Fragaria virginiana, Geranium, Heracleum,
Lomatium dissectum, Malus, Melilotus, Prunus americana, P. demissa, P.
pennsylvanica, P. virginiana, Ptelea, Pyracantha, Pyrus, Rhamnus, Rhododendron,
Rubus deliciosus, Salix, Senecio, Spiraea, Stellaria pubera, Stenophragma thaliana,
Taraxacum officinale, Viburnum.

Andrena perplexa Smith, 1853. Cat. Hym. Brit. Mus. v. 1, p. 148. 2.

Andrena belfragei Cresson, 1872. Amer. Ent. Soc., Trans. 4: 256. @.

Andrena brunniventris Cresson, 1872. Amer. Ent. Soc., Trans. 4: 258. 6.

Andrena viburnella Graenicher, 1903. Canad. Ent. 35: 165. 2, 6.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 150 (type). —Viereck, 1904. Canad.
Ent. 36: 189, 194. 2, ¢ (key, as viburnella). —Cockerell, 1906. Psyche 13: 10 (type).
—Viereck, 1907. Ent. News 18: 365 (as viburnella). —Lanham, 1941. Ent. Soe. Amer.,
Ann. 34: 705, 708. 2, ¢ (as viburnella). —Michener, 1947. Amer. Midland Nat. 38: 445. 9,
3 (as viburnella). —Lanham, 1949. Ent. News 60: 67 (as belfragei). —Mitchell, 1958. In
Krombein, U.S. Dept. Agr., Agr. Monog 2, Suppl. 1: 215 (synonymy). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 116-117, figs. 18-19, 21, table 3 (redescription).
—Stephen, 1966. Kans. Ent. Soc., Jour. 39: 51-53, fig. 1 (larva, pupa, as viburnella).
—LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 559-564, figs. 1, 3, 16-20
(redescription, synonymy).

Biology: Parker and Boving, 1924. U. S. Natl. Mus., Proc. 64: 1 (nesting habits). —Stephen,
1966. Kans. Ent. Soc., Jour. 39: 42-51, 2 figs. (nest architecture, life history, as viburnella).
subaustralis Cockerell. B. C. to Calif., east to Idaho, Utah and east. N. Mex. Pollen: Oligolege

of Salix including S. gooddingii, S. hindsiana, S. laevigata, S. lasiolepis, S. nigra, but
visits other flowers for nectar including Baccharis viminea, Capsella bursa-pastoris,
Descourainia sophia, Hackelia floribunda, Malus, Prunus americanus, Pyrus, Ribes
aureum, R. inerme, Senecio salignus, Sisymbrium irio, Tamarix, Taraxacum officinale.
Predator: Philanthus solivagus Say.

Andrena subaustralis Cockerell, 1898. Canad. Ent. 30: 146. 2, 6.

Andrena viridinitens Cockerell, 1936. Pan-Pacifie Ent. 12: 152. °.

Taxonomy: LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 556-559, figs. 1, 2,
11-15 (redescription, synonymy).

Superfamily APOIDEA 1843

sublayiae LaBerge and Bouseman. Cent. Calif. Pollen: Unknown, but visits flowers of
Hemizonia, Layia.
Andrena (Tylandrena) sublayiae LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans.
96: 597, figs. 1, 3, 71-75. 2, 6.
subtilis Smith. B. C. to Calif., east to Mont., Wyo. and Colo. Pollen: Polylectic, visits flowers of
Acer macrophylla, Baccharis viminea, Brassica campestris, Calandrinia caulescens,
Caragana, Conium maculatum, Crepis, Gilia, Grossularia, Hackelia floribunda,
Lasthenia, Limnanthes douglasii var. douglasii, Lomatium dissectum, Prunus demissa,
P. subcordata, Purshia tridentata, Pyracantha, Pyrus, Ranunculus californicus, Rosa,
Salix lasiolepis, Senecio, Spiraea, Tamarix gallica, Taraxacum.
Andrena subtilis Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 55. 2.
Andrena seminigra Viereck, 1904. Canad. Ent. 36: 190. 9.
Andrena semicyanea Cockerell, 1924. Pan-Pacifie Ent. 1: 58. °.

Taxonomy: Morice and Cockerell, 1901. Canad. Ent. 33: 155 (type). —Cockerell, 1906. Psyche
13: 6 (type). —LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 574-580, figs. 1, 5,
36-40 (redescription, synonymy).

Biology: Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 19 (floral relationships).
waldmerei LaBerge and Bouseman. South. Calif. Pollen: Unknown, but visits flowers of
Allium haematochiton, Encelia californica, Lomatium dasycarpum, Rhus trilobata.
Possibly a subspecies or synonym of A. hurdi Lanham.
Andrena (Tylandrena) waldmerei LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans.
96: 580, figs. 1, 41-45. 2, 6.
wilmattae Cockerell. Pa. to N. C., west to Wyo. and Colo. Pollen: Unknown, but visits flowers
of Amorpha canescens, Crataegus, Crepis runcinata, Erigeron, Melilotus alba, M.
officinale, Prunus angustifolia, Spiraea vanhouteti, Pastinaca sativa, Viburnum
lentago.
Andrena Wilmattae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 224. °.
Andrena subaustraliformis Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 28. d.
Andrena beutenmuelleri Viereck, 1916. Amer. Mus. Nat. Hist., Bul. 35: 729. °.
Andrena (Bythandrena) acra Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 111, figs.
18,19) 21-0:

Taxonomy: LaBerge and Bouseman, 1970. Amer. Ent. Soc., Trans. 96: 567-571, figs. 1, 4,
26-30 (redescription, synonymy).

Genus ANDRENA Subgenus XIPHANDRENA LaBerge

Andrena subg. Xiphandrena LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 504.
Type-species: Andrena (Trachandrena) mendica Mitchell. Monotypic and orig.

desig.
Revision: LaBerge, 1971. Amer. Ent. Soc., Trans. 97: 504-507, figs. 1, 4, 54-58.
mendica Mitchell. Ill., Ohio, Tenn., Va., N. C., Ga. Pollen: Unknown, but visits flowers of
Aruncus aruncus, Ceanothus including C. americanus.
Andrena (Trachandrena) mendica Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
189, figs. 35-37, 40. 2, d.

Genus ANDRENA Subgenus UNASSIGNED

Although some of these species have been assigned and even reassigned to some of the sub-
genera in Andrena, their correct assignments must await further studies of the genus.

agricolarum Viereck and Cockerell. Colo.
Andrena agricolarum Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 29. ¢.

albuginosa (Viereck). Oreg.
Pterandrena albuginosa Viereck, 1904. Canad. Ent. 36: 227, 228. °.

angustella angustella Cockerell. Calif.
Andrena angustella Cockerell, 1936. Pan-Pacific Ent. 12: 136. d.

1844 Hymenoptera in America North of Mexico

angustella quercina Cockerell. Calif. (Live Oak Canyon, Redlands). Pollen: Unknown, but

visits flowers of Ceanothus, Salix.
Andrena angustella quercina Cockerell, 1939. Ann. and Mag. Nat. Hist. (11) 3: 183. 6.

angustifovea Viereck. B. C. to Oreg.
Andrena angustifovea Viereck, 1904. Canad. Ent. 36: 194. 6.

atala Viereck. N. Mex.
Andrena atala Viereck, 1903. Amer. Ent. Soc., Trans. 23: 54. 2.
Taxonomy: Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 430. 2, 6.

azygos Viereck. N. Mex.
Andrena (Andrena) azygos Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 550. d.

banksi Malloch. N. J. to Fla., west to Tex. Pollen: Unknown, but visits flowers of Prunus
angustifolia.
Andrena banksi Malloch, 1917. Brooklyn Ent. Soce., Bul. 12: 89. d, 2.
Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 254-255, figs. 51, 54, 56-57,
table 7 (redescription).
caeruleonitens Viereck. Calif.
Andrena caeruleonitens Viereck, 1926. Pomona Jour. Ent. Zool. 18: 1. 6.
casadae Cockerell. N. Mex., Colo.
Andrena casadae Cockerell, 1896 Ann. and Mag. Nat. Hist. (6) 18: 83. 3.
Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48. ¢ (key).
cerebrata Mitchell. N. C., Ark. Pollen: Unknown, but visits flowers of Malus, Philadelphus,

Raphanus sativa, Spiraea.
Andrena cerebrata Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 255, figs. 51, 56, 57.

3.

Taxonomy: Chandler, 1966. Pan-Pacifie Ent. 42: 89-90. °.
cragini Cockerell. Kans.
Andrena cragini Cockerell, 1899. Ent. News 10: 254. ¢.
dallasiana Cockerell. Tex.
Andrena dallasiana Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 262.
didelta Viereck. N. Mex.
Andrena delta Viereck, 1903. Amer. Ent. Soc., Trans. 29: 55. 9. Preoce.
Andrena didelta Viereck, 1908. Ent. News 19: 42. N. name.
dimorpha Mitchell. N. J. N. C., Fla.
Andrena (Pterandrena) dimorpha Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 148,
figs. 24-26, 28. 2, 6.
Taxonomy: LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 294 (subgeneric status).
dolomellea Lanham. Tex. Pollen: Unknown, but visits flowers of Crataegus.
Andrena dolomellea Lanham, 1949. Ent. News 60: 65. 2, 6.
dreisbachi Mitchell. Mich.
Andrena (Gonandrena) dreisbachi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 2380.
De

Taxonomy: Knerer and Atwood, 1963. Canad. Ent. 95: 586, figs. 6, 8. ¢.

elongatula Viereck. Calif.
Andrena (Andrena) elongatula Viereck, 1917. Amer. Ent. Soe., Trans. 43: 378. d.
enceliarum Cockerell. Ariz. Pollen: Unknown, but visits flowers of Encelia farinosa.
Andrena enceliarwm Cockerell, 1937. Amer. Mus. Novitates 948: 13. 6.
enocki (Cockerell). Calif.
Parandrena enocki Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 189. o.
hirsutula Cockerell. N. W. T. (Hay River).
Andrena hirsutula Cockerell, 1936. Canad. Ent. 68: 282. °.

inclinata Viereck. Calif.
Andrena (Andrena) inclinata Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 559. 9.

Superfamily APOIDEA = 1845

interrogationis Viereck and Cockerell. Colo.

Andrena interrogationis Viereck and Cockerell, 1914. U. S. Natl. Mus., Proce. 48: 35. d.
jennei Viereck. Wash.

Andrena (Andrena) jennei Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 561. d.

lauracea Robertson. II]. Pollen: Unknown, but visits flowers of Sassafras variifolium.
Andrena lauracea Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 331. 9.
Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 194. ?. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 148 (redescription).
littlefieldi Viereck. Colo.
Andrena (Andrena) littlefieldi Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 563. 3.
mellitarsis Viereck. Nev.
Andrena (Andrena) mellitarsis Viereck, 1917. Amer. Ent. Soe., Trans. 43: 384. d.
microsoma Viereck. Pacific Northwest.
Andrena microsoma Viereck, 1904. Canad. Ent. 36: 194. 6.

monilicornis Cockerell. N. Mex.
Andrena monilicornis Cockerell, 1896. Canad. Ent. 28: 181. 6.

Taxonomy: Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 48. ¢ (key).

moquiorum Viereck and Cockerell. Ariz.

Andrena moquiorum Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 21. d.
mustelicolor Viereck. Idaho, Wash. to Calif.

Andrena mustelicolor Viereck, 1904. Canad. Ent. 35: 189, 195. 2, 6.
navajorum Viereck and Cockerell. N. Mex.

Andrena navajorum Viereck and Cockerell, 1914. U.S. Natl. Mus., Proc. 48: 29. d.
neurona Viereck. B. C., Wash.

Andrena neurona Viereck, 1904. Canad. Ent. 36: 191. 9.

nigerrima Casad. N. Mex.
Andrena nigerrima Casad, 1896. Ann. and Mag. Nat. Hist. (6) 18: 83. 2.

Taxonomy: Linsley, 1938. Calif. Acad. Sci., Proc. (4) 23: 265, 272. 2 (key).
nothoscordi Robertson. III. (Carlinville). Pollen: Unknown, but visits flowers of Nothoscordum

bivalve.
Andrena nothoscordi Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 331. 9.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 191. 2 (key). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 165 (redescription of type). —Ribble, 1968. Nebr. Univ.
State Mus., Bul. 8: 333 (subgeneric position).
nubilifascia Viereck. Calif.

Andrena (Ptilandrena) nubilifascia Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 597. ?.
obscuripostica Viereck. Nev.

Andrena obscuripostica Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 568. °.
occidentalis Cockerell. N. Mex.

Andrena platyparia occidentalis Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 87. ¢.
oniscicolor (Viereck). Oreg.

Pterandrena oniscicolor Viereck, 1904. Canad. Ent. 36: 227, 228. °.
paduocorum Viereck and Cockerell. Colo.

Andrena paduocorum Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 38. ¢.
pediculihirta Viereck. Calif.

Andrena (Ptilandrena) pediculihirta Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 598. 2.
perindotata Viereck. Wash.

Andrena indotata Viereck, 1904. Canad. Ent. 36: 190. 2. Preoce.

Andrena perindotata Viereck, 1908. Ent. News 19: 42. N. name.
pineti Cockerell. Wyo.

Andrena nigerrima pineti Cockerell, 1931. Amer. Mus. Novitates 458: 13. °.

1846 Hymenoptera in America North of Mexico

plumiscopa Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Cryptantha
intermedia, Platystemon californicus.
Andrena (Pterandrena) plumiscopa Timberlake, 1951. U.S. Natl. Mus., Proe. 101: 376. °,
3.
Taxonomy: LaBerge, 1967. Nebr. Univ. State Mus., Bul. 7: 294 (subgeneric position).
polygoni Viereck and Cockerell. Colo.
Andrena polygoni Viereck and Cockerell, 1914. U.S. Natl. Mus., Proce. 48: 18. d.
potentillarum Viereck. Alta. to Colo. Pollen: Unknown, but visits flowers of Potentilla.
Andrena (Andrena) potentillarum Viereck, 1924. Canad. Ent. 56: 79. 2.
pulverea Viereck. Calif.
Andrena (Andrena) pulverea Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 569. 2.
purpurina Viereck and Cockerell. Colo.
Andrena purpurina Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 16. 9.
robertsonii Dalla Torre. Colo. and Minn. to N.S., south to Ark. and Ga. Pollen: Unknown, but
visits flowers of Apocynum, Aronia, Aruncus, Brassica, Ceanothus, Cryptotaenia,
Pastinaca, Ptelea, Pyracantha, Rhus, Rubus, Taenidia, Viburnum. Predator:
Philanthus albopilosus Cress.
Andrena serotina Robertson, 1893. Amer. Ent. Soc., Trans. 20: 148. 2 (¢ misdet.). Preocc.
Anthrena robertsonii Dalla Torre, 1896. Cat. Hym., v. 10, p. 149. N. name.

Taxonomy: Robertson, 1902. Amer. Ent. Soc., Trans. 28: 193. 2, ¢ (key). —Atwood, 1934.
Canad. Jour. Res. 10: 208, 209. 2, d (key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 237-238, figs. 46-47, 49, table 6 (redescription).

Biology: Evans, 1975. Ent. Soc. Amer., Ann. 68: 891 (predator).

rufojugata Cockerell. Wyo.
Andrena rufojugata Cockerell, 1931. Amer. Mus. Novitates 458: 17. 9.

sancta Viereck. Calif.

Andrena (Andrena) sancta Viereck, 1916. Acad. Nat. Sci. Phila., Proc. 68: 571. 2.
scotoptera Cockerell. Tex.

Andrena scotoptera Cockerell, 1934. Amer. Mus. Novitates 732: 1. °.
stenosoma Viereck. Nev.

Andrena (Andrena) stenosoma Viereck, 1925. Amer. Ent. Soc., Trans. 51: 135. ¢.

stictigastra Viereck. Calif.
Andrena (Andrena) stictigastra Viereck, 1917. Acad. Nat. Sci. Phila., Proc. 68: 579. °.

Taxonomy: Linsley and MacSwain, 1961. Pan-Pacific Ent. 37: 129 (subgeneric position).

subarctica Cockerell. Alta. (Hay River).

Andrena subarctica Cockerell, 1936. Canad. Ent. 68: 282. °.
tetonorum Viereck and Cockerell. Nebr.

Andrena tetonorum Viereck and Cockerell, 1914. U. S. Natl. Mus., Proc. 48: 24. 3.
trivialis Viereck. Calif.

Andrena (Andrena) trivialis Viereck, 1917. Amer. Ent. Soc., Trans. 43: 388. 2.
variantia Linsley. B. C.

Andrena (Andrena) varia Viereck, 1924. Canad. Ent. 56: 81. 2. Preoce.

Andrena variantia Linsley, 1951. U. S. Dept. Agr., Agr. Monog. 2: 1084. N. name.
vicinoides Viereck. B. C., Wash.

Andrena vicinoides Viereck, 1904. Canad. Ent. 36: 191. °.

NOMINA NUDA IN ANDRENA FABRICIUS

Andrena alienoides (Viereck) Smith, 1910. Ins. New Jersey, p. 690.
Andrena delawarearum (Viereck). Smith, 1910. Ins. New Jersey, p. 690.
Andrena sparsipilosa Pierce, 1908. U. S. Natl. Mus., Bul. 66: 108.
Andrena tuberculata (Ashmead) Smith, 1910. Ins. New Jersey, p. 690.

Superfamily APOIDEA 1847
Genus MEGANDRENA Cockerell

Genus MEGANDRENA Subgenus MEGANDRENA Cockerell

Andrena subg. Megandrena Cockerell, 1927. Pan-Pacific Ent. 4: 42.
Type-species: Andrena (Megandrena) enceliae Cockerell. Monotypic.
enceliae (Cockerell). Ariz., south. Calif., deserts. Pollen: Oligolectie on flowers of Larrea
tridentata, but visits other flowers for nectar including Argemone platyceros, Aster,
Cirsium, Encelia californica, E. farinosa, Eschscholzia darwinensis, Geraea canescens,
Ferrocactus ancanthodes, Opuntia basilaris, O. echinocarpa.
Andrena (Megandrena) enceliae Cockerell, 1927. Pan-Pacific Ent. 4: 43. °.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 23-24, table 14 (floral
relationships, sleep).

Genus MEGANDRENA Subgenus ERYTHRANDRENA Zavortink

Megandrena subg. Erythrandrena Zavortink, 1972. Ent. Soc. Wash., Proc. 74: 61.
Type-species: Megandrena (Erythrandrena) mentzeliae Zavortink. Monotypie and
orig. desig.
mentzeliae Zavortink. Nev. (Spring Mts., northwest of Las Vegas). Pollen: Oligolectic on
flowers of Mentzelia tricuspis.
Megandrena (Erythrandrena) mentzeliae Zavortink, 1972. Ent. Soc. Wash., Proc. 74: 64,
figs. 1-3. 2, 6.

Biology: Zavortink, 1972. Ent. Soc. Wash., Proc. 74: 69-75, figs. 2-3 (floral relationships,
mating, sleep).

Genus ANCYLANDRENA Cockerell

Andrena subg. Ancylandrena Cockerell, 1930. Pan-Pacifie Ent. 7: 5.
Type-species: Andrena (Ancylandrena) heterodoxa Cockerell. Monotypic.

Revision: Zavortink, 1974. Calif. Acad. Sci., Occas. Papers, 109: 1-36, 6 figs. (N. Amer. spp.).
atoposoma (Cockerell). South. Calif.; Mexico (Baja California and Sonora). Pollen: Apparently
collects pollen from the flowers of the leguminous genera Lotus and Lupinus including
Lotus glaber, L. scoparius, Lupinus formosus, L. hallii, L. paynei, but visits these and
other flowers for nectar including Calochortus splendens, Ceanothus, Convolvulus,
Cryptantha intermedia, Eriodictyon trichocalyx, Eriogonum fasciculatum, Gilia
virgata, Penstemon, Phacelia ramosissima, Sphaeralcea ambigua.
Andrena (Ancylandrena) heterodoxa Cockerell, 1930. Pan-Pacifie Ent. 7: 6. 6, 2. Preoce.
Andrena atoposoma Cockerell, 1934. Pan-Pacific Ent. 10: 82. N. name.
koebelei (Timberlake). Southeast. Calif. and south. Nev. Pollen: Possibly oligolectic on flowers
of Dalea including D. fremontii, but visits other flowers presumably for nectar including
Echinocactus, Oenothera refracta, Stanleya pinnata.
Megandrena (Ancylandrena) koebelei Timberlake, 1951. U. S. Natl. Mus., Proce. 101: 413.
é.

Biology: Zavortink, 1974. Calif. Acad. Sci., Occas. Papers, 109: 28 (matinal behavior).
larreae Timberlake. N. Mex. (Lordsburg), west to south. Calif. and south. Nev., deserts.
Parasite: Hexepeolus rhodogyne Linsley and Michener? Pollen: Oligolectic on flowers of
Larrea tridentata, but visits these flowers and others for nectar including Dalea greggii,
Prosopis juliflora.
Megandrena (Ancylandrena) larreae Timberlake, 1951. U. S. Natl. Mus., Proc. 101: 411. 2,
Or
Biology: Zavortink, 1974. Calif. Acad. Sci., Occas. Papers 109: 23 (floral relationships).
—Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 22-23, fig. 4, tables 2, 9, 11, 12-14
(floral relationships, parasite).
timberlakei Zavortink. Ariz. to south. Calif. and south. Nev. Pollen: Apparently polylectic,
visits flowers of Dalea fremontii, Eriodictyon trichocalyx, Hyptis emoryi, Larrea

1848 | Hymenoptera in America North of Mexico

tridentata, Lepidium fremontii, Malacothrix, Mentzelia involucrata, M. tricuspis,
Phacelia, Sphaeralcea ambigua, Verbesina.

Ancylandrena timberlakei Zavortink, 1974. Calif. Acad. Sci. Occas. Papers 109: 16, figs. 1,
aCe o:

Biology: Zavortink, 1974. Calif. Acad. Sci., Occas. Papers 109: 19 (floral relationships).

SUBFAMILY PANURGINAE

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 1-22, 54 figs. (larvae). —Yager and
Rozen, 1966. Amer. Mus. Novitates 2265: 6-13, figs. 7-18 (pupae). —Rozen, 1970. Amer.
Mus. Novitates 2416: 1-16, 19 figs. (immature stages).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 1-44, 18 figs. (review of N. Amer. spp.).
Genus PROTANDRENA Cockerell

Protandrena Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 91.
Type-species: Andrena maurula Cockerell. Desig. by Sandhouse, 1943.
Austrandrena Timberlake, 1906. Psyche 13: 37.
Type-species: Andrena modesta Smith. Monotypic and orig. desig.
Protoandrena Cresson, 1928. Amer. Ent. Soc., Mem. 5: 58. Emend.

Revision: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 133-227, 93 figs. (N. Amer. spp.).

Taxonomy: Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 92 (tax. characters). —Cockerell,
1896. Canad. Ent. 28: 184 (tax. characters). —Timberlake, 1955. Bol. Lab. Zool. Gen. Agr.
33: 398-409 (key to females, as Psaenythia). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 286-287, figs. 58, 67-68, table 8 (east. U. S. spp.). —Timberlake, 1973. Calif. Univ.
Pubs. Ent. 72: 1 (tax. status).
amplipennis Timberlake. Ariz. (10 mi. S.W. of Patagonia). Pollen: Unknown, but visits flowers
of Croton.
Protandrena amplipennis Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 146. ?.
bancrofti Dunning. Ariz., N. Mex., Tex., Kans., Mo., Okla., Nebr., Colo., Wyo., Iowa, N. Dak.,
Ind., Ill., Wis. Pollen: Unknown, but visits flowers of Amorpha, Asclepias, Astragalus,
Baccharis, Eysenhardtia, Guardiola, Helianthus, Heterotheca, Lepidium, Medicago,
Melilotus, Monarda, Petalostemon, Psoralea, Solanum, Symphoricarpos.
Protandrena Bancrofti Dunning, 1897. Canad. Ent. 29: 264. 2.

Taxonomy: Cockerell, 1921. Amer. Mus. Novitates 24: 13. 2. —Timberlake, 1976. Amer. Ent.
Soc., Trans. 102: 150, figs. 4, 5, 6 (geogr. and floral records).

Biology: Chandler, 1962. Kans. Ent. Soc., Jour. 35: 314 (nest).
bicolor (Timberlake). Tex. to Ariz.; Mexico (Coahuila, Durango). Pollen: Collects pollen from
flowers of the composite genus Bahia, but also visits flowers of Verbesina.
Psaenythia bicolor Timberlake, 1955. Bol. Lab. Zool. Gen. Agr. 33: 401. °.

Taxonomy: Rozen, 1970. Amer. Mus. Novitates 2416: 9-13, figs. 9-17 (larva, pupa).
—Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 150-152, figs. 7, 8, 9. 3.

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 17-19, fig. 6 (nesting activity, nest
architecture, floral relationships). —Rozen, 1970. Amer. Mus. Novitates 2416: 6-7, fig. 2
(nest architecture, nesting habits).

bishoppi Crawford. Tex.
Protandrena bishoppi Crawford, 1916. Ent. Soc. Wash., Proc. 19: 128. 9.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 152-153 (notes on type).
cockerelli Dunning. N. J., Ind., Ill., Iowa, Mich., Nebr., Mo., Kans., Colo., N. Mex., Tex. Pollen:
Unknown, but visits flowers of Acerates, Amphiachyris, Asclepias, Cassia, Euphorbia

corollata, Gaillardia, Melilotus, Monarda, Petalostemon, Solanum, Teucriuwm.
Protandrena Cockerelli Dunning, 1897. Canad. Ent. 29: 47. °.

Superfamily APOIDEA 1849

Taxonomy: Cockerell, 1899. Ent. News 10: 3. 6. —Timberlake, 1955. Bol. Lab. Zool. Gen.
Agr. 33: 400 (tax. status). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 287 (tax.
characters and status). —Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 156-157 (tax.
characters and status).

cognata Timberlake. Ariz. (Cochise County).
Protandrena cognata Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 157. °.
duplonotata (Timberlake). Ariz. (Pinal Co.).; Mexico (Sonora). Pollen: Unknown, but visits
flowers of Hymenothrix.
Psaenythia duplonotata Timberlake, 1955. Bol. Lab. Zool. Gen. Agr. 33: 407. °.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 159 (geogr. and floral record).
euphorbiae (Timberlake). Ariz. (Hope and Picacho Pass); Mexico (Sonora). Pollen: Unknown,

but visits flowers of Euphorbia, including E. capitellata.
Psaenythia euphorbiae Timberlake, 1955. Bol. Lab. Zool. Gen. Agr. 33: 406. °.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 161-162, figs. 16, 17, 18. 3.
heteromorpha (Cockerell). N. Mex., Ariz. Pollen: Unknown, but visits flowers of Aplopappus

gracilis, Gutierrezia microcephala, Heterotheca subaxillaris, Verbesina encelioides.
Andrena heteromorpha Cockerell, 1896. Canad. Ent. 28: 180. 9, 3.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 168-171, figs. 25, 26, 27
(redescription).
hurdi Timberlake. N. Mex. (Rodeo and 25 miles east of Steins), Ariz. (Chiricahua and
Huachuca Mts.). Pollen: Unknown, but visits flowers of Asclepias subverticillata.
Protandrena hurdi Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 171, figs. 28, 29, 30. 3.
kansensis Timberlake. Kans. (Douglas and Greenwood Counties). Pollen: Unknown, but visits
flowers of Amphiachyris dracunculoides.
Protandrena kansensis Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 175, figs. 34, 35, 36.
OPIGe
leucopus Timberlake. Ariz. (Cochise and Pima Counties).
Protandrena leucopus Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 178, figs. 40, 41, 42.
é.
maurula (Cockerell). Tex.
Andrena maurula Cockerell, 1896. Canad. Ent. 28: 179. °.
mexicanorum (Cockerell). N. Dak., Nebr. Colo. and Tex., west to Ariz.; Mexico (Chihuahua,
Coahuila, Durango and Sonora). Pollen: Unknown, but visits flowers of Asclepias,
Kallstroemia grandiflora, Solanum. Protandrena durangoensis Timberlake is possibly
a subspecies of this.
Andrena mexicanorum Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 85. 9
Andrena asclepiadis Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 86. 6.

Taxonomy: Timberlake, 1955. Bol. Lab. Zool. Gen. Agr. 33: 400 (synonymy, tax. status, flower
records). —Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 188-189, figs. 55, 56, 57 (tax.
characters and status).

pectidis (Timberlake). Ariz., N. Mex.; Mexico (Sonora). Pollen: Unknown, but visits flowers of
Pectis papposa, Viguiera annua.

Psaenythia pectidis Timberlake, 1955. Bol. Lab. Zool. Gen. Agr. 33: 405. 2.
Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 191-192, figs. 58, 59, 60. 3.

sorocula Timberlake. N. Mex. (7 miles southeast of Rodeo).

Protandrena sorocula Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 205. °.
sphaeralceae Timberlake. Ariz. (Cochise County). Pollen: Unknown, but visits flowers of

Sphaeralcea.

Protandrena sphaeralceae Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 207. °.
swenki Crawford. Nebr.

Protandrena swenki Crawford, 1913. U.S. Natl. Mus., Proce. 45: 241. 2.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans: 102: 191-192, figs. 58, 59, 60. ¢.

1850 Hymenoptera in America North of Mexico

texana Timberlake. Tex. (Archer, Gillespie and Throckmorton Counties). Pollen: Unknown, but
visits flowers of Teucriwm laciniatum.
Protandrena texana Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 211, figs. 76, 77, 78. d.
tidestromiae Timberlake. N. Mex., Ariz. Pollen: Collects pollen from flowers of Tidestromia
including T. lanuginosa, but visits other flowers presumably for nectar including
Verbesina.
Protandrena tidestromiae Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 212, figs. 79, 80,
Be M5 Ge
trifoliata (Cockerell). N. Mex., Ariz. Pollen: Unknown, but visits flowers of Baccharis
glutinosa, Euphorbia capitellata, Solanum, Trifolium, Verbesina.
Andrena trifoliata Cockerell, 1896. Canad. Ent. 28: 179. @.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 215-217, figs. 82, 83, 84. 3.
verbesinae (Timberlake). West Tex. to Ariz.; Mexico (Coahuila and Durango). Pollen:
Unknown, but visits flowers of Asclepias, Baileya, Heterotheca, Pectis, Verbesina

encelioides.
Psaenythia verbesinae Timberlake, 1955. Bol. Lab. Zool. Gen. Agr. 33: 402. 2, d.

Taxonomy: Timberlake, 1976. Amer. Ent. Soc., Trans. 102: 222-224, figs. 88, 89, 90 (geogr. and
floral records).

Genus PANURGINUS Nylander

Panurginus Nylander, 1848. Notiser Sallskapet Flora Fauna Fenn. 1: 223.
Type-species: Panurginus niger Nylander. Monotypic.
Greeleyella Cockerell, 1904. Entomologist 37: 235.
Type-species: Greeleyella beardsleyi Cockerell. Monotypic and orig. desig.
Birkmania Viereck, 1909. Ent. Soc. Wash. Proce. 11: 50.
Type-species: Panurginus polytrichus Cockerell. Monotypic and orig. desig.
(=Birkmania andrenoides Viereck).

Revision: Crawford, 1926. Ent. Soc. Wash., Proc. 28: 207-214. —Michener, 1935. Canad. Ent.
67: 275-278 (species with blackfaced males).

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 5, 10-12, figs. 17-28 (larvae).
armaticeps Cockerell. South. Calif. Pollen: Unknown, but visits flowers of Ceanothus
integerrimus, Cryptantha intermedia, Gilia integrifolia, Phacelia davidsonii, P. distans,
Potentilla glandulosa, Prunus demissa, Rhamnus crocea, Salix laevigata.
Panurginus armaticeps Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 280. “2” = 6.

Taxonomy: Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 322. 3, °.
atramontensis Crawford. Md. to Ga., Miss. Pollen: Unknown, but visits flowers of Polycodium.
Panurginus atramontensis Crawford, 1926. Ent. Soc. Wash., Proc. 28: 210. d, 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 259-260, figs. 59-61, table 8
(redescription).
atriceps (Cresson). Idaho, Wash., Oreg., Calif. Pollen: Collects pollen from flowers of Dowingia

bella or D. cuspidata, but also visits flowers of Ceanothus dentatus, C. integerrimus,
Cryptantha intermedia, Lasthenia chrysostoma, Montia perfoliata, Nemophila
maculata, N. pulchella, Ranunculus, Rhamnus crocea, Rhus diversiloba, Tamarix
gallica. Predator: Philanthus pulcher Dalla Torre.

Calliopsis atriceps Cresson, 1878. Amer. Ent. Soc., Trans. 7: 67. d.

Taxonomy: Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 194. 2. —Cockerell, 1925. Calif. Acad.
Sci., Proc. (4) 14: 198. 6, 2. —Rust, 1976. Pan-Pacifie Ent. 52: 165, figs. 3, 10-16 (larva).

Biology: Rust, 1976. Pan-Pacific Ent. 52: 161-166, figs. 2-9 (nest site, nest architecture, cell
provisions, adult and immature habits).
beardsleyi (Cockerell). N. Dak., Nebr., Colo. Pollen: Unknown, but visits flowers of

Malvastrum.
Greeleyella beardsleyi Cockerell, 1904. Entomologist 37: 236. 9.

Superfamily APOIDEA 1851

Panurginus malvastri Swenk and Cockerell, 1907. Ent. News 18: 179. 6, 9.

Taxonomy: Cockerell, 1906. Canad. Ent. 38: 164. ¢.
bilobatus Michener. South. Calif. Pollen: Unknown, but visits flowers of Astragalus,
Cryptantha intermedia, Gilia integrifolia, Lasthenia gracilis, Phacelia distans, P.
hispida, Rhamnus crocea, R. ilicifolia, Salvia columbariae, Salix laevigata, Sisymbrium
irio.
Panurginus bilobatus Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 322. 3, 2.
ceanothi Michener. South. Calif. Pollen: Unknown, but visits flowers of Ceanothus.
Panurginus ceanothi Michener, 1935. Pan-Pacific Ent. 11: 180. 3.
cressoniellus Cockerell. Colo., N. Mex. Predator: Philanthus pulcher Dalla Torre.
Calliopsis clypeata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 67. 6. Preoce.
Panurginus cressoniellus Cockerell, 1898. Canad. Ent. 30: 29. N. name.
Panurginus cressoniellus calochorti Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 222.
OF
?Panurginus verus Cockerell, 1901. Psyche 9: 163. 9.

Taxonomy: Cockerell, 1897. Acad. Nat. Sci. Phila., Proce. 49: 350. 6, 2. —Viereck, 1903. Amer.
Ent. Soc., Trans. 29: 49. 6. —Cockerell, 1912. Ent. News 23: 445 (synonymy).
emarginatus Michener. Calif.
Panurginus emarginatus Michener, 1935. Pan-Pacific Ent. 11: 179. 6, °.
gabrielis Michener. South. Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers
of Arctostaphylos glauca, Ceanothus crassifolius, C. greggii, C. leucodermis, C.

verrucosus, Rhamnus californica, R. crocea.
Panurginus gabrielis Michener, 1935. Canad. Ent. 67: 276. 6.

Taxonomy: Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 321 (tax. characters).
gracilis Michener. Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Arctostaphylos pungens, Baccharis, Ceanothus crassifolius, C. verrucosus, Lasthenia
chrysostoma, Rhamnus crocea, Salix lasiolepis, Salvia mellifera.
Panurginus gracilis Michener, 1935. Canad. Ent. 67: 276. 6, 9.
ineptus Cockerell. Colo., Wash., Oreg., Utah.
Panurginus ineptus Cockerell, 1922. Amer. Mus. Novitates 36: 8. 2.
Panurginus bakeri Crawford, 1926. Ent. Soc. Wash., Proc. 28: 213. ¢. Preoce.
Panurginus rohweri Crawford, 1932. Ent. Soc. Wash., Proc. 34: 78. N. name.

Taxonomy: Timberlake, 1971. Pan-Pacific Ent. 47: 148 (tax. status, synonymy).

maritimus Michener. Calif.
Panurginus maritimus Michener, 1935. Pan-Pacific Ent. 11: 178. ¢.

Taxonomy: Michener, 1935. Canad. Ent. 67: 277. 2.
melanocephalus (Cockerell). Calif. Parasite: Zodion cinereiventre Van Duzee, Z. nigrifrons
Krober. Pollen: Collects pollen of Ranunculus californicus, but visits these and other
flowers for nectar.
Panurgus melanocephalus Cockerell, 1926. Pan-Pacific Ent. 3: 80. d, 9.
Panurginus morrisoni Crawford, 1926. Ent. Soc. Wash., Proc. 28: 209. d.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1036, figs. 86-88 (larva). —Rozen, 1966.
Amer. Mus. Novitates 2259: 12, figs. 21-23 (larva).

Biology: MacSwain and Bohart, 1947. Pan-Pacific Ent. 23: 30 (parasite). —Linsley and
MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 20-21, 28 pl. 6 (nest, life history, mating, sex
ratio, floral relationships, parasite).

nigrellus Crawford. Calif. Pollen: Unknown, but visits flowers of Arctostaphylos, Baccharis
glutinosa, Cryptantha intermedia, Lasthenia chrysostoma, Nemophila pulchella,
Ranunculus californicus, Salix lasiolepis, Tamarix gallica.
Panurginus nigrellus Crawford, 1926. Ent. Soe. Wash., Proc. 28: 210. ¢.

Taxonomy: Michener, 1935. Pan-Pacifie Ent. 11: 180. °.

nigrihirtus Michener. Calif.
Panurginus nigrihirtus Michener, 1935. Pan-Pacific Ent. 11: 179. 6, °.

1852 Hymenoptera in America North of Mexico

occidentalis (Crawford). Calif. Pollen: Collects pollen from flowers of Limnanthes douglasii.
Greeleyella occidentalis Crawford, 1916. Insecutor Inscitiae Menstruus 4: 105. d, 2.

Biology: Rust, 1976. Pan-Pacifie Ent. 52: 159, 165-166, fig. 1 (mating behavior, pollen and
nectar source).
polytrichus Cockerell. Miss., La., Tex.
Panurginus polytrichus Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 28. d.
Birkmania andrenoides Viereck, 1909. Ent. Soc. Wash., Proc. 11: 50. @.
Greeleyella resinata Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 361. °.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 260-262, figs. 59-61, table 8
(redescription).
potentillae (Crawford). N. J., Md., Va., N. C. Pollen: Collects pollen from the flowers of
Potentilla, but visits these and other flowers for nectar including Fragaria,
Ranunculus.
Greeleyella potentillae Crawford, 1916. Insecutor Inscitiae Menstruus 4: 104. 6, @.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 262-263, figs. 59-61, table 8
(redescription). —Rozen, 1966. Amer. Mus. Novitates 2259: 10-12, figs. 17-20 (larva).
—Rozen, 1967. Amer. Mus. Novitates 2297: 28-29, figs. 12-13, table 2 (egg, larva).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 27-30, tables 1-2 (nest architecture, life
history, floral relationship).

Genus PSPEUDOPANURGUS Cockerell

Pseudopanurgus Cockerell, 1897. Canad. Ent. 29: 290.
Type-species: Panurgus aethiops Cresson. Orig. desig.
Protandrenopsis Crawford, 1903. Canad. Ent. 35: 337.
Type-species: Panurgus aethiops Cresson. Monotypic and orig. desig.
(=Protandrenopsis fuscipennis Crawford).

Revision: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 1-58, 4 plates, 56 figs. (N. Amer. spp.).

Taxonomy: Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 195-198. —Cockerell, 1922. Amer.
Mus. Novitates 36: 9. —Robertson, 1922. Psyche 29: 164 (Illinois spp.). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 266-270, figs. 62, 65-66, table 8 (eastern U.S. spp.).
—Rozen, 1966. Amer. Mus. Novitates 2259: 5, 7-9, figs. 8-16 (larva). —Timberlake, 1967.
Amer. Mus. Novitates 2298: 1-10 (Ariz. spp.). —Timberlake, 1975. Calif. Univ. Pubs. Ent.
77: 1, 2 (key, tax. status).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 19-27, figs. 1-3, 6-8, tables 1-2 (nest
architecture, life history, floral relationships, parasites).
aethiops (Cresson). Nebr., Kans., Colo., Tex., N. Mex., Ariz. Pollen: Collects pollen from the

flowers of Compositae including Baileya, Encelia, Grindelia, Helianthus, Heterotheca,
Melampodium, Prionopsis, Verbesina encelioides, but visits these and other flowers for
nectar including Asclepias.

Panurgus aethiops Cresson, 1872. Amer. Ent. Soc., Trans. 4: 259. 2, d.

Protandrenopsis fuscipennis Crawford, 1903. Canad. Ent. 35: 337. °.

Taxonomy: Michener, 1951. Jn Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr.
Monog. 2: 1098 (synonymy). —Rozen, 1966. Amer. Mus. Novitates 2259: 9 figs. 9-10 (larva).
—Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 8-10 figs. 1-2 (redescription).

Biology: Hicks, 1931. Canad. Ent. 63: 172 (nest). —Rozen, 1967. Amer. Mus. Novitates 2297:
19-27, figs. 1-2, 7, tables 1-2 (nest architecture, life history, floral relationship).

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 10 (pollen sources).
bradleyi Timberlake. Tex. (Blanco County).
Pseudopanurgus bradleyi Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 10, figs. 3-4. ¢.
cazieri cazieri Timberlake. Ariz. (Cochise County). Another subspecies, P. c. coahwilensis
Timberlake, occurs in Mexico.
Pseudopanurgus cazieri cazieri Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 18, figs. 7-8.
2068

Superfamily APOIDEA 1853

dicksoni Timberlake. N. Mex., Ariz.; Mexico (Baja California and Sonora). Pollen: Unknown,
but visits flowers of Aplopappus, Argemone intermedia, Crotalaria pumila, Milleria
quinquefolia, Pectis papposa.
Pseudopanurgus dicksoni Timberlake, 1967. Amer. Mus. Novitates 2298: 8. 9, d.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 19, figs. 15-16 (tax. characters,
flower record).
fraterculus fraterculus (Cockerell). West. Tex., N. Mex.; Mexico (Aguascalientes, Coahuila,
Durango and San Luis Potosi). Pollen: Unknown, but visits flowers of Gutierrezia
lucida, Tidestromia.
Calliopsis fraterculus Cockerell, 1896. Canad. Ent. 28: 159. 3, °.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 21-23 (redescription, tax. status).

fraterculus timberlakei Cockerell. Colo., N. Mex., Ariz., Calif. (San Diego County); Mexico
(Baja California). Parasite: Holcopasites arizonicus (Linsley)?, H. insoletus (Linsley)?
Pollen: Apparently an oligolege of the Compositae, including Aplopappus gracilis,
Baileya multiradiata, B. pleniradiata, Gutierrezia californica, G. microcephala,
Heliopsis, Heterotheca subaxillaris, Hymenothrix wislizeni, Pectis papposa, but visits
these and other flowers for nectar including Allionia, Eriogonum.

Pseudopanurgus timberlakei Cockerell, 1931. Ent. Soc. Wash., Proc. 33: 201.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 23-24, figs. 2, 19-20 (redescription,
floral and geogr. records).

Biology: Rozen, 1965. N. Y. Ent. Soc., Jour. 73: 88 (parasite). —Rozen, 1967. Amer. Mus.
Novitates 2297: 19-27, figs. 1-2, tables 1-2 (nest architecture, life history, pollen source,
parasite). —Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 39 (parasites).

pectidellus Cockerell. N. Mex., Ariz.; Mexico (Chihuahua). Pollen: Apparently an oligolege of
the Compositae, visits flowers of Baileya, Pectis papposa, Verbesina.
Pseudopanurgus pectidellus Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 26.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 32-34, figs. 33-34 (redescription,
geogr. and floral records).
pectiphilus (Cockerell). N. Mex., Ariz. Pollen: Apparently an oligolege of the Compositae,
visits flowers of Baileya multiradiata, Heterotheca subaxillaris, Pectis papposa,

Tidestromia lanuginosa.
Panurginus pectiphilus Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 193. 2.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 34-35, figs. 35-36 (redescription,
geogr. and floral records).
perarmatus Timberlake. South. Ariz.; Mexico (Chihuahua and Sonora). Pollen: Apparently an
oligolege of the Compositae, visits flowers of Aplopappus gracilis, Heterotheca
subawxillaris, Verbesina encelioides, Zexmenia podocephala.
Pseudopanurgus perarmatus Timberlake, 1967. Amer. Mus. Novitates 2298: 5. 2, d.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 35, figs. 37-38 (tax. characters).

perpunctatus Timberlake. N. Mex., Ariz. Pollen: Presumably an oligolege of Compositae, visits
flowers of Heterotheca, Pectis papposa.
Pseudopanurgus perpunctatus Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 35, figs. 39-40.
OFIOr
rugosus (Robertson). Md. to Ga., west to IIl., Nebr., Kans., Tex. Pollen: Apparently an oligolege
of the Compositae, especially the Helianthae, visits flowers of Bidens aristosa,
Helianthus annuus, H. divaricatus, H. mollis, H. petiolaris, H. subaxillaris, Heliopsis
helianthoides, Pontederia cordata, Rudbeckia laciniata, R. triloba, Silphium
perfoliatum, S. speciosum, but visits flowers of Melilotus officinalis for nectar.
Calliopsis rugosus Robertson, 1895. Amer. Ent. Soc., Trans. 22: 121. 9, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 269-270, figs. 62, 65-66
(redescription). —Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 38, figs. 41-42 (tax.
characters, geogr. and floral records).

1854 Hymenoptera in America North of Mexico

texanus Timberlake. Tex.; Mexico (Tamaulipas).
Pseudopanurgus tecanus Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 40, figs. 45-46. °,
3

verticalis Timberlake. N. Mex., Ariz., south. Calif. (Riverside County); Mexico (Sonora).
Parasite: Holcopasites insoletus (Linsley). Pollen: Apparently an oligolege of the
Compositae; visits flowers of Bahia absinthifolia, Pectis papposa, Verbesina encelioides,
but also visits the flowers of Kallstroemia grandiflora presumably for nectar.
Pseudopanurgus verticalis Timberlake, 1967. Amer. Mus. Novitates 2298: 2. ?, d.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 9, figs. 11-16 (larva, as Pseudopanurgus
sp. B). —Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 50, figs. 55-56 (tax. characters).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 27 (parasite of Pseudopanurgus sp. B).
—Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 39 (parasite).

Genus ANTHEMURGUS Robertson

Anthemurgus Robertson, 1902. Canad. Ent. 34: 321.
Type-species: Anthemurgus passiflorae Robertson. Monotypic and orig. desig.

Taxonomy: Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 1 (tax. status).

passiflorae Robertson III. to N. C. Pollen: Unknown, but visits flowers of Passiflora.
Anthemurgus passiflorae Robertson, 1902. Canad. Ent. 34: 321. 2, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 265-266, figs. 62, 65-66
(redescription).

Genus HETEROSARUS Robertson

Heterosarus Robertson, 1918. Ent. News 29: 91.
Type-species: Calliopsis parvus Robertson. Monotypic and orig. desig.

Revision: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 1-56, 8 plates, 87 figs. (N. Amer. spp.).

Taxonomy: Timberlake, 1964. Amer. Mus. Novitates 2185: 2-5 (key to spp. in Amer. north of
Mexico). —Timberlake, 1973. Calif. Univ. Pubs. Ent. 72: 1, 2 (tax. status).
arizonicus (Timberlake). Ariz., N. Mex. Pollen: Unknown, but visits flowers of Eriogonum.
Pseudopanurgus arizonicus Timberlake, 1964. Amer. Mus. Novitates 2185: 23. d, 2.

Taxonomy: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 14, figs. 9-10 (tax. characters).
bakeri (Cockerell). Colo., N. Mex., Ariz.; Mexico (Chihuahua, Michoacan, Morelos and Puebla).
Pollen: Unknown, but visits flowers of Calochortus gunnisonii, Euphorbia, Frasera
speciosa, Helenium hoopesii, Heterotheca subaxillaris, Linum lewisti, Lopezia, Monarda
austromontana, Oxalis, Phacelia leucophylla, Potentilla filipes, P. fruticosa,
Pseudocymopterus montanus, Sedum stenopetalum, Solidago.
Calliopsis bakeri Cockerell, 1896. Ent. News 7: 221. 6.

Taxonomy: Viereck, 1903. Amer. Ent. Soc., Trans. 29: 50. —Cockerell, 1910. Psyche 17: 245.
3, 2. —Timberlake, 1964. Amer. Mus. Novitates 2185: 7-8 (geogr. and floral records).
—Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 15-16, figs. 13-14 (tax. characters, geogr.
and floral records).

dawsoni (Timberlake). Tex., N. Mex., Colo. Pollen: Unknown, but visits flowers of an
unspecified crucifer.
Pseudopanurgus dawsoni Timberlake, 1964. Amer. Mus. Novitates 2185: 10. 2, d.

Taxonomy: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 20, figs. 23-24 (tax. characters).
euphorbiae Timberlake. Ariz. (Pima County); Mexico (Chiapas, Michoacan, Morelos, Nayarit
and Puebla).
Heterosarus euphorbiae Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 24, figs. 29-30. d, @.
flavotinctus (Cockerell). N. Mex., Colo., Ariz.; Mexico (Zacatecas). Pollen: Unknown, but visits
flowers of Bigelovia, Sphaeralcea.
Panurginus pauper var. flavotinctus Cockerell, 1898. Denison. Univ. Sci. Labs., Bul. 11: 51.
é.

Superfamily APOIDEA 1855

Taxonomy: Cockerell, 1898. N. Mex. Univ., Bul. 1: 51. 6. —Cockerell, 1899. Entomologist 32:
129. 2. —Timberlake, 1964. Amer. Mus. Novitates 2185: 20-23 (redescription).
—Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 26, figs. 33-34 (tax. characters and status).

illinoiensis (Cresson). Que. to Fla., west to IIl., Kans., Okla. and Tex. Pollen: Unknown, but
visits flowers of Amphiachyris dracunculoides, Bellis integrifolia, Chrysanthemum
leucanthemum, Descourainia, Erigeron, Gaillardia, Heterotheca subaxillaris, Lepidium.
Calliopsis illinoiensis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 66. 3.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 274, figs. 64-66, table 8
(redescription). —Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 30-31, figs. 43-44 (tax.
status, geogr. and floral records).

nanulus (Timberlake). N. Mex., Ariz.; Mexico (Baja California, Chihuahua, Durango and
Sonora). Parasite: Holcopasites tegularis Hurd and Linsley. Pollen: Unknown, but visits
flowers of Euphorbia, Physalis, Wislizenia refracta.
Pseudopanurgus nanulus Timberlake, 1964. Amer. Mus. Novitates 2185: 5. 2, ¢.

Taxonomy: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 37, figs. 55-56 (tax. characters).

Biology: Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 26, 39 (parasite).
neomexicanus (Cockerell). N. Mex., Ariz.; Mexico (Distrito Federal, Durango, Guanajuato,
Hidalgo, Mexico, Michoacan, Morelos, Puebla and Tlaxcala). Pollen: Unknown, but visits
flowers of Cacalia decomposita, Ceanothus, Erigeron, Geranium, Helenium hoopesii,
Monarda austromontana, Penstemon, Solidago.
Panurginus neomexicanus Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 451. ¢.
Panurginus nigrinus Viereck, 1903. Amer. Ent. Soc., Trans. 29: 50. 2.

Taxonomy: Timberlake, 1964. Amer. Mus. Novitates 2185: 8-10 (redescription, geogr. and
floral records). —Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 37-38, figs. 57-58 (tax.
characters).

opacicollis (Timberlake). Ariz. Pollen: Unknown, but visits flowers of Eriogonum.
Pseudopanurgus opacicollis Timberlake, 1964. Amer. Mus. Novitates 2185: 15. 9.

Taxonomy: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 38 (tax. characters).

opaculus (Cockerell). Ariz. (Mud Springs, Santa Catalina Mts.).
Panurginus opaculus Cockerell, 1922. Amer. Mus. Novitates 36: 3. °.

Taxonomy: Timberlake, 1964. Amer. Mus. Novitates 2185: 18-20 (redescription).
—Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 39, fig. 87 (tax. characters).
parvus (Robertson). Pa., N. C., Ga., Miss., Mo., Ill, Wis., N. Dak., Alta., Colo., N. Mex. Pollen:
Unknown, but visits flowers of Aster, Eulophus, Geranium, Gerardia tenuifolia,
Gillenia, Monarda, Solidago, Thaspium.
Calliopsis parvus Robertson, 1892. Amer. Nat. 26: 273. 2, 6.
Pseudopanurgus gerardiae Crawford, 1932. Ent. Soc. Wash., Proc. 34: 77. d, °@.
Pseudopanurgus stevensi Crawford, 1932. Ent. Soc. Wash., Proc. 34: 78. d, °.
Panurginus borealis Cockerell, 1937. Canad. Ent. 69: 33. °.
Taxonomy: Crawford, 1932. Ent. Soc. Wash., Proc. 34: 77 (tax. characters). —Mitchell, 1960.
N. C. Agr. Expt. Sta. Tech. Bul. 141: 277-278, figs. 64-66, table 8 (synonymy, redescription).

—Timberlake, 1964. Amer. Mus. Novitates 2185: 25 (synonymy, as stevens?).
—Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 40, figs. 63-64 (synonymy, tax. characters).

pauper (Cresson). New England to Fla., west to Ill. and Mo. Pollen: Collects pollen especially
from the flowers of Ceanothus americanus, but also visits flowers of Rubus presumably

for nectar.
Calliopsis pauper Cresson, 1878. Amer. Ent. Soc., Trans. 7: 66. 2, d.
Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 278-279, figs. 64-66, table 8
(redescription). —Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 40, figs. 65-66 (tax.
characters).

Biology: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 40 (pollen source).

1856 Hymenoptera in America North of Mexico

pernitens (Cockerell). Ariz., N. Mex.
Panurginus pernitens Cockerell, 1922. Amer. Mus. Novitates 36: 4. °.

Taxonomy: Timberlake, 1964. Amer. Mus. Novitates 2185: 14-15 (redescription).
—Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 40-41 (tax. characters).
subglaber Timberlake. Tex. (Del Rio).
Heterosarus subglaber Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 45, figs. 71-72. 6.

townsendi (Cockerell). Tex., N. Mex., Ariz.
Calliopsis townsendi Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 152. 6.

Taxonomy: Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 48-49, figs. 77-78. ¢, 2
(redescription).
virginicus (Cockerell). Md. to Ga., W. Va. and Ala. Pollen: Unknown, but visits flowers of
Ceanothus, Houstonia purpurea.
Panurginus virginicus Cockerell, 1907. Entomologist 40: 137. d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 281-282, figs. 64-66, table 8
(redescription). —Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 52, figs. 83-84 (tax.
characters).

Genus PTEROSARUS Timberlake

Pterosarus Timberlake, 1967. Amer. Mus. Novitates 2298: 10.
Type-species: Calliopsis rudbeckiae Robertson. Orig. desig.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 5-8, figs. 1-8 (larvae, as
Pseudopanurgus). —Timberlake, 1967. Amer. Mus. Novitates 2298: 19-22 (Key to spp.).
—Timberlake, 1975. Calif. Univ. Pubs. Ent. 77: 1, 2 (key, tax. status).

aestivalis (Provancher). Que.
Panurgus aestivalis Provancher, 1882. Nat. Canad. 13: 205. 9, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 271-272 (tax. characters).
albitarsis (Cresson). Conn. to Ga., west to IIl., Colo., N. Mex. and Tex., ?Calif. Pollen:
Unknown, but visits flowers of Aster ericoides villosus, Brauneria pallida, B. purpurea,
Coreopsis palmata, Helianthus divaricatus, H. mollis, Lepachys pinnata, Rudbeckia
hirta, R. laciniata, R. submentosa, Verbesina helianthoides.
Panurgus albitarsis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 260. 2, 6.
Panurgus picipes Cresson, 1872. Amer. Ent. Soc., Trans. 4: 261. d.

Taxonomy: Cockerell, 1900. Canad. Ent. 32: 364. d. —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 266-267, figs. 62, 65-66, table 8 (synonymy, redescription). —Timberlake,
1967. Amer. Mus. Novitates 2298: 11 (tax. position).

altissimus (Cockerell). Colo.
Panurginus altissimus Cockerell, 1922. Amer. Mus. Novitates 36: 3. d, 2.
andrenoides (Smith). N. B. to N. C., west to Minn. and IIl., ?N. Mex. Pollen: Unknown, but
visits flowers of Aster, Rudbeckia triloba, Solidago. Predator: Philanthus lepidus
Cress.
Scrapter Andrenoides Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 121. °.
Calliopsis asteris Robertson, 1895. Amer. Ent. Soc., Trans. 22: 121. 2, d.

Taxonomy: Cockerell, 1904. Canad. Ent. 36: 303. 2. —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 272-273, figs. 64-66, table 8 (synonymy, redescription).

atricornis (Cresson). Colo., N. Mex.
Calliopsis atricornis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 67. 3.
Panurginus atricomis (!) Viereck, 1903. Amer. Ent. Soc., Trans. 29: 50 (tax. characters).

Taxonomy: Timberlake, 1967. Amer. Mus. Novitates 2298: 11 (tax. position).
aurifodinae (Michener). South. Calif., mountains.
Pseudopanurgus aurifodinae Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 320. 6.
barberi (Cockerell). N. Mex.
Panurginus barberi Cockerell, 1899. Entomologist 32: 129. ¢.

Superfamily APOIDEA 1857

boylei (Cockerell). Colo., N. Mex., Ariz. Parasite: Holcopasites insoletus (Linsley)? Pollen:
Collects pollen from the flowers of Verbesina encelioides .
Calliopsis boylei Cockerell, 1896. Canad. Ent. 28: 161. d.

Taxonomy: Cockerell, 1897. Acad. Nat. Sci., Phila., Proc. 49: 350. —Crawford, 1912. Canad.
Ent. 44: 368 (tax. characters). —Rozen, 1966. Amer. Mus. Novitates 2259: 6-7, figs. 1-7
(larva).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 19-27, figs. 1-2, tables 1-2 (nest
architecture, life history, parasite).
californicus (Cresson). Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Chaenactis glabriuscula, Encelia californica, Gilia multicaulis, Lasthenia californica,
L. gracilis, Layia elegans, L. platyglossa, Ranunculus californicus.
Calliopsis californicus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 67. ¢.
citrinifrons (Viereck). N. Mex.
Panurginus citrinifrons Viereck, 1903. Amer. Ent. Soc., Trans. 29: 51. ¢.
citripes (Ashmead). Colo.
Calliopsis citripes Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 5. d.
compositarum (Robertson). Md. to Ga., west to Ill. Pollen: Unknown, but visits flowers of
Aster, Bidens aristosa, Boltonia asteroides, Polygonum scandens, Rudbeckia triloba,
Solidago nemoralis, S. ulmifolia.
Calliopsis compositarum Robertson, 1893. Amer. Ent. Soc., Trans. 20: 274. °.

Taxonomy: Robertson, 1895. Amer. Ent. Soc., Trans. 22: 121. 6. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 273-274, figs. 64-66 (redescription).
didirupa (Cockerell). Idaho, Colo., N. Mex.
Panurginus didirupa Cockerell, 1908. Entomologist 41: 293. d, 9.
expallidus (Swenk and Cockerell). Nebr.
Panurginus expallidus Swenk and Cockerell, 1907. Ent. News 18: 181. d.
helianthi (Mitchell). Ind. (Tippecanoe Co.). Pollen: Unknown, but visits flowers of Helianthus
annuus.
Pseudopanurgus (Pseudopanurgus) helianthi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 267. 2.
horizontalis (Swenk and Cockerell). Nebr
Panurginus horizontalis Swenk and Cockerell, 1907. Ent. News 18: 188. d.
illustris (Timberlake). Ariz. (Chiricahua Mts.). Pollen: Unknown, but visits flowers of
Verbesina encelioides, Viguiera.
Pseudopanurgus illustris Timberlake, 1967. Amer. Mus. Novitates 2298: 18. 2.
innuptus (Cockerell). Alta., N. Dak., S. Dak., Nebr., Colo., N. Mex., Ariz. Pollen: Unknown, but
visits flowers of Helianthus.
Calliopsis innuptus Cockerell, 1896. Ent. News 7: 222. “2” = d.
Panurginus albitarsis var. fortior Cockerell, 1899. Entomologist 32: 129. 6.
Panurginus innuptus var. absonus Cockerell, 1912. Ent. News 23: 446. d.

Taxonomy: Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 194. 6, °.
irregularis (Cockerell). Colo.
‘Panurginus irregularis Cockerell, 1922. Amer. Mus. Novitates 36: 2. d.
labrosiformis distractus (Cockerell). Nebr.
Panurginus labrosiformis distractus Cockerell, 1912. Ent. News 23: 447. ¢.
labrosifromis labrosiformis (Robertson). N. C., Ga., Ill. Pollen: Unknown, but visits flowers of
Actinomeris alternifolia, Bidens aristosa, Coreopsis tripteris, Helianthus divaricatus,
H. tuberosus, Heliopsis helianthoides, Rudbeckia laciniata, R. triloba, Silphium
perfoliatum, Solidago canadensis.
Panurginus labrosiformis Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 49. °, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 274-275, figs. 64-66, table 8
(redescription).

1858 Hymenoptera in America North of Mexico

labrosus (Robertson). N. Y. to N. C., west to Minn. Pollen: Unknown, but visits flowers of
Helianthus divaricatus, H. tuberosus, Heliopsis, Rudbeckia lanceolata, R. triloba.
Calliopsis labrosus Robertson, 1895. Amer. Ent. Soe., Trans. 22: 122. 2, 3.

Taxonomy: Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 48, ?, ¢. —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 268-269, figs. 62, 65-66, table 8 (redescription).
leucopterus (Cockerell). Colo.
Panurginus leucopterus Cockerell, 1923. Ent. News 34: 49. 6.
lutzae (Cockerell). Colo.
Panurginus lutzae Cockerell, 1922. Amer. Mus. Novitates 36: 7. 2.
nebrascensis muesebecki (Michener). N. C. to Fla., Miss. Pollen: Unknown, but visits flowers
of Aster, Chrysopsis, Haplopappus.
Pseudopanurgus nebrascensis timberlakei Michener, 1947. Amer. Midland Nat. 38: 446. 6,
9. Preoce.
Pseudopanurgus (Heterosarus) nebrascensis muesebecki Michener, 1951. In Muesebeck,
Krombein and Townes, U.S. Dept. Agr., Agr. Monog. 2: 1100. N. name.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 276-277, fig. 65, table 8
(redescription, geogr. range).
nebrascensis nebrascensis (Crawford). Maine to N.J., west to Colo. and Alta. Pollen:
Unknown, but visits flowers of Aster, Solidago.
Panurginus nebrascensis Crawford, 1903. Canad. Ent. 35: 335. d, 2.

Taxonomy: Crawford, 1912. Canad. Ent. 44: 368 (tax. characters). —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 275-276, figs. 64-66, table 8 (redescription).

nubis (Cockerell). N. Mex.
Panurginus nubis Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 193. 9.

occidus (Timberlake). Ariz. Parasite: Holcopasites arizonicus (Linsley)? Pollen: Aplopappus
gracilis, Helianthus, Heterotheca subaxillaris, Verbesina encelioides.
Pseudopanurgus (Pterosarus) occidus Timberlake, 1967. Amer. Mus. Novitates 2298: 14.
hn
Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 7-8, fig. 8 (larva, as species A).
Biology: Rozen, 1965. N. Y. Ent. Soc., Jour. 73: 88 (parasite). —Hurd and Linsley, 1972.
Smithsn. Contrib. Zool. 114: 39 (parasite).
ornatipes (Cresson). Nebr., Colo., Tex.
Calliopsis ornatipes Cresson, 1872. Amer. Ent. Soc., Trans. 4: 260. ¢.
Taxonomy: Crawford, 1912. Canad. Ent. 44: 368. (tax. characters).

pecki (Cockerell). Alta.
Panurginus pecki Cockerell, 1937. Canad. Ent. 69: 114. 6.

perlaevis (Cockerell). Colo., N. Mex., Ariz. Parasite: Holcopasites insoletus (Linsley)?
Calliopsis perlaevis Cockerell, 1896. Canad. Ent. 28: 160. °.
Taxonomy: Cockerell, 1922. Amer. Mus. Novitates 36: 5.

Biology: Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 39 (parasite).
piercei albertensis (Cockerell). Alta.
Panurginus piercei albertensis Cockerell, 1937. Canad. Ent. 69: 33. °.
piercei piercei (Crawford). N. Dak., Nebr., Colo., N. Mex.
Panurginus piercei Crawford, 1903. Canad. Ent. 35: 335. 6, 9.

Biology: Pierce, 1904. Nebr. Univ., Studies 4: 185. —Hicks, 1936. Canad. Ent. 68: 47 (nesting

habits).
porterae (Cockerell). N. Mex., Colo., Ariz.

Panurginus Porterae Cockerell, 1900. Canad. Ent. 32: 364. 6, 2.
pulchricornis (Cockerell). Colo.

Panurginus pulchricornis Cockerell, 1922. Amer. Mus. Novitates 36: 8. °.
renimaculatus (Cockerell). N. Dak., Wyo., Nebr., Colo., N. Mex., Tex.

Calliopsis renimaculatus Cockerell, 1896. Ent. News 7: 222. °.

Superfamily APOIDEA 1859

Taxonomy: Swenk and Cockerell, 1907. Ent. News 18: 180. 2 (only). —Crawford, 1915. U.S.
Natl. Mus., Proc. 94: 579. 2, ¢6. —Stevens, 1919. Canad. Ent. 51: 208. ¢.
rudbeckiae (Robertson). Wis., Ill. Pollen: Unknown, but visits flowers of Aster, Bidens
aristosa, Helianthus divaricatus, Rudbeckia hirta, R. laciniata, R. submentosa, R.

triloba.
Calliopsis rudbeckiae Robertson, 1895. Amer. Ent. Soc., Trans. 22: 122. 2, 6.

Biology: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 279-280, figs. 64-66, table 8
(redescription).
simulans (Swenk and Cockerell). N. Dak., Nebr.
Panurginus simulans Swenk and Cockerell, 1907. Ent. News 18: 182. 6, 9.
solidaginis (Robertson). New England west to IIl., south to Miss. Pollen: Unknown, but visits
flowers of Bidens aristosa, Boltonia asteroides, Coreopsis tripteris, Helianthus

grosse-serratus, H. tuberosus, Rudbeckia submentosa, Solidago canadensis.
Calliopsis solidaginis Robertson, 1893. Amer. Ent. Soc., Trans. 20: 274. 9.

Taxonomy: Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 50. ¢. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 280-281, figs. 64-66, table 8 (redescription).
stathamae (Timberlake). Ariz. (Southwestern Research Station).
Pseudopanurgus (Pterosarus) stathamae Timberlake, 1967. Amer. Mus. Novitates 2298:
165.92
stigmalis (Swenk and Cockerell). Nebr.
Panurginus stigmalis Swenk and Cockerell, 1907. Ent. News 18: 180. d, 2.
vicinus (Timberlake). Ariz. (Cochise County). Pollen: Unknown, but visits flowers of
Aplopappus gracilis.
Pseudopanurgus (Pterosarus) vicinus Timberlake, 1967. Amer. Mus. Novitates 2298: 11.
ORIG
ximenesiae (Cockerell). N. Mex.
Panurginus ximenesiae Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 195. 2.

Genus METAPSAENYTHIA Timberlake

Metapsaenythia Timberlake, 1969. Ent. News 80: 89.
Type-species: Calliopsis abdominalis Cresson. Orig. desig.

Taxonomy: Timberlake, 1969. Ent. News 80: 89-92 (N. Amer. spp.). —Timberlake, 1973. Calif.
Univ. Pubs. Ent. 72: 1 (tax. relationships).
abdominalis abdominalis (Cresson). Tex., Kans. Pollen: Collects pollen from flowers of
Monarda, but visits other flowers for nectar including Aster tanacetifolius, Brazoria
truncata, Dalea aurea, Gaillardia, Helianthus annuus.
Calliopsis abdominalis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 68. °, d.
Camptopoeum semirufum Cockerell, 1937. Amer. Mus. Novitates 899: 1. ?.

Biology: Timberlake, 1969. Ent. News 80: 90 (floral relationships).
abdominalis tricolor (Cockerell). N. J. and Pa. to Ga. Pollen: Apparently an oligolege of
Monarda including M. pwnctata.
Calliopsis tricolor Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 151. °.
Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 270-271, figs. 62-63, 66, table
8 (redescription). —Timberlake, 1969. Ent. News 80: 90-91 (tax. status).

Genus NOMADOPSIS Ashmead

Revision: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 1-202, 218 figs., 17 maps (monograph,
includes information on biology, systematics and ecology).

Taxonomy: Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 195-198 (tax. status and characters).
—Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 104-107 (tax. status). —Rozen, 1959. Ent.
Soe. Wash., Proc. 61: 255-259 (geogr. and floral records). —Rozen, 1966. Amer. Mus.
Novitates 2259: 5, 16 (larvae). —Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 6-8,
figs. 3-10 (pupae).

1860 Hymenoptera in America North of Mexico

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 4-54 (life history, nest architecture, adult
behavior, comparative ecology, floral relationships, parasites, predators). —Rozen, 1963.
Amer. Mus. Novitates 2142: 1-17, 20 figs., 2 tables (nest architecture, life history, floral
relationships). —Rozen, 1967. Amer. Mus. Novitates 2297: 1-44, 18 figs. 2 tables (review).

Genus NOMADOPSIS Subgenus NOMADOPSIS Ashmead

Nomadopsis Ashmead, 1898. Psyche 8: 285.
Type-species: Peridita zonalis Cresson. Monotypic and orig. desig. Lapsus for
Calliopsis zonalis Cresson.
Spinoliella subg. Claremontiella Cockerell, 1933. Pan-Pacifie Ent. 9: 25.
Type-species: Calliopsis zonalis Cresson. Monotypic and orig. desig. (=Spinoliella
euxantha Cockerell).

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 60-63 (tax. status).
cincta cincta (Cresson). Calif. (east side of Sierra Nevada, Tulare and Trinity Counties),
Nevada. Pollen: Apparently an oligolege of Calochortus including C. luteus.
Calliopsis cinctus Cresson, 1879. Amer. Ent. Soc., Trans. 7: 201. 2, 6.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 83, figs. 94-98, map 4 (redescription, tax.

status).
cincta hurdi Rozen. Calif. (Sacramento). Pollen: Collects pollen from the flowers of
Calochortus luteus.
Nomadopsis cincta hurdi Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 85. 6, 2.
comptula (Cockerell). Calif. (Tulare, Inyo, and San Bernardino Counties). Pollen: Collects
pollen from the flowers Potentilla, including P. bolanderi var. bernardina, P. b. var.
parryi, P. gracilis, but visits these and other flowers including Calochortus palmeri var.
paludicola for nectar.
Spinoliella comptula Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 284. @.
Spinoliella edwardsii media Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 325. 2, ¢.
Spinoliella edwardsii bernardina Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 326. °,

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 75-77, figs. 79-83, map 3 (redescription,
synonymy).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 7, 31, 75 (nest site, mating, floral
relationships).
edwardsii (Cresson). Calif., Oreg., Wash., Idaho. Pollen: Collects pollen from flowers of
Calochortus, Chamaebatia foliolosa, Potentilla including P. bolanderi var. parryi, P.
congesta, P. fusca, P. glandulosa, P. tridentata, but visits these and other flowers for
nectar including Clarkia rhomboidea, Eriogonum marifolium, Fragaria californica, Iris
hartwegui, Lathyrus sulphureus, Spraguea umbellata.
Calliopsis Edwardsii Cresson, 1878. Amer. Ent. Soce., Trans. 7: 64. 2, 6.
Calliopsis lateralis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 65. @.
Spinoliella triangulifera Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 195. 9.

Taxonomy: Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 325, 3 (as triangulifera).
—Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 77-81, figs. 4-5, 28-29, 64, 84-88, map 3
(redescription, synonymy, larva).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 6, 28-31, 81 (nest site, mating behavior,
floral relationships).

linsleyi Rozen. Calif.; Mexico (Baja California). Pollen: Collects pollen from the flowers of
Eriodictyon, including E. angustifolium, E. crassifolium, E. parryi, E. trichocalyzx, but
visits these and other flowers for nectar including Arctostaphylos, Ceanothus.
Nomadopsis linsleyi Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 85, figs. 6, 7, 30, 31, 53,
99-103, map 5. 6, 2.
obscurella (Cresson). Calif., Oreg., Ariz. (Tucson), ?Wash.; Mexico (Baja California). Pollen:
Apparently oligolectic on E'schscholzia, especially E. californica, but visits other flowers

Superfamily APOIDEA 1861

including Agoseris glauca, Calendula, Ceanothus, Chaenactis glabriuscula, Coreopsis
lanceolata, Cryptantha intermedia, Lasthenia, Layia platyglossa, L. p. breviseta,
Malacothrix californica, Nemophila menziesii, Phacelia davidsonii, P. distans, P.
tanacetifolia, Sida malvaeflora, Sphaeralcea.

Calliopis obscurellus Cresson, 1879. Amer. Ent. Soc., Trans. 7: 201. 2, 6.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 91-93, figs. 104-108, map 7 (redescription).

Biology: Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 107 (mating). —Rozen, 1958. Calif.
Univ. Pubs. Ent. 15: 32, 93 (mating, floral relationships).
puellae (Cockerell). Tex., N. Mex., Ariz., Nev., Calif., Idaho.; Mexico (Baja California). Parasite:
Oreopasites vanduzeei Cockerell. Pollen: Evidently collects pollen chiefly from the
flowers of Malacothrix including M. californica, M. exigua, M. glabrata, but visits other
flowers including Anisocoma acaulis, Aster abatus, Astragalus, Baileya pleniradiata,
Calycoseris wrightii, Chaenactis fremontii, Encelia farinosa, Haplopappus cooper, H.
linearifolius, Rafinesquia neomexicana.
Spinoliella puellae Cockerell, 1933. Pan-Pacifie Ent. 9: 25. 9°, d.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 89-91, figs. 89-93, map 6 (redescription).
—Rozen, 1959. Ent. Soc. Wash., Proc. 61: 255 (geogr. records). —Yager and Rozen, 1966.
Amer. Mus. Novitates 2265: 8 (pupa).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 8, 31, 54, 89 (nest site, mating behavior,
parasites, floral relationships).
timberlakei Rozen. Calif. (Los Angeles County), desert. Pollen: Possibly an oligolege of
Monardella exilis.
Nomadopsis timberlakei Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 78, map 2. 6, °.
zonalis sierrae Rozen. Calif. (Sierra Nevada Mts.). Parasite: Oreopasites sp. Pollen: Collects
pollen from the flowers of Monardella lanceolata, but visits these and other flowers for
nectar and or pollen including Solidago californica.
Nomadopsis zonalis sierrae Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 71-73, map 2. d, °.

Taxonomy: Rozen, 1963. Amer. Mus. Novitates 2142: 3-5, figs. 1-2, table 1 (immature stages).
—Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 7-8, figs. 7-8 (pupa).

Biology: Rozen, 1963. Amer. Mus. Novitates 2142: 2-6 (nest site, nest architecture, floral
relationships, parasite).
zonalis zonalis (Cresson). Calif. (central and south coasts). Pollen: Collects pollen from the
flowers of Monardella douglasii, M. lanceolata.
Calliopsis zonalis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 201. d.
Spinoliella euxantha Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 283. ?.

Taxonomy: Timberlake, 1951. In Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr.
Monog. 2: 1102 (synonymy). —Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 69-71, figs. 66, 74-78,
map 2 (redescription).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 70-71 (floral relationships).

Genus NOMADOPSIS Subgenus MACRONOMADOPSIS Rozen

Nomadopsis subg. Macronomadopsis Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 93.
Type-species: Nomadopsis micheneri Rozen. Orig. desig.
anthidia anthidia (Fowler). Oreg., Calif. (west slope of Sierra Nevada, northern Coast Ranges,
Sacramento Valley, part of San Joaquin Valley). Parasite: Euphytomima nomiivora
James, Oreopasites vanduzeii melanantha Linsley. Pollen: Collects pollen from the
flowers of Trifolium including T. involucratum, T. melananthum, T. m icrocephalum, T.
monanthum var. parvum, T. repens, T. tridentatum, T. variegatum, but visits these and
other flowers for nectar including Eriodictyon trichocalyx, Lupinus bicolor. Predator:
Solenopsis sp.
Calliopsis anthidius Fowler, 1899. Psyche 8: 407. d (? misdet.).

1862 Hymenoptera in America North of Mexico

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 99-102, figs. 8, 9, 56, 63, 119-123, map 9
(tax. status, redescription, larva). —Rozen, 1959. Ent. Soe. Wash., Proc. 61: 258 (gender of
name). —Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 8, figs. 9-10 (pupa).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 8-9, 32-34, 53, 101 (nest site, nest
architecture, mating behavior, floral relationships, parasite, predator, larva).
—Moradeshaghi and Bohart, 1968. Kans. Ent. Soc., Jour. 41: 456-473, 15 figs., 3 tables
(parasite).

anthidia lutea Rozen. Calif. (east of Sierra and southern Calif.), Oreg., Idaho, Utah. Pollen:
Collects pollen from the flowers of Trifolium including T. involucratum, T. repens.
Nomadopsis anthidius lutea Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 102, figs. 2, 3, 10, 11,
34, 35, 57, 124-128, map 9. 6, &.
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 9, 34, 101 (nest site, mating behavior, floral
relationships).
barri Rozen. Idaho, Oreg. Pollen: Unknown, but visits flowers of Melilotus.
Nomadopsis (Macronomadopsis) barri Rozen, 1959. Ent. Soc. Wash., Proc. 61: 256, figs. 1,
AM Gra or
filiorum Rozen. Calif. (Mono Co.). Pollen: Probably collects pollen from the flowers of
Trifolium.
Nomadopsis (Macronomadopsis) filiorum Rozen, 1963. Amer. Mus. Novitates 2142: 6, figs.
3-6. 3, @.
Biology: Rozen, 1963. Amer. Mus. Novitates 2142: 3-6, tables 1-2 (nest site, nest architecture,
floral relationships).
micheneri Rozen. Calif. (Sierra Nevada and Trinity Mt.). Pollen: Collects pollen from the
flowers of Trifolium including T. gracilentum, T. monanthus var. parvum, T. repens.
Nomadopsis micheneri Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 95, figs. 12, 13, 32, 338, 55,
67, 109-113, map 8. 6, ¢&.
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 9-12, 34-36, 43, 53, 98 (nest site, nest
architecture, mating behavior, floral relationships).
zebrata bobbae Rozen. Ariz. (Navajo and Coconino Counties).
Nomadopsis zebrata bobbae Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 107, map 8. 6, 2.
zebrata zebrata (Cresson). S. Dak., Wyo., Colo., N. Mex. Pollen: Collects pollen from the
flowers of Astragalus.
Calliopsis zebratus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 64. 2.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 103-107, figs. 114-118, map 8 (tax. status,

redescription).

Biology: Cockerell, 1915. Ent. News 26: 366 (nest site).—Hicks, 1936. Canad. Ent. 68: 47
(nest).— Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 35 (nest site).

Genus NOMADOPSIS Subgenus MICRONOMADOPSIS Rozen

Nomadopsis subg. Micronomadopsis Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 107.
Type-species: Nomadopsis fracta Rozen. Orig. desig.
australior (Cockerell). S. Dak., Nebr., Colo., Utah, N. Mex., Tex., Ariz.; Mexico (Chihuahua).
Pollen: Collects pollen from the flowers of Lepidium alyssoides, L. montanum, but may
also obtain pollen from the flowers of Dithyrea wislizeni.
Calliopsis australior Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 349. 2 (¢ misdet.).

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 149-152, figs. 71, 182-185, map 15. 6, 2
(redescription). —Rozen, 1959. Ent. Soc. Wash., Proc. 61: 259 (geogr. rec.).

Biology: Custer, 1927. Psyche 34: 199 (nesting activities). —Custer, 1929. Canad. Ent. 61: 49
(nest). —Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 8, 152 (nest site, floral relationships).

Superfamily APOIDEA 1868

barbata Timberlake. South. Calif., north into Central Valley. Parasite: Oreopasites sp. Pollen:
Collects pollen from the flowers of Phacelia including P. distans, P. tanacetifolia, but
visits these and other flowers for nectar including Cryptantha, Linanthus dianthiflorus.

Nomadopsis barbata Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 107. d, °.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 114-117, figs. 18-19, 42-43, 59, 133-136,
map 11 (redescription, larva).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 8, 36, 40, 117, table 2 (nest site, nesting
activity, mating behavior, floral relationships, parasite).
beamerorum Rozen. Tex. (Reeves and Pecos Counties). Pollen: Unknown, but holotype male
was visiting flowers of Prosopis.
Nomadopsis (Micronomadopsis) beamerorum Rozen, 1963. Amer. Mus. Novitates 2142: 13,
figs. 12-15. 6, 2.
callosa Timberlake. Tex. (E] Paso Co.); Mexico (Baja California).
Nomadopsis callosa Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 110. d.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 140-142, figs. 174-177, map 8
(redescription). —Rozen, 1959. Ent. Soc. Wash., Proc. 61: 259 (type).
cazieri Rozen. Calif. (San Jacinto Mts. and San Diego County).
Nomadopsis cazieri Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 135, figs. 165-169, map 14. 6,
oF
foleyi Timberlake. South. Calif., deserts. Pollen: Collects pollen from the flowers of Larrea
tridentata, but visits these and other flowers for nectar including Heliotropium, Nama
hispida.
Nomadopsis foleyi Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 112. 9, d.
Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 119-121, figs. 141-144, map 10
(redescription).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 121 (floral relationships).
fracta Rozen. Calif. (widely distributed at lower altitudes). Pollen: Collects pollen from the
flowers of Eriodictyon angustifolium, E. californicum, E. crassifolium, E. trichocalyz,
but visits these and presumably other flowers for nectar including Ceanothus
integerrimus, Chaenactis, Chamaebatia foliolosa, Cryptantha intermedia, Erigeron
divergens, Eriogonum fasciculatum, Gilia exilis, Mimulus fremontii, Nama parry),
Phacelia brachyloba, P. douglasii.
Nomadopsis fracta Rozen, 1952. Kans. Ent. Soc., Jour: 25: 144, 4 figs. d, 2.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1036, figs. 92-98 (larva). —Rozen, 1958.
Calif. Univ. Pubs. Ent. 15: 111-113, figs. 36-37, 68-69, 129-132, map 10 (redescription, larva).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 36, 39, 48-50, 55, 60, 113 (mating behavior,

sleep, flower relationships, voltinism).

helianthi (Swenk and Cockerell). Nebr. and N. Mex., west to Calif.; Mexico (Baja California).

Parasite: Oreopasites euphorbiae Ckll. Pollen: Collects pollen from the flowers of
Euphorbia albomarginata, E. polycarpa, but visits these and other flowers for nectar
including Cryptantha intermedia, Eremocarpus setigerus, Eriogonum fasciculatum, E.
gracile, Gutierrezia sarothrae, Helianthus, Heliotropium curassavicum, Hemizonia
paniculata.

Spinoliella helianthi Swenk and Cockerell, 1907. Ent. News 18: 178. °.

Spinoliella euphorbiae Cockerell, 1925. Pan-Pacifie Ent. 1: 179. °.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1036, figs. 99, 106-108 (larva, as
euphorbiae). —Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 133, map 13 (type). —Rozen, 1958.
Calif. Univ. Pubs. Ent. 15: 129-133, figs. 16-17, 40-41, 61, 65, 157-160, map 13 (as
euphorbiae, redescription, larva). —Rozen, 1963. N. Y. Ent. Soe., Jour. 7: 142-143

(synonymy).

1864 Hymenoptera in America North of Mexico

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 11, 16-17, 37, 40, 48, 48-49, 50, 55, 59, 133
(adult activity, defecation, floral relationships, habitat, hibernation, mating behavior, sleep,

parasitism, voltinism).
hesperia equina (Cockerell). Calif. (Great Valley), Nev. (Virginia City). Parasite: Oreopasites
vanduzeei Ckll. Pollen: Collects pollen from the flowers of Heliotropium curassavicum,
but visits these and other flowers for nectar including Erodium cicutarium, Hemizonia

pungens, Medicago sativa.
Spinoliella equina Cockerell, 1925. Calif. Acad. Sci., Proe. (4) 14: 196. 2, 6.
Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 148-149 (redescription). —Yager and
Rozen, 1966. Amer. Mus. Novitates 2265: 6-7, figs. 5-6 (pupa).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 49, 59, 147 (flower relationships, parasite).
hesperia hesperia (Swenk and Cockerell). South. Calif. Pollen: Collects pollen from the flowers
of Heliotropium curassavicum oculatum.
Spinoliella hesperia Swenk and Cockerell, 1907. Ent. News 18: 186. °, 6.
Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 147-148, figs. 20-21, 44-46, 70, 178-181,
map 15 (redescription).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 49, 59, 147 (floral relationships).
larreae Timberlake. South. Calif. (Colorado and Mojave Deserts). Ecology: Nest sites are
established in sand dune areas. Pollen: Collects pollen from the flowers of Larrea

tridentata.
Nomadopsis larreae Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 111. @.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 126-128, figs. 153-156, map 12

(redescription).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 11, 37, 41, 48, 49, 55, 128 (mest site, nest
architecture, nesting activities, mating behavior, floral relationships).
macswaini Rozen. Ariz. Pollen: Evidently collects pollen from the flowers of Wislizenia

refracta.
Nomadopsis macswaini Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 152, figs. 195-198, map 16.

Gpmce
meliloti (Cockerell). Tex., N. Mex., Ariz. (Willcox). Pollen: Pollen-laden females have been
collected at the flowers of Dalea scoparia, but also visits flowers of Melilotus.
Calliopsis meliloti Cockerell, 1896. Canad. Ent. 28: 158. 9.

Taxonomy: Cockerell, 1933. Pan-Pacific Ent. 9: 158. 6, 2. —Rozen, 1958. Calif. Univ. Pubs.
Ent. 15: 138-140, figs. 170-173, map 14 (redescription). —Rozen, 1959. Ent. Soc. Wash.,
Proc. 61: 259 (geogr. record).

mellipes Timberlake. South. Calif. Pollen: Apparently collects pollen from the flowers of Lotus
including L. glaber, L. hamatus, L. salsuginosus, L. scoparius, L. strigosus, but visits
these and apparently other flowers for nectar including Camissonia dentata, Encelia.
Nomadopsis mellipes Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 109. 6, @.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 125-126, figs. 149-152, map 12
(redescription).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 126 (floral relationships).

nigromaculata Timberlake. South. Calif., Ariz., deserts. Pollen: Apparently collects pollen from
the flowers of Euphorbia polycarpa including E. p. var. hirtella, but visits other flowers
including Eriogonum trichopes, Pectis papposa possibly for nectar.
Nomadopsis nigromaculata Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 115. 2, 6.
Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 133-135, figs. 161-164, map 14
(redescription).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 37-38, 50, 52, 1385 (mating behavior, adult
behavior, floral relationships, voltinism).

Superfamily APOIDEA 1865

personata (Cockerell). Wash., Oreg., Idaho, Wyo. Pollen: Pollen laden females have been
collected at the flowers of Cleome lutea, C. serrulata.
Calliopsis personatus Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 349. 9.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 165-167, figs. 186-189, map 15
(redescription). —Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 7 (pupa).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 167 (floral relationships).
phaceliae Timberlake. Calif. (chiefly east side of Sierra Nevada and Mojave Desert). Pollen:
Apparently collects pollen from the flowers of Phacelia fremontii, but visits these and
other flowers for nectar including Euphorbia albomarginata, Hesperochiron
californicum, Nama demissum.
Nomadopsis phaceliae Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 116. 3.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 117-119, figs. 137-140, map 11
(redescription). —Rozen, 1959. Ent. Soc. Wash., Proc. 61: 258-259 (geogr. and floral
records).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 119 (floral relationships).
scitula lawae (Michener). Calif. (Inyo, Lassen and Mono Counties), Nev. (Fallon). Pollen: A
female laden with pollen has been taken at the flowers of Thelypodium brachycarpum,
others at flowers of Cleome and a male at the flowers of Cleomella oocarpa.
Spinoliella lawae Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 324. 9°.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 156-157, 159-160, map 16 (redescription).

Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 160 (floral relationships).
scitula scitula (Cresson). Wyo., Idaho, Utah, Colo., N. Mex.; variants occur in BrC., Oreg.,
northeast. Calif., northeast. Nev., Colo. and Wyo. Parasite: Oreopasites scituli Ckll.
Pollen: Females with pollen-laden hind legs have been collected at the flowers of
Cleome serrulata, Sisymbrium altissimum.
Calliopsis scitulus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 64. 2.
Calliopsis pictipes Cresson, 1878. Amer. Ent. Soc., Trans. 7: 65. 6.

Taxonomy: Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 348 (tax. characters).
—Timberlake, 1951. In Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog.
2: 1102 (synonymy). —Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 21, 154-159, figs. 72, 199-203,
map 16 (redescription, tax status, larva). —Yager and Rozen, 1966. Amer. Mus. Novitates
2265: 7 (pupa).

Biology: Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 440 (parasite). —Rozen, 1958. Calif.
Univ. Pubs. Ent. 15: 12, 40, 47, 159 (nest site, nesting activities, floral relationships.)

scutellaris scutellaris (Fowler). Wyo., Idaho, Utah, Nev., Wash., Oreg., Calif., Ariz.; Mexico

(northern Baja California). Parasite: Euphytomima nomiivora James, Myrmosula
rutilans (Fowler), Oreopasites sp. Pollen: Apparently polylectic, pollen-laden females
have been collected from the flowers of Heliotropium curassavicum,
Mesembryanthemum, Sesuvium sessile, Wislizenia refracta, alsike clover, ladino clover,
carrots, but also visits other flowers including Alyssum maritimum, Brassica incana,
Chrysanthemum, Cressa truxillense, Gutierrezia sarothrae, Melilotus indica, Phacelia
fremontii, Raphanus sativus, Sesuvium verrucosum, Solidago, Tamarix. Another
subspecies, N. scutellaris peninsularis (Cockerell), occurs in southern Baja California.

Calliopsis scutellaris Fowler, 1899. Psyche 8: 406. d.

Calliopsis visaliensis Fowler, 1899. Psyche 8: 406. °.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 20, 23, 24, 25, 27, 30, 160-165, figs. 24-25,
47-48, 62, 73, 190-194, map 17 (redescription, larva). —Yager and Rozen, 1966. Amer. Mus.
Novitates 2265: 7 (pupa).

Biology: Snelling, 1954. Pan-Pacific Ent. 30: 124 (parasite). —Rozen, 1958. Calif. Univ. Pubs.
Ent. 15: 12, 15, 38, 40, 48, 53-54, 164 (nest, site, nest architecture, life history, mating
behavior, parasite). —Moradeshaghi and Bohart, 1968. Kans. Ent. Soc., Jour. 41: 456-473,
15 figs., 3 tables (parasite).

1866 Hymenoptera in America North of Mexico

snellingi Rozen. Calif. (Squaw Valley). Pollen: Unknown, but visits flowers of Salvia.
Nomadopsis (Micronomadopsis) snellingi Rozen, 1963. Amer. Mus. Novitates 2142: 9, figs.
(Al iy Oe
trifolii Timberlake. Calif. Pollen: Collects pollen from the flowers of Mimulus fremontii,
Trifolium melananthum, T. tridentatum, T. variegatum, but also visits other flowers
including Cryptantha, Eriodictyon, Salix.
Nomadopsis trifolii Timberlake, 1952. Ent. Soc. Amer., Ann. 45: 113. 2, 6.
Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 14, 19, 23, 26, 30, 121-128, figs. 14-15,
38-39, 60, 145-148, map 12 (redescription, egg, larva).
Biology: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 10, 15, 17, 35, 40, 48-50, 123 (nest site, nest
architecture, life history, mating behavior, hibernation, sleep, voltinism, floral
relationships).

Genus NOMADOPSIS Subgenus UNASSIGNED

boharti Rozen. Calif. (Eldorado, Napa and Tuolumne Counties).
Nomadopsis boharti Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 171, figs. 204-208, map 7. 6,
OF

Taxonomy: Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 8 (pupa).
interrupta (Provancher). Ont. (Toronto).
Calliopsis interrupta Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 320. 2.
Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 174. (tax. status).

quadrilineata (Provancher). Ont. (Toronto).
Calliopsis 4-lineata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 319. 2.
Calliopsis quadrilineata Dalla Torre, 1896. Cat. Hym., v. 10, p. 174. Emend.

Taxonomy: Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 174 (tax. status).
smithi Rozen. Calif. (San Benito County). Pollen: Unknown, but visits flowers of Cryptantha.
Nomadopsis smithi Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 169, figs. 214-218, map 7. 6, °.
solitaria Rozen. Calif. (Lassen County).
Nomadopsis solitaria Rozen, 1963. Amer. Mus. Novitates 2142: 15, figs. 16-20. 6.

xenus Rozen. Wash., Calif. (Lake Tahoe). Pollen: Pollen-laden females have been collected
from the flowers of Phacelia.
Nomadopsis xenus Rozen, 1958. Calif. Univ. Pubs. Ent. 15: 167, figs. 209-213. dg, °@.

Genus CALLIOPSIS Smith

Revision: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 753-936, 156 figs., 8 maps, 9 tables (N. and
Cent. Amer. spp., includes biological information).

Taxonomy: Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 195-198 (U. S. spp.). —Mitchell, 1960.
N. C. Agr. Expt. Sta. Tech. Bul. 141: 287-293, figs. 58, 69-71, table 8 (eastern U. S. spp.).
—Rozen, 1966. Amer. Mus. Novitates 2259: 5, 13-16, figs. 29-40 (larvae). —Yager and
Rozen, 1966. Amer. Mus. Novitates 2265: 8-10, figs. 11-12 (pupae).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 1-44, 18 figs., 2 tables (review).

Genus CALLIOPSIS Subgenus CALLIOPSIS Smith

Calliopsis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 128.
Type-species: Calliopsis andreniformis Smith. Desig. by Ashmead, 1899.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 782-816, figs. 5, 6, 8-45, maps 1-3
(redescription, included spp.).
andreniformis Smith. Southeast. Canada including the Maritime Provinces, south to Fla., west
to Mont., S. Dak., Colo., Utah, Okla. and east Tex. Parasite: Holcopasites calliopsidis
(Linsley), H. illinoiensis (Robertson), Sphecodes brachycephalus Mitchell?, Sphecodes
sp., Parabombylius coquilletti (Will.)?, Trochometridium tribulatum Cross, Villa

Superfamily APOIDEA 1867

sinuosa (Wied.). Pollen: Polylectic, known sources include Aster ericoides villosus,
Convolvulus, Desmodium marilandicum, D. paniculatum, Erigeron canadensis,
Gerardia tenuifolia, Hedyotis purpurea, Lippia lanceolata, Lycopus sinuatus, Lythrum
alatum, Malva neglecta, M. rotundifolia, Melilotus alba, M. officinalis, Oxalis stricta,
Polygala sanguinea, Polygonum buxiforme, Psoralea onybrichis, Pycnanthemum
pilosum, Trifolium pratense, T. procumbens, T. repens, Verbena bracteata, V. hastata,
V. urticifolia, Verbesina helianthoides, but visits these and other flowers for nectar
including Achillea, Ailanthus altissima, Ammania coccinea, Amorpha canescens,
Anaphalis margaritacea, Asclepias, Bidens aristosa, Boltonia asteroides, Brassica,
Castanea pumila, Ceanothus americanus, Chrysanthemum leucanthemum, Chrysopsis,
Cleome, Coreopsis palmata, C. tripteris, Cryptotaenia canadensis, Cucumis, Dianthera
americana, Epilobium, Eryngium yuccifolium, Geum album, Gillenia stipulacea,
Hedeoma pulegioides, Hedyotis nigricans, Helenium, Hypericum perforatum, Lindernia
dubia riparia, Lespedeza capitata, L. procumbens, L. repens, L. reticulata, Ligustrum,
Medicago sativa, Nepeta cataria, Oenothera laciniata, Oxalis dillenii, Penstemon,
Petalostemon candidum, Polygonum convolvulus, P. pennsylvanicum, Portulaca,
Potentilla monspeliensis, P. recta, Prunella vulgaris, Prunus, Psoralea tenuiflora, P.
tenuiflora floribunda, Pycnanthemum flexuosum, P. virginianum, P. lanceolatum,
Raphanus sativus, Rhus glabra, Rosa, Rubus, Rudbeckia triloba, Serinea oppositifolia,
Sisymbrium repandum, Solanum carolinense, Solidago canadensis, Spiranthes gracilis,
Stachys palustris, Stellaria, Strophostyles pauciflora, Stylosanthes biflora,
Symphoricarpos, Tamarix, Trifolium hybridum, Verbena bracteosa, Veronica spicata,
Vicia. Predator: Philanthus gibbosus (Fabr.), P. politus Say.

Calliopsis andreniformis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 128. °.

Calliopsis flavipes Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 128. d.

Calliopsis lepidus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 68. °.

Panurgus vernalis Provancher, 1882. Nat. Canad. 13: 204. °, d.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1038, figs. 100-105 (larva). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 288-289, figs. 69-71, table 8 (redescription,
synonymy). —Rozen, 1966. Amer. Mus. Novitates 2259: 16, fig. 40 (larva). —Yager and
Rozen, 1966 Amer. Mus. Novitates 2265: 8-10, figs. 11-12 (pupa). —Shinn, 1967. Kans. Univ.
Sci. Bul. 46: 786-791, figs. 5-6, 8-13, map 1 (redescription).

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 159 (nest). —Swenk, 1907. Ent. News
18: 297 (nest, parasite). —Reinhard, 1924. Ann. Rept. Smithsn. Inst. 2738: 371-373
(predator). —Robertson, 1926. Psyche 33: 118 (nest, parasite). —Hendrickson, 1930. lowa
State Coll. Jour. Sci. 4: 163 (nest). —Pearson, 1933. Ecol. Monog. 3: 387 409-411 (floral
relationships). —Ainslie, 1937. Canad. Ent. 69: 97-100 (nest, floral relationships, parasite).
—Lovell and Lovell, 1989. Rhodora 41: 185 (ecology). —Crandall and Tate, 1947. Amer.
Soe. Agronomy, Jour. 39: 161-163 (nest site, floral relationships). — Mitchell, 1956. Elisha
Mitchell Sci. Soc. 72: 207 (parasite). —Michener and Rettenmeyer, 1956. Kans. Univ. Sci.
Bul. 37: 645 (age of nest site). —Montgomery, 1957. Ind. Acad. Sci., Proc. 66: 129 (ecology).
—Byers, 1962. Kans. Ent. Soc., Jour. 35: 320 (ecology). —Rozen, 1966. Amer. Mus.
Novitates 2244: 33 (parasite). —Shinn, 1967. Kans. Univ. Sci. Bul. 46: 895-935 figs. 144-156,
tables 4-9 (nest architecture, life history, immature stages, odor production, sleep, floral
relationships, parasites). —Rozen, 1967. Amer. Mus. Novitates 2297: 32-33, fig. 9, table 1
(nest architecture, life history, parasite). —Hurd and Linsley, 1972. Smithsn. Contrib. Zool.
114: 39, table 1 (parasites).

empelia Shinn. Ariz. (Douglas and Mt. Lemmon).
Calliopsis (Calliopsis) empelia Shinn, 1967. Kans. Univ. Sci. Bul. 46: 811, figs. 38-41, map
Py. Ce
helenae Shinn. Tex. (10 mi. s. Alice); Mexico (Piedras Negras, Coahuila).
Calliopsis (Calliopsis) helenae Shinn, 1967. Kans. Univ. Sci. Bul. 46: 814, map 1. °.
hondurasica Cockerell. La. (Robson), Tex.; Mexico to Panama. Pollen: Unknown, but visits
flowers of Cassia, Kallstroemia hirsutissima, Lippia, Nama undulatum, Phyla
strigosa, Teucrium, Trifolium repens, Verbena.
Calliopsis hondurasica Cockerell, 1949. U. S. Natl. Mus., Proc. 98: 487. 2.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 800-804, figs. 30-33, map 2 (redescription).

1868 | Hymenoptera in America North of Mexico

rhodophila Cockerell. N. Mex., Ariz. (Chino Valley), south. Calif. (Riverside, San Bernardino
and San Diego Counties); Mexico (Durango and Zacatecas). Pollen: Unknown, but visits

flowers of Solidago, Sphaeralcea.
Calliopsis andreniformis rhodophilus Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 350.

Cnaee
Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 796-799, figs. 22-25, map 1 (redescription).
squamifera Timberlake. Ariz. Pollen: Unknown, but visits flowers of Baccharis glutinosa,
Euphorbia, Medicago sativa.
Calliopsis squamifera Timberlake, 1947. Pan-Pacific Ent. 23: 28. 6.
Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 804-807, figs. 42-45, map 3 (redescription).
teucrii Cockerell. N. Mex., Ariz., Colo.; Mexico (Distrito Federal, Durango, Guerrero, Mexico,
Michoacan, Morelos, Nuevo Leon, Oaxaca, Puebla, Queretaro, San Luis Potosi, Tlaxcala
and Zacatecas). Pollen: Unknown, but visits flowers of Asclepias, Convolvulus incanus,
Heterotheca chrysopsidis, Melilotus officinalis, Oxalis, Potentilla, Stylosanthes,

Taraxacum, Teucrium laciniatum.
Calliopsis teucrii Cockerell, 1899. In Cockerell and Porter, Ann. and Mag. Nat. Hist. (7) 4:

A QEIOR
Genus CALLIOPSIS Subgenus PERISSANDER Michener

Calliopsis subg. Perissander Michener, 1942. N. Y. Ent. Soc., Jour. 50: 275.
Type-species: Calliopsis (Perissander) anomoptera Michener. Monotypic and orig.
desig.
Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 816-834, figs. 7, 46-57, maps 3-4
(redescription, included spp.).
anomoptera Michener. East. N. Mex., west to south. Calif.; Mexico (Baja California and
Sonora). Pollen: Polylectic, collects pollen and nectar from a variety of flowers including
Cladothrix lanuginosa, Eriogonum, Euphorbia albomarginata, E. capitellata, E.
hirtella, E. pleniradiata, E. polycarpa hirtella, E. polycarpa typica, Lepidium thurberi,
Tidestromia.
Calliopsis (Perissander) anomoptera Michener, 1942. N. Y. Ent. Soc., Jour. 50: 275. dg, 2.
Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 818-823, figs. 7, 46-49, map 4 (redescription).

Biology: Krombein, 1961. Ent. News 72: 82-83 (floral relationships with Euphorbia
albomarginata). —Shinn, 1967 Kans. Univ. Sci. Bul. 46: 823 (floral relationships).
fulgida Shinn. N. Mex. (4.8 mi. N. of Rodeo), Ariz. (5 mi. E. Portal). Pollen: Unknown, but
visits flowers of Euphorbia, Tidestromia lanuginosa.
Calliopsis (Perissander) fulgida Shinn, 1967. Kans. Univ. Sci. Bul. 46: 831, map 3 °.
gilva Shinn. Ariz. (Douglas and Pima Counties), N. Mex. (Hidalgo County). Pollen: Unknown,
but visits flowers of Euphorbia, Verbesina, Tidestromia lanuginosa.
Calliopsis (Perissander) gilva Shinn, 1967. Kans. Univ. Sci. Bul. 46: 829, figs. 54-57, map 4.
ORO
limbus Shinn. Ariz. (Santa Catalina Mts.), N. Mex. (Granite Pass). Pollen: Unknown, but visits
flowers of Euphorbia.
Calliopsis (Perissander) limbus Shinn, 1967. Kans. Univ. Sci. Bul. 46: 827, map 4. 2, 6.
rogeri Shinn. Ariz. (Cochise County). Pollen: Unknown, but visits flowers of Euphorbia
albomarginata, Lepidium thurberi.
Calliopsis (Perissander) rogeri Shinn, 1967. Kans. Univ. Sci. Bul. 46: 823, figs. 50-53, map
AD Gis

Genus CALLIOPSIS Subgenus CALLIOPSIMA Shinn

Calliopsis subg. Calliopsima Shinn, 1967. Kans. Univ. Sci. Bul. 46: 834.
Type-species: Calliopsis rozeni Shinn. Orig. desig.

Superfamily APOIDEA 1869

bernardinensis Michener. South. Calif. (Riverside and San Bernardino Counties). Pollen:
Apparently an oligolege of autumnal flowering Compositae including Gutierrezia
sarothrae, Heterotheca grandiflora, Senecio ionophyllus.
Calliopsis bernardinensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 323. d, 9.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 852-855, figs. 77-82, map 6 (redescription,
floral records).

chlorops Cockerell. N. Mex., Colo., Wyo., Idaho, Utah, Nev., Ariz. Pollen: Apparently an
oligolege of the Compositae; visits flowers of Chrysopsis, Grindelia squarrosa,
Haplopappus gracilis, Medicago sativa, Ratibida, Verbesina, Viguiera annua.

Calliopsis chlorops Cockerell, 1899. In Cockerell and Porter, Ann. and Mag. Nat. Hist. (7)
4: 413. 3.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 860-862, figs. 95-98, map 6 (redescription,
floral records).
coloradensis Cresson. Alta. to south. Utah and south. N. Mex., east to Mississippi River,

thence through east. Tex. and Gulf States to east slope of the Appalachian Mts. in N. C.
Parasite: Holcopasites arizonicus (Linsley). Pollen: Apparently an oligolege of the
Astereae and Heliantheae (Compositae); collects pollen from the flowers of Bidens
aristosa, Boltonia asteroides, Coreopsis tripteris, Rudbeckia triloba, Solidago
canadensis, but visits other flowers some of which may also serve as pollen sources
including Anthemis cotula, Aplopappus pluriflorus, Aster dumosus, A. praetus, Bidens
laevis, Chrysopsis, Chrysothamnus nauseosus, Eriocarpum gracile, Grindelia perennis,
G. squarrosa, Haplopappus, Helenium nudiflorum, Heterotheca subawxillaris, Macrotera,
Silphium, Solidago rigida, S. serotina.

Calliopsis coloradensis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 63. 2, 6.

Calliopsis coloratipes fedorensis Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 28. 2.

Taxonomy: Cockerell, 1908. Canad. Ent. 40: 147. 6. —Timberlake, 1951. Jn Muesebeck,
Krombein and Townes, U. S. Dept. Agr., Agr. Monog. 2: 1103 (synonymy). — Mitchell, 1960.
N. C. Agr. Expt. Sta. Tech. Bul. 141: 289-290, figs. 69-71 (redescription, floral records).
—Shinn, 1967. Kans. Univ. Sci. Bul. 46: 841-846, figs. 63-66, map 5, tables 2, 3
(redescription).

Biology: Hicks, 1926. Colo. Univ., Studies 15: 223 (nest). —Robertson, 1922. Psyche 29: 169
(floral relationships). —Shinn, 1967. Kans. Univ. Sci. Bul. 46: 846 (floral relationships).
—Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 39, table 1 (parasite).

coloratipes Cockerell. N. Mex., Ariz., Utah, Nev. Pollen: Unknown, but visits flowers of
Asclepias subverticillata, Bigelovia hariwegii, B. wrightti, Chrysothamnus nauseosus
consimilis, Heterotheca.
Calliopsis flavifrons race coloratipes Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 52.
3.

Taxonomy: Cockerell, 1898. Univ. N. Mex., Bul. 1: 52. 6. —Cockerell. 1899. Entomologist 33:
64. 2. —Shinn, 1967. Kans. Univ. Sci. Bul. 46: 862-865, figs. 99-102, map 5 (redescription,
floral records).

crypta Shinn. Ariz. (Chiricahua Mts.); Mexico (Chihuahua). Parasite: Holcopasites stevensi
(Cwfd.). Pollen: Collects pollen from Heterotheca subawxillaris, but also visits the flowers
of Cirsium, Helianthus.
Calliopsis crypta Shinn, 1965. Amer. Mus. Novitates 2211: 15, figs. 9-12. 2, d.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 14, figs. 29-33 (larva). —Shinn, 1967.
Kans. Univ. Sci. Bul. 46: 858-860, figs. 87-90, map 6 (redescription).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 30-31, fig. 10, tables 1-2 (nest architecture,
life history, floral relationship, parasite). —Hurd and Linsley, 1972. Smithsn. Contrib. Zool.
114: 39, table 1 (parasite).

deserticola Shinn. South. Calif., deserts. Pollen: Unknown, but visits flowers of Encelia
farinosa.
Calliopsis (Calliopsima) deserticola Shinn, 1967. Kans. Univ. Sci. Bul. 46: 865, figs.
103-106, map 5. 2, 6.

1870 Hymenoptera in America North of Mexico

pectidis Shinn. west. N. Mex., Ariz., south. Calif.; Mexico (Baja California). Parasite:
Holcopasites arizonicus (Linsley)? Pollen: Unknown, but visits flowers of Aplopappus,
Baccharis, Baileya pleniradiata, Helianthus, Heterotheca subaxillaris, Hymenothrix
wislizeni, Melilotus alba, Pectis angustifolia, P. papposa, Tidestromia lanuginosa,
Verbesina encelioides.
Calliopsis pectidis Shinn, 1965. Amer. Mus. Novitates 2211: 10, figs. 5-8. 2, d.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 846-850, figs. 67-71, map 6 (redescription).

Biology: Shinn, 1965. Amer. Mus. Novitates 2211: 15 (parasite). —Shinn, 1967. Kans. Univ.
Sci. Bul. 46: 850, 920 (floral records, parasite). —Hurd and Linsley, 1972. Smithsn. Contrib.
Zool. 114: 39, table 1 (parasite).

pugionis Cockerell. South. Calif. (cismontane). Pollen: Apparently an oligolege of Compositae,
visits flowers of Chaenactis artemisiaefolia, C. glabriuscula, Coreopsis (cultivated), C.
lanceolata, Encelia farinosa, Hemizonia wrightii, Oenothera veitchiana.
Calliopsis pugionis Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 197. @.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 868-870, figs. 107-110, map 5 (redescription).

rozeni Shinn. Tex., N. Mex., Ariz.; Mexico (Chihuahua and Coahuila). Parasite: Holcopasites
stevensi (Cwfd.)? Pollen: Collects pollen from Heterotheca subaxillaris primarily, but
visits other flowers including Baccharis, Baileya pleniradiata, Chamaesaracha
conioides, Eriogonum gracile, Gaillardia, Helianthus, Hymenoxys odorata, Melilotus
alba, Parkinsonia, Pectis papposa, Psilostrophe cooperi, Sphaeralcea emoryi, Verbesina
exauria.

Calliopsis rozeni Shinn, 1965. Amer. Mus. Novitates 2211: 2, figs. 1-4. 2, d.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 14-16, figs. 34-39 (larva, as rozeni?).
—Shinn, 1967. Kans. Univ. Sci. Bul. 46: 838-841, figs. 58-62, map 5 (redescription).

Biology: Rozen, 1965. N. Y. Ent. Soc., Jour. 73: 88 (parasite). —Rozen, 1967. Amer. Mus.
Novitates 2297: 31-32, tables 1-2 (nest architecture, life history, floral relationships,
parasite). —Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 39, table 1 (parasite).

timberlakei Shinn. West. Tex., N. Mex., Ariz., Utah. Pollen: Apparently an oligolege of
Compositae, visits flowers of Bahia absinthifolia, Bigelovia, Chrysothamnus nauseosus,
Gutierrezia longifolia, G. microcephala, G. sarothrae, Haplopappus spinulosus, H.
wrightii.
Calliopsis (Calliopsima) timberlakei Shinn, 1967. Kans. Univ. Sci. Bul. 46: 850, figs. 72-76,
map 7. °, 6.
unca Shinn. N. Mex. (3 mi. W. Bingham). Pollen: Unknown, but visits flowers of Baileya
pleniradiata.
Calliopsis (Calliopsima) unca Shinn, 1967. Kans. Univ. Sci. Bul. 46: 855, figs. 83-86, map 7.
é.

Genus CALLIOPSIS Subgenus VERBENAPIS Cockerell and Atkins
Calliopsis subg. Verbenapis Cockerell and Atkins, 1902. Ann. and Mag. Nat. Hist. (7) 10:
44,

Type-species: Calliopsis verbenae Cockerell and Porter. Monotypic.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 876-891, figs. 123-143, map 8 (redescription,
included spp.).
hirsutifrons Cockerell. West. Tex. (El Paso), N. Mex.; Mexico (Chihuahua, Guanajuato, Hidalgo

and Mexico).
Calliopsis hirsutifrons Cockerell, 1896. Canad. Ent. 28: 158. 3.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 887-889, figs. 134-138. ?, d (redescription).
micheneri Shinn. Tex. (Carrizo Springs). Pollen: Unknown, but visits flowers of Verbena
cloveri.
Calliopsis (Verbenapis) micheneri Shinn, 1967. Kans. Univ. Sci. Bul. 46: 889, figs. 139-143,
map 8. @, dc.

Superfamily APOIDEA 1871

nebraskensis Crawford. N.J., Wis. west to N. Dak., 8S. Dak., Nebr., Colo., south to Il., Ark.
and Kans. Parasite: Holcopasites heliopsis (Robt.), Sphecodes sp.? Pollen: Apparently
polylectic since mixed pollen loads from flowers of Leguminosae and Verbenaceae have

been observed; visitation records include flowers of Ambrosia, Asclepias, Medicago
sativa, Verbena hastata, V. stricta, V. urticaefolia, Vernonia.

Calliopsis verbenae var. Nebraskensis Crawford, 1902. Canad. Ent. 34: 240. 2, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 291-292, figs. 69-71, table 8
(redescription). —Shinn, 1967. Kans. Univ. Sci. Bul. 46: 882-887, figs. 129-133, map 8
(redescription).

Biology: Robertson, 1914. Ent. News 25: 72 (floral relationships, as verbenae). —Rau and
Rau, 1916. Jour. Anim. Behavior 6: 368 (nesting habits, mating). —Robertson, 1922. Psyche
29: 171 (floral relationships, as verbenae). —Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7):
33 (nesting habits, mating behavior, parasite). —Shinn, 1967. Kans. Univ. Sci. Bul. 46:
886-887 (summary of bionomics including floral relationships). —Hurd and Linsley, 1972.
Smithsn. Contrib. Zool. 114: 39, table 1 (parasite).

verbenae Cockerell and Porter. West. Tex. to southeast. Ariz.; Mexico (Durango). Pollen:

Unknown, but visits flowers of Chamaesaracha conioides, Sphaeralcea lobata, Verbena
bipinnatifida, V. macdougalli.

Calliopsis verbenae Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 412. 9, d.

Taxonomy: Shinn, 1967. Kans. Univ. Sci. Bul. 46: 878-881, figs. 123-128, map 8 (tax. status
redescription, flower records).

NOMEN NUDUM IN CALLIOPSIS SMITH

Calliopsis bridwelli Bridwell, 1899. Kans. Acad. Sci., Trans. 16: 210.

UNPLACED TAXON IN CALLIOPSIS SMITH
flavifrons Smith. Va., Fla.

Calliopsis flavifrons Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 129. ¢.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 321. 2. —Mitchell, 1960. N. C. Agr

Expt. Sta. Tech. Bul. 141: 293 (tax. status). —Shinn, 1967. Kans. Univ. Sci. Bul. 46: 392
(tax. status).

Genus HYPOMACROTERA Cockerell and Porter

Hypomacrotera Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 418.
Type-species: Hypomacrotera callops Cockerell and Porter. Orig. desig.
Biology: Rozen, 1970. Amer. Mus. Novitates 2416: 1-16, 19 figs., table 1 (nest architecture,
life history, mating habits, immature stages, nest associates).
callops callops Cockerell and Porter. Colo., N. Mex., Ariz. Parasite: Holcopasites illinoiensis
minimus (Linsley)? Pollen: Collects pollen from the flowers of Sphaeralcea.

Hypomacrotera callops Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 419. d,
2.

Taxonomy: Cockerell, 1937. Amer. Mus. Novitates 948: 3. —Rozen, 1970. Amer. Mus.
Novitates 2416: 7-9, 15-16, figs. 3-8, 18-19 (larva, pupa).

Biology: Rozen, 1970. Amer. Mus. Novitates 2416: 2-5, table, 1 (nest architecture, nesting
habits, pollen source). —Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 36, table 1
(parasite).

callops persimilis Cockerell. Ariz., south. Calif.; Mexico (Baja California and Sonora). Parasite:
Holcopasites illinoiensis minimus (Linsley). Pollen: Collects pollen from the flowers of
Physalis, but visits other flowers for nectar including Melilotus alba, Sida hederacea.
Hypomacrotera callops persimilis Cockerell, 1899. Entomologist 33: 64. 2, d.

Biology: Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 36, table 1 (parasite).
subalpinus andradensis Cockerell. N. Mex. to south. Calif.; Mexico (Baja California and
Sonora). Pollen: Collects pollen from the flowers of Sphaeralcea including S. ambigua, S*

1872 Hymenoptera in America North of Mexico

angustifolia, S. emoryi, S. orcutti, but visits other flowers for nectar including Baileya
multiradiata, B. pleniradiata.
Hypomacrotera andradensis Cockerell, 1937. Amer. Mus. Novitates 948: 3. °, 3.
subalpinus subalpinus (Cockerell). N. Mex., Ariz.; Mexico (Sonora). Parasite: Oreopasites sp.
Pollen: Collects pollen from the flowers of Sphaeralcea including S. ambigua, S. orcutti.
Calliopsis subalpinus Cockerell, 1894. Ent. News 5: 235. 6.
Calliopsis semirufus Cockerell, 1896. Ent. Monthly Mag. 32: 219. 2.

Biology: Rozen, 1970. Amer. Mus. Novitates 2416: 2-5, table 1 (nest architecture, nesting
habits, pollen source, parasite).

Genus XENOPANURGUS Michener

Xenopanurgus Michener, 1952. Kans. Ent. Soc., Jour. 25: 24.
Type-species: Xenopanurgus readioi Michener. Orig. desig.

Taxonomy: Shinn, 1964. Ent. News 75: 73-78 (included sp.).

readioi Michener. Ariz. (Huachuca Mts.); Mexico (Toluca).
Xenopanurgus readioi Michener, 1952. Kans. Ent. Soc., Jour. 25: 25, 5 figs. ¢.

Taxonomy: Shinn, 1964. Ent. News 75: 74-78. 2, ¢.
Genus PERDITA Smith

This is a very large genus of pollen-collecting bees that has established through its component
species a predominantly oligolectic relationship with the flora of North America. Even though
the genus occurs transcontinentally in southern Canada, the United States and Mexico to as far
south as Central America (Guatemala), it is centered in the more arid areas of northern Mexico
and adjacent southwestern United States. The nesting habits of only a few species have been
studied and these, like other species of the family, make their nests in the ground, although one
species, Perdita (Cockerellula) opuntiae Cockerell, is known to excavate its nests in sandstone.
Several females of at least some species occupy a single nest, but there is no evidence of caste
development.

In recognition of utilitarian advantages, the subgenera are arranged alphabetically in this
catalog.

Revision: Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 25-107. —Cockerell, 1922. Amer.
Mus. Novitates 33: 14 (partial key). —Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 345-432,
172 figs. (all subgenera except Hexaperdita Timberlake, Perditella Cockerell, Alloperdita
Viereck, and typical Perdita Smith). —Timberlake, 1956. Calif. Univ. Pubs. Ent. 11:
247-350, figs. 173-367 (treats subgenera Alloperdita Viereck, Perditella Cockerell,
Hexaperdita Timberlake, and Pygoperdita Timberlake). —Timberlake, 1958. Calif. Univ.
Pubs. Ent. 14: 303-410, figs. 368-521 (treats zonalis and halictoides groups of subgenus
Perdita). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 1-156, figs., 522-732 (treats
octomaculata group of subgenus Perdita). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28:
1-124, figs. 733-904, 2 text figs. (treats ventralis group of subgenus Perdita, and suppl. to
Parts I-IV). —Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 125-388, figs. 905-1190, 77 text
figs. (treats sphaeralceae group of subgenus Perdita). —Timbelake, 1968. Calif. Univ.
Pubs. Ent. 49: 1-196, figs. 1191-1360, 31 text figs. (suppl. to preceding parts, bibliography,
index to Parts I-VII, corrigenda). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 1-63, figs.
1361-1417 (suppl. to Parts I-VII, treats subgenera Cockerellia, Epimacrotera,
Hexaperdita, Pentaperdita, Perdita, Procockerellia, Pygoperdita, Xeromacrotera).

Taxonomy: Cockerell, 1922. Amer. Mus. Novitates 33: 1-15 (west. U.S. spp., species
formation). —Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 199-204 (eastern U. S. spp.).
—Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 961-985 (treats spp. of the subgenera
Cockerellia, Cockerellula, Macroteropsis, Xerophasma). —Mitchell, 1960. N. C. Agr. Expt.
Sta. Tech. Bul. 141: 295-330, figs. 58, 72-79, table 9 (eastern U. S. spp.). —Rozen, 1966.
Amer. Mus. Novitates 2259: 5, 16-21, figs. 41-54 (larvae). —Yager and Rozen, 1966. Amer.
Mus. Novitates 2265: 12 (pupae).

Superfamily APOIDEA 1873

Biology: Linsley, 1958. Hilgardia 27: 562, tables 4, 5 (pollen sources of some subgenera).
—Rozen, 1967. Amer. Mus. Novitates 2297: 1-44, 18 figs., 2 tables (review). —Hurd and
Linsley, 1975. Smithsn. Contrib. Zool. 193: 24-26, figs. 5-7 (Larrea visiting spp.).

Genus PERDITA Subgenus ALLOMACROTERA Timberlake

Perdita subg. Allomacrotera Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 131.
Type-species: Perdita stephanomeriae Timberlake. Monotypic and orig. desig.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 5-6 (key to spp. of subgenera
Allomacrotera, Procockerellia).
stephanomeriae Timberlake. Calif. (Colorado and Mojave Deserts). Pollen: Possibly an
oligolege of Stephanomeria including S. pauciflora, a male has been collected at the
flowers of Dalea spinosa.
Perdita stephanomeriae Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 404. 9.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 60: 131-132, figs. 658, 659, 731. ¢.

Genus PERDITA Subgenus ALLOPERDITA Viereck

Perdita subg. Alloperdita Viereck, 1917. Amer. Mus. Nat. Hist., Bul. 37: 241.
Type-species: Perdita novae-angliae Viereck. Monotypic.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 348 (key). —Timberlake, 1956. Calif.
Univ. Pubs. Ent. 11: 271-275 (redescription, key to spp.). —Timberlake, 1958. Calif. Univ.
Pubs. Ent. 14: 393 (modification of key). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17:
134. —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 30.

bradleyi Viereck. N. J. to Fla. Pollen: Unknown, but visits flowers of Batodendron, Crataegus,
Hydrocotyle wmbellatum, Ilex, Nyssa sylvatica, Pyracantha, Sarracenia flava.
Perdita bradleyi Viereck, 1907. Ent. News 18: 393. 6.

Taxonomy: Timberlake, 1928. Amer. Mus. Novitates 321: 7 (as obscurata). —Timberlake,
1956. Calif. Univ. Pubs. Ent. 11: 274, figs. 211, 212, 306 (as obscwrata). —Timberlake, 1958.
Calif. Univ. Pubs. Ent. 14: 394 (as obscurata). —Timberlake, 1960. Calif. Univ. Pubs. Ent.
17: 134 (type). —Mitchell, 1964. N. C. Agr. Expt. Sta. Tech. Bul. 141: 310-311, figs. 72-73,
table 9 (redescription).

floridensis Timberlake. N. C. to Fla. Pollen: Unknown, but visits flowers of Batodendron, Ilex
glabra, Leucothoe.
Perdita floridensis Timberlake, 1928. Amer. Mus. Novitates 321: 7. d, 9.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 278, 274 (key, geogr. records).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 393 (key). —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 311, figs. 72-74, 79, table 9 (redescription).

gertschi Timberlake. Tex. (Brewster County); Mexico (Coahuila). Pollen: Unknown, but visits
flowers of Prosopis juliflora.
Perdita (Alloperdita) gertschi Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 398, figs. 468,
469, 519. 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 30-31. °.
mitchelli Timberlake. N. C., Fla., Ala., Miss. Pollen: Unknown, but visits flowers of Ceanothus,

Cyrilla racemiflora, Oxydendrum.
Perdita mitchelli Timberlake, 1947. Ent. Soc. Wash., Proc. 49: 81. 2, d.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 273, 274, figs. 213-214, 307 (key,
geogr. records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 311-312, figs. 73-74,
table 9 (redescription).

novaeangliae Viereck. Mass. to Fla. Pollen: Unknown, but visits flowers of Batodendron,
huckleberry, Lyonia ligustrina.
Perdita novae-angliae Viereck, 1907. Ent. News 18: 394. 2, d.
Perdita novaeangliae Viereck, 1917. Amer. Mus. Nat. Hist., Bul. 37: 241. Emend.

1874 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 273, 274, 275, figs. 219, 220, 310 (key,
genitalia). —Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 393 (key). —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 312-313, figs. 72-73, table 9 (redescription).

obscurata Cresson. N. C. to Tex. Parasite: Neolarra cockerelli (Cwfd.)? Pollen: Unknown, but
visits flowers of Batodendron, Diospyros virginiana, Stillingia sylvatica.
Perdita obscurata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 70. 2, 6.
Perdita carolina Timberlake, 1952. Ent. Soe. Wash., Proc. 54: 199. 2, do.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 273, 274, figs. 215, 216, 308 (key,
genitalia, geogr. and floral records, as carolina). —Timberlake, 1960. Calif. Univ. Pubs.
Ent. 17: 134 (synonymy). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 313-314,
figs. 72-74, 79, table 9 (synonymy, redescription).

townesi Timberlake. N. C. to Fla. Pollen: Unknown, but visits flowers of Batodendron,
Ceanothus microphyllus, Diospyros virginiana, Ilex, Nyssa sylvatica.
Perdita townesi Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 31, figs. 1209-1210, 1308. 6,
oe

Genus PERDITA Subgenus COCKERELLIA Ashmead

Cockerellia Ashmead, 1898. Psyche 8: 284.
Type-species: Perdita albipennis Cresson. Monotypic and orig. desig.
(=Perdita hyalina Cresson).
Philoxanthus Ashmead, 1898. Psyche 8: 285.
Type-species: Perdita beata Cockerell. Monotypic and orig. desig.

The species of this subgenus are found almost exclusively at flowers of Compositae, mostly of
the genera Baileya, Coreopsis, Gaillardia, Geraea, Helianthus, Ratibida, Rudbeckia and Ver-
besina, but also occur at the flowers of Erigeron, Heterotheca, Pectis and Prionopsis.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 971-974 (key to females of included
spp.). —Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 347, 385-402 (key to included spp.).
—Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 326. —Timberlake, 1958. Calif. Univ. Pubs.
Ent. 14: 382-383. —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 19-22 (modification of
key). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 2-5 (key to spp. allied to hilaris).

albihirta albihirta Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an
oligolege of Geraea canescens, but visits the flowers of Encelia farinosa and Melilotus
presumably for nectar.
Perdita albthirta albihirta Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 395, fig. 79. 9, 3.
albihirta geraeae Timberlake. Calif. (Colorado and Mojave Deserts), Nev. (Stateline). Pollen:
Apparently an oligolege of Geraea canescens, but visits the flowers of Acamptopappus
sphaerocephalus var. hirtellus and Larrea tridentata presumably for nectar.
Perdita albihirta geraeae Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 396. 2, ¢.

albipennis albipennis Cresson. N. Dak. south to Tex., west to Wyo., Colo. and Ariz. Pollen:
Unknown, but visits flowers of Helianthus annuus, H. petiolaris, Hymenopappus,
Nolina, Prionopsis ciliata. Predator: Philanthus ventilabris Fabr.
Perdita(?) albipennis Cresson, 1868. Amer. Ent. Soc., Trans. 1: 386. 9.
Perdita hyalina Cresson, 1878. Amer. Ent. Soc., Trans. 7: 68. 3.
Perdita albipennis var. helianthi Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 104. 6,
g

Perdita albipennis var. helianthi mut. pasonis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc.
48: 105. d.
Perdita lacteitpennis Swenk and Cockerell, 1907. Ent. News 18: 51. 2, ¢.

Taxonomy: Cockerell, 1899. Psyche 8: 419. 3 (as hyalina). —Cockerell, 1899. Psyche 8: 419.
6 (as pasonis). —Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 977-978 (geogr. and floral
records, as lacteipennis lacteipennis). —Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 388,
390, 391, 398, 400 (geogr. and floral records, in part, as albipennis, lacteipennis
lacteipennis, pasonis and hyalina). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 130-131
(types, synonymy, status discussed). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 20
(synonymy, status discussed).

Superfamily APOIDEA 1875

Biology: Aleorn and Gamboa, 1975. Ariz. Acad. Sci. 10: 163 (predator).

albipennis canadensis Crawford. Alta., Idaho, N. Dak., Utah (Uinta and Wasatch Counties).
Perdita canadensis Crawford, 1912. Canad. Ent. 44: 360. °.

Taxonomy: Stevens, 1919. Canad. Ent. 51: 205. 6. —Timberlake, 1954. Calif. Univ. Pubs. Ent.
9: 388, 390, 398-399 (key, tax. status, as lacteipennis canadensis). —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 130-131 (status discussed).

albipennis heliophila Cockerell. Utah, Colo., N. Mex.; Mexico (Chihuahua). Pollen: Unknown,
but visits flowers of Helianthus including H. annuus.
Perdita heliophila Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 281. @.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 390, 397, figs. 82, 83, 158 (tax.
characters and tax. status). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 20 (tax.
characters). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 2 (tax. characters).

albipennis pallidipennis Graenicher. Ind. and Wis., west to east. Kans. and Nebr. Pollen:
Unknown, but visits flowers of Helianthus annuus.
Perdita pallidipennis Graenicher, 1910. Canad. Ent. 42: 101. 2, 6.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 978 (tax. characters and status, as
lacteipennis pallidipennis). —Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 388, 390, 399
(key, geogr. records). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 131 (status
discussed). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 302-303, table 9
(redescription).

beata beata Cockerell. Tex. (Davis Mts. and El Paso), N. Mex. (Las Cruces and Mesilla Park).
Pollen: Unknown, but visits flowers of Verbesina encelioides. Another subspecies,
Perdita beata signata Timberlake, occurs in Mexico (Chihuahua, Coahuila and Durango).
Perdita beata Cockerell, 1895. Psyche 1 (sup.): 10. °.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 386, 391 (key, geogr. and floral
record). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 130 (tax. status).
bequaerti bequaerti Viereck. N. J. to Fla., west to Minn. and Miss. Pollen: Unknown, but visits
flowers of Bidens, Helianthus divaricatus, Physalis lanceolata, P. virginiana.
Perdita (Cockerellia) bequaerti Viereck, 1917. Amer. Mus. Nat. Hist., Bul. 37: 241. 2, d.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 397, figs. 80, 157 (tax. characters).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 300, figs. 72-74, 79, table 9
(redescription).

bequaerti indianensis Cockerell. Ind. (Gary and Lafayette), Okla. (South McAlester), Tenn.
(Knoxville).
Perdita pallidipennis indianensis Cockerell, 1922. Amer. Mus. Novitates 33: 2. d, ?.
Perdita (Cockerellia) wickhami Cockerell, 1922. Amer. Mus. Novitates 33: 13. °.

Taxonomy: Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 204 (tax. status). —Timberlake, 1953.
Kans. Univ. Sci. Bul. 35: 976 (tax. characters, synonymy). —Timberlake, 1954. Calif. Univ.
Pubs. Ent. 9: 388, 390, 397 (key, tax. status, geogr. range). —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 302, fig. 72 (status discussed).

coreopsidis collaris Cockerell. N. Mex.; Mexico (Chihuahua).
Perdita verbesinae collaris Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 281. ¢.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 389, 390, 401 (key, redescription).
coreopsidis coreopsidis Cockerell. Kans., Tex.; Mexico (Coahuila). Pollen: Unknown, but visits
flowers of Gaillardia which it apparently prefers, but also has been taken at the

flowers of Coreopsis and Monarda punctata cory.
Perdita coreopsidis Cockerell, 1906. Entomologist 39: 126. ?, d.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 983-984 (geogr. and floral records).
—Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 389, 390, 401 (key, geogr. and floral records).
—Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 4 (key).

1876 Hymenoptera in America North of Mexico

coreopsidis kansensis Timberlake. Kans. (Grove and Ford Counties), Tex. (Van Horn), N.
Mex. (Roosevelt County), Utah (Iron County). Pollen: Unknown, but visits flowers of
Gaillardia.

Perdita coreopsidis kansensis Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 984. 2, d.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 389, 390, 401 (key, geogr. range).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 20 (geogr. records).
coreopsidis obscurior Timberlake. Gulf Coast of Tex. (Galveston to Brownsville, but a male
has been taken inland at Lytle, Atacosa County). Pollen: Unknown, but visits flowers of
Gaillardia.
Perdita coreopsidis obscurior Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 984. 2, 6.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 389, 390, 401 (key, geogr. records).

fracticincta Timberlake. Tex. (Sarita). Pollen: Unknown, but visits flowers of Coreopsis.
Perdita fracticincta Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 976. female.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 388, 397 (key).
hilaris Timberlake. Utah (Dixie State Park), N. Mex. (San Juan County). Pollen: Unknown, but

visits flowers of Chrysothamnus nauseosus.
Perdita hilaris Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 21, figs. 1207, 1208, 1307. 6.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 2-3. 9.
imbellis Timberlake. Ariz. (Coconino County).
Perdita imbellis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 21, figs. 1205, 1206, 1306. ¢.
incana Timberlake. Tex. (Culberson County), N. Mex. (Quay County).
Perdita incana Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 383. 2, ¢.
lepachidis lepachidis Cockerell. Kans., Tex., Colo., N. Mex. Pollen: Unknown, but visits

flowers of Ratibida tagetes.
Perdita lepachidis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 106. 6.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 982-983. 2. —Timberlake, 1954. Calif.
Univ. Pubs. Ent. 9: 389, 391, 401 (key). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 4
(key).

lepachidis levifrons Timberlake. Tex. (Brownsville).
Perdita lepachidis levifrons Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 983. 2, 6.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 389, 391, 401 (key).
lingualis Cockerell. Nebr., Kans., Tex., N. Mex., Colo., Utah; Mexico (Tamaulipas). Pollen:
Apparently an oligolege of Helianthus including H. annuus, H. petiolaris, but may also

visit other large flowered Compositae.
Perdita albipennis var. lingualis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 105. @.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 130, 181 (tax. status). —Michener,
1963. Kans. Ent. Soc., Jour. 36: 117-118, figs. 2-6 (prepupa). —Rozen, 1966. Amer. Mus.
Novitates 2259: 21, figs. 52-54 (larva). —Yager and Rozen, 1966. Amer. Mus. Novitates
2265: 12, figs. 17-18 (pupa). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 4 (key).

Biology: Michener, 1963. Kans. Ent. Soc., Jour. 36: 114-118, 6 figs. (nest architecture, colonial
behavior, floral relationships).
luculenta Timberlake. South. Calif. (Imperial County).
Perdita luculenta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 19. @.
perpulchra flavidior Timberlake. Kans., (Barton and Rice Counties), Tex. (Somervel County).
Pollen: Unknown, but visits flowers of Aphanostephus skirrhobasis, Heterotheca

subaxillaris, Rudbeckia bicolor.
Perdita perpulchra flavidior Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 975. 9.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 395. —Timberlake, 1956. Calif. Univ.
Pubs. Ent. 11: 327 (geogr. and floral records).
perpulchra perpulchra Cockerell. Tex. (E] Paso), N. Mex. (Mesilla Valley), Ariz. (Holbrook and
Tucson); Mexico (Aguascalientes and Durango). Pollen: Unknown, but visits flowers of
Asclepias, Baileya, Haplopappus heterophylla, Verbesina encelioides.
Perdita perpulchra Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 99. °.

Superfamily APOIDEA 1877

Perdita baileyae Cockerell, 1909. Canad. Ent. 41: 151. 6.

Taxonomy: Cockerell, 1897. Ann. Mag. and Nat. Hist. (6) 19: 398. 6. —Timberlake, 1953.
Kans. Univ. Sci. Bul. 35: 975 (tax. characters). —Timberlake, 1954. Calif. Univ. Pubs. Ent.
9: 387, 389, 395 (key, tax. characters). —Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 327
(synonymy, tax. status).
perpulchra punctissima Timberlake. Tex. (Bexar, Wood, Robertson, Lamar Counties). Pollen:
Unknown, but visits flowers of Heterotheca subaxillaris.
Perdita perpulchra punctissima Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 327. 2, ¢.
pulchella Timberlake. Tex. (Carrizo Springs).
Perdita pulchella Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 393. 9.
purpurascens Timberlake. Tex. Pollen: Unknown, but visits flowers of Aster tanacetifolius,
Brazoria truncata, Gaillardia, Monarda.
Perdita purpurascens Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 328. 6.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 382-383. 2 (tax. characters, geogr.
and floral records).
scopata Timberlake. Gulf Coast of Tex. (Galveston.to Brownsville). Pollen: Unknown, but

visits flowers of Coreopsis, Helianthus annuus.
Perdita scopata Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 980. 2, ¢.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 401, fig. 88 (tax. characters).
shinnersi Timberlake. Tex. Pollen: Unknown, but visits flowers of Brazoria truncata,

Coreopsis basalis, Gaillardia, Hymenopappus artemisaefolius, Monarda.
Perdita shinnersi Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 326. 9, 6.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 383 (geogr. and floral records).
tricincta Timberlake. South. Tex. (Cameron, Hidalgo and Starr Counties). Pollen: Unknown,
but visits flowers of Helianthus annuus. This may prove to be a race of Perdita

albipennis Cress.
Perdita tricincta Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 978. °, 6.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 388, 390, 400, fig. 86 (key, tax.
status).
utahensis Cockerell. N. Mex. (Rodeo), Ariz. (Tucson), southwest. Utah, south. Calif. (Colorado
Desert). Pollen: Unknown, but visits flowers of Pectis papposa, Tidestromia, Verbesina

encelioides.
Perdita utahensis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 106. 9.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 974. 6. —Timberlake, 1954. Calif.
Univ. Pubs. Ent. 9: 387, 391, 394, figs. 76, 155 (tax. characters, geogr. and floral records).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 20 (geogr. and floral records).

verbesinae Cockerell. Tex. (Barstow), N. Mex. (Deming and Las Cruces), Ariz. (Douglas,

Portal, Tucson and Santa Rita Mts.); Mexico (Chihuahua). Pollen: Usually visits flowers
of Verbesina encelioides, but has been recorded from the flowers of Helianthus.

Perdita verbesinae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 102. 2, d.

Perdita verbesinae mut. intermedia Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 103. °.

Perdita verbesinae mut. nigrior Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 103. °.

Perdita verbesinae mut. cyanella Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 104. d.

Perdita verbesinae mut. maculata Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 104. ¢.

Taxonomy: Cockerell, 1899. Psyche 8: 323 (color variation). —Timberlake, 1954. Calif. Univ.
Pubs. Ent. 9: 388, 391, 397, fig. 81 (key, tax. characters, geogr. and floral records).
Genus PERDITA Subgenus COCKERELLULA Strand

Perdita subg. Lutziella Cockerell, 1922. Amer. Mus. Novitates 47: 1. Preoce.
Type-species: Perdita (Lutziella) opuntiae Cockerell. Monotypic and orig. desig.
Perdita subg. Cockerellula Strand, 1932. Folia Zool. Hydrobiol. Riga 4: 196. N. name.

1878 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 963-968 (tax. characters, key to
included spp.). —Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 346, 354-356 (key to included
spp., tax. characters). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 122-123 (key to males
of included spp.). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 2 (modified key to males).

bidenticauda Timberlake. Tex. (Big Bend Park). Pollen: Unknown, but visits flowers of
Opuntia.
Perdita bidenticauda Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 965. d, °.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 356, figs. 18, 14, 122 (tax. characters,
key). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 122 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 2 (key).

knulli Timberlake. Tex. (Davis Mts.). Pollen: Unknown, but the female allotype bears dark
brown pollen, with spherical and moderately fine grains.
Perdita knulli Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 128, figs. 652, 653, 728. 3, 2.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 2. ¢ (key).
laticauda Timberlake. Tex. (Dryden); Mexico (Coahuila). Pollen: Unknown, but visits flowers

of Chamaesaracha conioides, Gaillardia, Gilia acerosa.
Perdita laticauda Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 967. 6, @.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 356, figs. 15, 16, 123 (tax. characters,
key). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 122. 3 (key).
lobata Timberlake. Tex. (Hidalgo and Starr Counties). Pollen: Unknown, but visits flowers of
Opuntia.
Perdita lobata Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 967. 6.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 354, 356, figs. 17, 18, 124 (tax.
characters, key). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 122. 6, 2 (tax. characters,
key).

opuntiae Cockerell. Colo., Wyo., S. Dak. (Pine Ridge). Ecology: Nests in soft sandstone rocks
of the Laramie formation. Pollen: Collects pollen from the flowers of Opuntia.
Perdita (Lutziella) opuntiae Cockerell, 1922. Amer. Mus. Novitates 47: 2. 6, 2.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 964 (key). —Timberlake, 1954. Calif.
Univ. Pubs. Ent. 9: 354, 355, figs. 9, 10, 120 (tax. characters, key). —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 122. 5 (key)

Biology: Custer, 1928. Psyche 35: 67-84 (nesting habits). —Custer, 1929. Canad. Ent. 61: 49
(nesting habits). —Custer, 1929. Psyche 36: 293 (nesting habits).

Genus PERDITA Subgenus EPIMACROTERA Timberlake

Perdita subg. Epimacrotera Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 377.
Type-species: Perdita ainsliei Crawford. Orig. desig.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 378-379 (key to included spp.).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 380 (modified key to included spp.).
—Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 129 (tax. characters, relationship to
Glossoperdita). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 92-93 (revised key to
included spp.). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 14-15 (key to males).

ainsliei Crawford. Iowa, N. Mex. (Bernalillo County).
Perdita ainsliei Crawford, 1932. Ent. Soc. Wash., Proc. 34: 74. d, @.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 378, 379, figs. 55, 56, 144 (key, tax.
characters). —Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 380 (key, geogr. record).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 93 (key). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 14 (key).

biguttata Timberlake. Ariz. (Cochise County). Pollen: Unknown, but visits flowers of
Eriogonum abertianum var. neomexicanum, Euphorbia.
Perdita biguttata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 95. @.
Perdita parvula Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 15, figs. 1202, 1304. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 15 (synonymy in footnote).

Superfamily APOIDEA 1879

binotata Timberlake. Ariz.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Physalis.
Perdita binotata Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 385, figs. 78, 74, 153. 2, d.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 93 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 14 (key).
crassula Timberlake. N. Mex. (Las Cruces), Ariz. (Willcox). Pollen: Unknown, but visits

flowers of Euphorbia.
Perdita crassula Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 381, figs. 462, 463, 516. d.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 93-94. 2 (key). —Timberlake, 1968.
Calif. Univ. Pubs. Ent. 49: 14 (key).
diversa Timberlake. Tex. (E] Paso); Mexico (Coahuila, Nuevo Leon). Pollen: Unknown, but
visits flowers of Coldenia greggit.
Perdita diversa Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 384, figs. 69, 70, 151. d, 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 15, 19 (key).

euphorbiae Timberlake. Ariz. (Picacho Pass); Mexico (Sonora). Pollen: Unknown, but visits
flowers of Euphorbia, Kallstroemia.
Perdita euphorbiae Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 380, figs. 59, 60, 146. 2, 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 130 (geogr. and floral records).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 93 (key). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 15, 17 (key, geogr. record).

namatophila Timberlake. Calif. (Riverside County). Pollen: Unknown, but visits flowers vf

Nama demissum.
Perdita namatophila Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 379, figs. 57, 58, 145. 3.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 380 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 15 (key).
nanula Timberlake. Ariz. (Douglas). Pollen: Unknown, but visits flowers of Euphorbia

albomarginata.
Perdita nanula Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 129. 9.
Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 93 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 15, 16. ¢ (key).
nigrocaerulea Timberlake. Calif. (Oro Grande Wash). Pollen: Unknown, but visits flowers of
Salazaria mexicana.
Perdita nigrocaerulea Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 382, figs. 65, 66, 149. ¢.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 15 (key).
peculiaris Timberlake. Tex. (Terrell County). Pollen: Unknown, but visits flowers of
Chamaesaracha conioides.
Perdita peculiaris Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 970. d.
Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 384, figs. 71, 72, 152 (tax. characters). —
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 15 (key).
polycarpae Timberlake. South. Calif.; Mexico (Baja California). Pollen: Possibly an oligolege of
Euphorbia including E. polycarpa, but also visits flowers of Pectis papposa.
Perdita polycarpae Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 381, figs. 61, 62, 147. d, °.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 14 (key).
Genus PERDITA Subgenus GLOSSOPERDITA Cockerell
Glossoperdita Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 43.
Type-species: Glossoperdita pelargoides Cockerell. Monotypic.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 347, 392-394 (redescription, key to
included spp.). —Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 379 (key to included spp.).

blaisdelli Timberlake. Calif. (Calaveras County).
Perdita blaisdelli Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 373. °.

1880 Hymenoptera in America North of Mexico

giliae Timberlake. Ariz. Pollen: Unknown, but visits flowers of Gilia, possibly G. aggregata.
Perdita giliae Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 374, figs. 51, 52, 141. 2, ¢.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 379 (key).
hurdi Timberlake. South. Calif., Ariz.; Mexico (Sonora). Ecology: Cuts unopened flowers of
Proboscidea in order to obtain pollen. Pollen: Collects pollen from the flowers of
Proboscidea althaeafolia, P. arenaria.
Perdita hurdi Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 324, figs. 283, 284, 3385. 2, d.
Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 379 (key). —Timberlake, 1960. Calif.
Univ. Pubs. Ent. 17: 127-128 (tax. characters, geogr. and floral records).
Biology: Hurd and Linsley, 1963. Kans. Ent. Soc., Jour. 36: 248-252, 5 figs. (corolla cutting
habits, floral relationships).
navarretiae angusticeps Timberlake. Calif. (Tuolumne County).
Perdita navarretiae angusticeps Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 128. 6, 9.
navarretiae navarretiae Timberlake. Calif. (Mariposa County). Pollen: Unknown, but visits

flowers of Navarretia viscidula.
Perdita navarretiae Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 379, figs. 460, 461, 515. 3.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 128 (tax. status).
navarretiae powelli Timberlake. Calif. (Nevada County).
Perdita navarretiae powelli Timberlake, 1962. Calif. Univ. Pubs. Ent. 91. 5, °.
pelargoides (Cockerell). South. Calif. (cismontane). Pollen: Unknown, but visits flowers of
Hugelia virgata, Navarretia atractyloides.
Glossoperdita pelargoides Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 48. °.
Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 378, figs. 49, 50, 140. ¢.
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 379 (key).

Genus PERDITA Subgenus HESPEROPERDITA Timberlake

Perdita subg. Hesperoperdita Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 374.
Type-species: Perdita ruficauda Cockerell. Orig. desig.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 375 (key to included spp.).

pyrifera Cockerell. Calif. (Pleyto, Monterey County).
Perdita pyrifera Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 192. @.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 376, fig. 142. ¢.
trisignata ornata Timberlake. North. Calif. Pollen: Possibly an oligolege of Lotus, visits
flowers of L. glaber, L. nevadensis, L. strigosus var. hirtellus.
Perdita ruficauda ornata Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 377. 2, 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 127 (tax. status).
trisignata trisignata Cockerell. South. Calif.; Mexico (Baja California). Pollen: Possibly an
oligolege of Lotus including L. americanus, L. argophyllus, L. davidsonii, L. scoparius,
L. strigosus, but also visits other flowers including Cryptantha, Heteromeles.
Perdita trisignata Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 76. °.
Perdita ruficauda Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 44. 9°.

Taxonomy: Cockerell, 1922. U. S. Natl. Mus., Proc. 60: 17. 2, d (as ruficauda).
—Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 375, 376 (key, type). —Timberlake, 1954.
Calif. Univ. Pubs. Ent. 9: 375, 376, figs. 53, 54, 143 (key, tax. characters, geogr. and floral
records, as ruficauda). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 127 (synonymy).

Genus PERDITA Subgenus HETEROPERDITA Timberlake

Perdita subg. Heteroperdita Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 365.
Type-species: Perdita rhodogastra Timberlake. Orig. desig.

The species of this subgenus apparently collect pollen exclusively from flowers of the
boraginaceous genera Coldenia and Heliotropium.

Superfamily APOIDEA 1881

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 347, 366 (key to included spp.).
—Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 126 (modified key to included spp.).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 10-11 (key to females of included spp.).

arenaria Timberlake. Ariz. (Yuma), Nev. (Washoe County), south. Calif. (Colorado Desert).
Pollen: Apparently an oligolege of Coldenia including C. nuttallii, C. palmeri.
Perdita arenaria Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 368, figs. 39, 40, 135. d, 9.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 126 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 11, 13 (key, geogr. and floral records).
bellula Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an oligolege of
Coldenia including C. palmeri, C. plicata.
Perdita bellula Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 369, figs. 41, 42, 186. 6, @.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11 (key).
coldeniae Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an oligolege of
Coldenia including C. palmeri, but visits flowers of Pectis papposa presumably for

nectar.
Perdita coldeniae Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 368, figs. 37, 38, 134. d, °.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11 (key).
frontalis Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an oligolege of
Coldenia including C. plicata, but visits flowers of Eriogonum deserticola presumably
for nectar.
Perdita frontalis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 12. 2.
maculosa Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an oligolege of
Coldenia including C. plicata.
Perdita maculosa Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 378, fig. 514. 2, 6.
Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 126 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 11 (key).
optiva Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an oligolege of Coldenia
including C. palmeri.
Perdita optiva Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 371, figs. 47, 48, 139. d (?
misdet.).

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 13. ¢ (tax. status, as seafasciata).
rhodogastra Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently collects pollen
from the flowers of Coldenia palmeri and Heliotropium oculatum, but visits these and
other flowers for nectar including Pectis papposa, Pluchea sericea.
Perdita rhodogastra Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 367, figs. 35, 36, 133. °,
3.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11 (key).
scutellaris Timberlake. Southern Calif. (Colorado Desert). Pollen: Apparently an oligolege of
Coldenia including C. plicata, but visits the flowers of Hriogonwm deserticola

presumably for nectar.
Perdita scutellaris Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 89, figs. 829, 880, 893, text

fig. B. 2, d.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11 (key).
sexfasciata Timberlake. Ariz. (Yuma County), south. Calif. (Riverside and San Bernardino
Counties). Pollen: Apparently an oligolege of Coldenia including C. palmeri, but visits

these and other flowers for nectar including Cryptantha, Nama. _
Perdita sexfasciata Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 370, figs. 43, 44, 137. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11, 13. ¢ (key).
trifasciata Timberlake. Tex. (Dryden), N. Mex. (White Sands); Mexico (Coahuila).
Perdita trifasciata Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 969. ¢.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 366, 370, figs. 45, 46, 138. °.
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11, 14 (key).

1882 Hymenoptera in America North of Mexico

vesca Timberlake. Nevada (Wadsworth). Pollen: Possibly an oligolege of Coldenia, visits
flowers of C. nuttallir.
Perdita vesca Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11, figs. 1199, 1200, 1303. °, d.
wasbaueri Timberlake. South. Calif. (Colorado Desert). Pollen: Apparently an oligolege of
Coldenia plicata.
Perdita wasbaueri Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 126, figs. 656, 657, 730. °,
é.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 11 (key).
Genus PERDITA Subgenus HEXAPERDITA Timberlake

Perdita subg. Hexaperdita Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 416.
Type-species: Perdita ignota Cockerell. Orig. desig.

These bees are exclusively oligoleges of the family Compositae.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 348 (key). —Timberlake, 1956. Calif.
Univ. Pubs. Ent. 11: 247-252 (key to included spp.).
agasta Timberlake. Tex. (Hidalgo Co.).
Perdita agasta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 389. °.
alexi Timberlake. Tex. (College Station), Kans. (Barber County). Pollen: Unknown, but visits
flowers of Helianthus petiolaris, Heterotheca subaxillaris.
Perdita alexi Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 26. °.
asteris Cockerell. West. Tex., N. Mex., Ariz. Pollen: Unknown, but visits flowers of
Aplopappus, Aster canescens, Chrysopsis, Heterotheca subaxillaris, Nolina microcarpa,
Verbesina encelioides.
Perdita asteris Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 93. 2.
Perdita vespertilio Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 95. ¢.
Perdita crassiceps Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 97. ¢.
Perdita vagans Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 101. d.
Perdita mellina Cockerell, 1900. Entomologist 33: 62. 3.

Taxonomy: Cockerell, 1906. Entomologist 39: 148. 2 (as vespertilio). —Timberlake, 1956.
Calif. Univ. Pubs. Ent. 11: 249, 258 (key, tax. characters, geogr. and floral records).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 389-390 (tax. characters, geogr. and floral
records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 27 (synonymy, geogr. and floral
records).

bebbiae Timberlake. Calif. (San Diego County). Pollen: Unknown, but visits flowers of Bebbia
juncea.
Perdita bebbiae Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 255, figs. 179, 180, 292. 2, ¢.
bishoppi bishoppi Cockerell. N. C. to Fla., west to Tex. Pollen: Unknown, but visits flowers of
Heterotheca latifolia, H. subaxillaris, Isopappus divaricatus.
Perdita bishoppi Cockerell, 1906. Entomologist 39: 148. °, 6.

Taxonomy: Cockerell, 1922. Amer. Mus. Novitates 33: 7. —Timberlake, 1956. Calif. Univ.
Pubs. Ent. 11: 249, 250, 261, figs. 193, 194, 299 (key, tax. characters, geogr. and floral
records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 303-304, figs. 72-74, 79,
table 9 (redescription).

bishoppi planorum Timberlake. Kans., Tex. Pollen: Unknown, but visits flowers of
Heterotheca latifolia, H. subaxillaris, Isopappus divaricatus.
Perdita bishoppi planorum Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 261. 2, ¢.
blatchleyi Timberlake. Fla. (Dunedin). Pollen: Unknown, but visits flowers of Chrysopsis.
Perdita blatchleyi Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 201. 2.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 248, 258 (key). —Mitchell, 1960. N.
C. Agr. Expt. Sta. Tech. Bul. 141: 304, fig. 72, table 9 (redescription).
boltoniae boltoniae (Robertson). IIl., La., Miss. Pollen: Unknown, but visits flowers of

Boltonia.
Perditella boltoniae Robertson, 1902. Canad. Ent. 34: 321. °, 6.

Superfamily APOIDEA 1883

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 249, 252, 265 (key, tax. characters,
geogr. records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 304-305, fig. 73,
table 9 (redescription).

boltoniae chrysopsina Timberlake. N. J. to Fla. Pollen: Unknown, but visits flowers of
Chrysopsis mariana, Coreopsis, Erigeron, Gerardia, Haplopappus.
Perdita chrysopsina Timberlake, 1928. Hawaii. Ent. Soc., Proc. 7: 155. d, @.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 250, 252, 265, figs. 199, 200, 302 (key,
tax. characters). — Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 305-306, figs. 72, 74,
table 9 (redescription).

callicerata Cockerell. Tex. to south. Calif.; Mexico (Chihuahua, Durango). Pollen: Unknown,

most frequently has been collected at the flowers of Baileya multiradiata, but also
visits flowers of Baileya pleniradiata, Baccharis, Dithyrea wislizenii, Haplopappus.

Perdita callicerata Cockerell, 1896. N. Y. Ent. Soc., Jour. 4: 206. Nomen nudum, validated
by Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 153. male.

Perdita callicerata var. chrysoceras Cockerell, 1897. N. Mex. Agr. Expt. Sta., Bul. 24: 43.
Bio

Perdita callicerata var. lewcura Cockerell, 1909. Canad. Ent. 41: 152. °.

Perdita lutzi Cockerell, 1922. Amer. Mus. Novitates 33: 7. 2.

Taxonomy: Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 354. 9. —Timberlake, 1956. Calif.
Univ. Pubs. Ent. 11: 247, 248, 250, 255, figs. 181, 182, 293 (key, tax. characters, geogr. and
floral records).

cambarella cambarella Cockerell. Tex. Pollen: Unknown, but visits flowers of Heterotheca

subaxillaris, Isopappus divaricatus.
Perdita cambarella Cockerell, 1906. Entomologist 39: 150. 2, d.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 248, 251, 256-257 (key, tax.
characters, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 29
(color variation, geogr. records). 3

cambarella platyura Cockerell. Kans., Tex., N. Mex. Pollen: Unknown, but visits flowers of

Heterotheca latifolia, Isopappus divaricatus.
Perdita platyura Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 547. 9.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 248, 251, 257, figs. 183, 184, 294. ¢
(key, tax. characters, geogr. and floral records).
cara Timberlake. Tex. (Reeves County). Pollen: Unknown, but visits flowers of Prosopis.
Perdita cara Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 388, figs. 466, 467, 518. ¢.
compacta Timberlake. Ariz. (Graham County). Pollen: Unknown, but visits flowers of Pectis

papposa.
Perdita compacta Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 9. °.

fedorensis Cockerell. Tex. (Lee County).
Perdita fedorensis Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 280. °.
Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 249, 259 (key).
foveata brachycephala Timberlake. Tex. Pollen: Unknown, but visits flowers of

Aphanostephus skirrhobasis, Aster tanacetifolius, Coreopsis basalis.
Perdita foveata brachycephala Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 266. 9, d.
foveata foveata Timberlake. Fla. (Dade and Pasco Counties), Tex. (Montgomery County), Ark.
(Sevier and Scott Counties), Kans. (Reno County). Pollen: Unknown, but visits flowers

of Coreopsis tinctoria, Hymenopappus.
Perdita foveata foveata Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 265, figs. 201, 202,

303. 2, d.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 392 (geogr. and floral records).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 307, figs. 72-74, 79, table 9
(redescription).

foveata persimilis Timberlake. Tex. (Comanche County), N. Mex. (Chaves County).
Perdita foveata persimilis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 267. 2, d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 29 (geogr. record).

1884 Hymenoptera in America North of Mexico

georgica Timberlake. N. C. to Fla., west to Miss. Pollen: Unknown, but visits flowers of Aster,
Chrysopsis mariana, C. microcephala, Haplopappus, Heterotheca, Isopappus.
Perdita georgica Timberlake, 1928. Hawaii. Ent. Soc., Proc. 7: 158. 3, 9.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 249, 251, 257, figs. 185, 186, 295 (key,
tax. characters, geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 307-308, figs. 72-74, table 9 (redescription).

graenicheri Timberlake. Fla. (Miami). Pollen: Unknown, but visits flowers of Chrysopsis
tracy.
Perdita graenicheri Timberlake, 1947. Ent. Soe. Wash., Proce. 49: 82. 2, 6.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 249, 252, 264, figs. 197, 198, 301
(key). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 308-309, figs. 72, 73, table 9
(redescription).

heterothecae heterothecae Cockerell. Ariz., Utah (San Juan County), south. Calif. (Colorado
Desert), Mexico (Baja California). Pollen: Unknown, but visits flowers of Aster
tephrodes, Grindelia, Gutierrezia californica, G. sarothrae, Heliotropium curassavicum,
Heterotheca grandiflora, H. subaxillaris, Tamarix.
Perdita heterothecae Cockerell, 1900. Entomologist 33: 62. 2, 3.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 248, 251, 260-261, figs. 191, 192, 298
(key, tax. characters, geogr. and floral records). —Timberlake, 1971. Calif. Univ. Pubs. Ent.
66: 10 (geogr. and floral records).
heterothecae trizonata Timberlake. Ariz. (Willcox). Pollen: Unknown, but visits flowers of
Aster intricatus, Euphorbia.
Perdita heterothecae trizonata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 96. 2°, 6.
ignota basalis Timberlake. Tex. (Hidalgo County); Mexico (Tamaulipas).
Perdita ignota basalis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 391. 2, 6.
ignota crawfordi Cockerell. Colo., Nebr., Kans., Okla., Mo. Pollen: Unknown, but visits flowers
of Chrysopsis, Grindelia, Heterotheca subaxillaris, Isopappus divaricatus, Prionopsis
ciliata, Sideranthus rubiginosus.
Perdita crawfordi Cockerell, 1901. Entomologist 34: 190. 6, @.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 262-263, figs. 195, 196, 300 (tax.
characters, geogr. and floral records).
ignota ignota Cockerell. Tex., Colo., Ariz.; Mexico (Chihuahua and Zacatecas). Pollen:
Unknown, but visits flowers of Baccharis, Chrysopsis rutteri, Erigeron, Heterotheca,
Sideranthus, Solidago.
Perdita ignota Cockerell, 1896. Ent. Monthly Mag. 32: 220. 9.

Taxonomy: Cockerell, 1907. Colo. Univ., Studies 4: 247. 2, 6. —Cockerell, 1922. Amer. Mus.

Novitates 33: 7 (tax. characters). —Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 250, 252,

262 (key, tax. characters, geogr. and floral records). —Timberlake, 1971. Calif. Univ. Pubs.

Ent. 66: 10 (geogr. record).

ignota isopappi Timberlake. East. Tex., La. Pollen: Unknown, but visits flowers of

Aphanostephus skirrhobasis, Aster tanacetifolius, Chrysanthemum, Chrysopsis
nuttallii, Grindelia, Helenium tenuifolium, Heterotheca latifolia, H. pilosa, H.
subaxillaris, Isopappus divaricatus, Prionopsis ciliata, Xanthisma texanwm var.
drummondii.

Perdita isopappi Timberlake, 1928. Hawaii. Ent. Soc., Proc. 7: 152. d, °.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 249, 252, 263-264 (key, tax.
characters, geogr. and floral records).
nubila Timberlake. Fla., Ark., Kans. Pollen: Unknown, but visits flowers of Erigeron ramosus.
Perdita nubila Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 392. 2, 3.

Taxonomy: Timberlkake, 1968. Calif. Univ. Pubs. Ent. 49: 29 (geogr. records).

pratti Cockerell. Tex., Okla., Kans. Pollen: Unknown, but visits flowers of Helenium,
Helianthus cucumerifolius, H. petiolaris, Heterotheca latifolia, H. subaxillaris.
Perdita pratti Cockerell, 1906. Entomologist 39: 125. °.

Superfamily APOIDEA 1885

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 248, 250, 252-253, figs. 173, 174, 289.

3 (key, tax. characters, geogr. and floral records).
xanthisma Cockerell. Wyo., Nebr., Kans., Tex., N. Mex., Ariz. Pollen: Unknown, but visits

flowers of A plopappus divaricatus, A. subruginosus, Croton monanthogynus, Grindelia,
Gutierrezia, Helianthus annuus, Heterotheca pilosa, H. subaxillaris, Isopappus
divaricatus, Prionopsis ciliata, Sideranthus, Xanthisma texanum.

Perdita xanthismae Cockerell, 1905. Ent. News 16: 3381. °.

Perdita xanthismae sideranthi Cockerell, 1906. Entomologist 39: 178. °.

Perdita xanthisma Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 29. Emend.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 247, 249, 251, 258-259, figs. 187, 188,
296. 2, 3 (key, tax. characters, synonymy, geogr. and floral records). —Timberlake, 1958.
Calif. Univ. Pubs. Ent. 14: 390 (geogr. and floral records). —Timberlake, 1968. Calif. Univ.

Pubs. Ent. 49: 29-30 (geogr. and floral records).
Genus PERDITA Subgenus MACROTERA Smith

Macrotera Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 130.
Type-species: Macrotera bicolor Smith. Monotypic.
The species of this subgenus are apparently oligoleges of Opuntia.
Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 346, 352-353 (tax. characters, key to
included spp.). —Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 375-378 (key to included

spp.).
crassa Timberlake. Tex. (Sutton and Gillespie Counties). Pollen: Unknown, but possibly an

oligolege of Opuntia.
Perdita crassa Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 376, figs. 456, 457, 512. 9, d.

texana ablusa Timberlake. N. Mex., Tex. Pollen: Collects pollen from flowers of Opuntia

phaeacantha var. major.
Perdita texrana ablusa Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 376. °, d.

Biology: Barrows, Chabot, Michener and Snyder, 1976. Kans. Ent. Soc., Jour. 49: 275-279, 4
figs. (mating behavior, foraging, floral relationship, variation in head width, as texana).
texana texana (Cresson). Tex. (Bastrop, Gonzales, Lee and possibly Travis Counties). Pollen:

Unknown, but visits flowers of Opwntia.
Macrotera texana Cresson, 1878. Amer. Ent. Soc., Trans. 7: 70. °.
Macrotera megacephala Cresson, 1878. Amer. Ent. Soc., Trans. 7: 71. d.
Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 353, figs. 5, 6, 118 (tax. characters).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 375, 376 (tax. characters, status,

synonymy).
Genus PERDITA Subgenus MACROTERELLA Timberlake

Perdita subg. Macroterella Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 360.
Type-species: Perdita mortuaria Timberlake. Orig. desig.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 346, 361 (key to included spp.).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 7-8 (key to females of included spp.; key to

males of carinata, mortuaria, opacella).
carinata Timberlake. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers of

Eriogonum, Eschscholzia minutiflora, Ferrocactus acanthodes.
Perdita carinata Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 9, figs. 1197, 1198, 1302. 2,

é.
mellea Timberlake. Ariz., south. Calif. (Colorado Desert); Mexico (Baja California and Sonora)
Pollen: Apparently an oligolege of Ewphorbia including E. polycarpa, E. p. var. hirtella.

Perdita mellea Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 364, figs. 38, 34, 182. d, 9.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 125 (geogr. and floral records).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 7 (key).

1886 Hymenoptera in America North of Mexico

mortuaria Timberlake. South. Calif., Ariz.; deserts. Pollen: Unknown, but visits flowers of
Echinocereus engelmanni, Eriogonum inflatum, Eschscholzia californica, E.
minutiflora var. darwinensis, Eucnide urens, Larrea tridentata, Prosopis juliflora.
Perdita mortuaria Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 362, figs. 27, 28, 129. d, 9.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 324 (tax. characters, geogr. record,
variation). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 126 (geogr. and floral record,
variation). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 7, 8, 10 (key, geogr. and floral
records).

nigrella Timberlake. South. Calif.; deserts. Pollen: Unknown, but visits flowers of Phacelia
including P. crenulata.
Perdita nigrella Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 362, figs. 29, 30, 1380. d, °.

Taxonomy: Timberlake, 1968 Calif. Univ. Pubs. Ent. 49: 7 (key).
opacella Timberlake. Ariz. (Grand Canyon), Utah (Moab). Pollen: Unknown, but visits flowers
of Cleome lutea, Stanleya.
Perdita opacella Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 324. @.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 7, 8, figs. 3, 1195, 1196, 1801, 1353.
3.
solitaria Cockerell. N. Mex. (Organ Mts.), Ariz. (Santa Catalina Mts.).
Perdita solitaria Cockerell, 1896. N. Y. Ent. Soc., Jour. 4: 206. Nomen nudum.
Perdita solitaria Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 152. °.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 361, 365 (key, geogr. records).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 7 (key).
tristella Timberlake. South. Calif. (Riverside County). Pollen: Unknown, but visits flowers of
Calochortus splendens, Eriogonum fasciculatum, Hugelia virgata, Sphaeralcea.
Perdita tristella Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 363, figs. 31, 32, 131. d, 2.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 7 (key).

Genus PERDITA Subgenus MACROTEROPSIS Ashmead

Macroteropsis Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 85.
Type-species: Perdita latior Cockerell. Monotypic and orig. desig.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 346, 356-357 (redescription, key to
included spp.). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 87 (key to males of included
spp.).

arcuata arcuata Fox. Calif. (San Benito County); Mexico (Baja California). Pollen: Apparently
an oligolege of Sphaeralcea.
Perdita arcuata Fox, 1893. Calif. Acad. Sci., Proc. (2) 4: 18. 6.

Taxonomy: Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 54 (tax. characters).
—Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 357, 358 (key, tax. characters, tax. status).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 87 (key). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 4 (geogr. and floral record).

arcuata dinognatha Cockerell. Ariz. (Tempe), Nev. (Charleston Mts.), south. Calif.; deserts.
Pollen: Apparently collects pollen only from the flowers of Sphaeralcea ambigua, but
visits other flowers for nectar including Encelia frutescens, Eriogonum fasciculatum
var. polifolium.
Perdita dinognatha Cockerell, 1922. U. S. Natl. Mus., Proc. 60 (18): 19. ¢.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 968 (tax. status, geogr. and floral
records). —Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 357, 359, figs. 21, 22, 126. 2 (key,
geogr. and floral records).

echinocacti Timberlake. Ariz.; Mexico (Sinaloa and Sonora). Pollen: Unknown, but visits
flowers of Antigonon leptopus, Jacquemontia, Kallstroemia grandiflora.
Perdita echinocacti Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 360, figs. 25, 26, 128. d, @.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 125 (geogr. and floral records).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 87 (key).

Superfamily APOIDEA 1887

haplura Cockerell. Tex. (Terrell and Val Verde Counties), N. Mex. (Lincoln County). Pollen:
Unknown, but visits flowers of Sphaeralcea angustifolia var. cuspidata.
Perdita haplura Cockerell, 1922. U.S. Natl. Mus., Proc. 60 (18): 19. 2.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 124 (type). —Timberlake, 1962.
Calif. Univ. Pubs. Ent. 28: 87-89, text fig. A. ¢. —Timberlake, 1968. Calif. Univ. Pubs. Ent.
49: 4 (geogr. and floral records).
latior Cockerell. N. Mex., Ariz., Nev. (Charleston Mts.). Pollen: Apparently an oligolege of
Sphaeralcea including S. angustifolia, but also visits flowers of Sida hederacea

presumably for nectar.
Perdita latior Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 53. d, 9.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 357, 359, figs. 28, 24, 127.
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 87 (key).
magniceps Timberlake. N. Mex. (Luna County). Pollen: Unknown, but visits flowers of
Sphaeralcea.
Perdita magniceps Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 124, figs. 654, 655, 729. ¢.
portalis Timberlake. N. Mex. (Deming), Ariz. (Cochise and Coconino Counties); Mexico
(Zacatecas). Pollen: Apparently an oligolege of Sphaeralcea.
Perdita portalis Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 357, figs. 19, 20, 125. 6, 9.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 323. ¢ (variation). —Timberlake,
1960. Calif. Univ. Pubs. Ent. 17: 125 (geogr. and floral record). —Timberlake, 1962. Calif.
Univ. Pubs. Ent. 28: 87. ¢ (key). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 4 (geogr.
record).

robertsi Timberlake. Tex. (Val Verde County). Pollen: Possibly collects pollen from the flowers
of Opuntia.
Perdita robertsi Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 6, fig. 2. °.

Genus PERDITA Subgenus PENTAPERDITA Cockerell and Porter

Perdita subg. Pentaperdita Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 415.
Type-species: Perdita albovittata Cockerell. Monotypic.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 347, 404-406 (redescription, key to
included spp.). —Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 329 (modified key to
included spp.). —Timberlake, 1958. Calif. Uniy. Pubs. Ent. 384-385 (key to spp. allied to
melanochlora). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 22-24 (key to females of
included spp.).

albovittata Cockerell. N. Mex., Ariz., south. Calif. (Colorado and Mojave Deserts); Mexico
(Chihuahua, Coahuila, Durango). Pollen: Unknown, but visits flowers of A plopappus
spinulosus, Baileya multiradiata, B. pleniradiata, Cevallia sinuata, Coldenia greggii,
Eriogonum inflatum, Gutierrezia californica, G. lucida, Haplopappus acradenius, H.
gracilis, Pectis papposa, Physalis, Stephanomeria, Verbesina encelioides.
Perdita albovittata Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 15. 2.
Perdita laticeps Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 98. d.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 405, 406, 411, figs. 105, 106, 168 (key,
geogr. and floral records, possible synonymy). —Timberlake, 1968. Calif. Univ. Pubs. Ent.
49: 23, 24 (synonymy, geogr. and floral record).
amoena Timberlake. Ariz. (Cochise and Santa Cruz Counties), Utah (Washington County);
Mexico (Sonora).
Perdita amoena Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 330, figs. 287, 288, 336. d.
Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 385, 386. ° (key). —Timberlake,
1960. Calif. Univ. Pubs. Ent. 17: 132 (geogr. record). —Timberlake, 1968. Calif. Univ. Pubs.
Ent. 49: 23, 24 (key, geographic records).
annexa Timberlake. N. Mex. (Pecos and Santa Fe). Pollen: Unknown, but visits flowers of

Hymenoxys richardson.
Perdita annexa Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 133. d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 28, 24. 2 (key).

1888 | Hymenoptera in America North of Mexico

bradleyana Timberlake. Tex. (Eagle Pass and El Paso); Mexico (Zacatecas).
Perdita bradleyana Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 409, figs. 99, 100, 165. 6.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 329. 2. —Timberlake, 1958. Calif.
Univ. Pubs. Ent. 14: 385 (key). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 23, 25 (key,
geogr. record). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 7 (geogr. record).

chrysophila chrysophila Cockerell. Tex. (Culberson County). N. Mex. (Hollywood, Organ and
Pecos), Ariz. (Safford); Mexico (Hidalgo, San Luis Potosi and Zacatecas). Pollen:
Unknown, but visits flowers of Flouwrensia, Hymenoxys, Picradenia floribunda,
Verbesina encelioides, Zaluzania globosa. Another subspecies, Perdita chrysophila
quadricincta Timberlake, occurs in Mexico (Nuevo Leon).
Perdita chrysophila Cockerell, 1896. N. Y. Ent. Soc., Jour. 4: 206. d.
Perdita nigrifacies Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 406 °.

Taxonomy: Cockerell, 1904. Entomologist 37: 6. d. —Timberlake, 1954. Calif. Univ. Pubs.
Ent. 9: 406, figs. 93, 94, 162 (key, type). Timberlake, 1958. Calif. Univ. Pubs. Ent. 14:
386-387 (tax. characters, as nigrifacies). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17:
132-133, figs. 660, 661, 732. d (redescription). —Timberlake, 1962. Calif. Univ. Pubs. Ent.
28: 95-96 (tax. characters, as nigrifacies). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49:
23, 25 (key, synonymy, geogr. and floral records).

coahuilensis Timberlake. Tex. (E] Paso); Mexico (Coahuila).
Perdita coahuilensis Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 409, figs. 101, 102, 166. °,
3.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 329, 330 (key, geogr. record).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 385 (key). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 24 (key).

idahoensis Timberlake. Idaho (Cassia County), Calif. (Inyo County). Pollen: Unknown, but
visits flowers of Chrysothamnus, Gutierrezia microcephala.
Perdita idahoensis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 385, figs. 464, 465, 517. 6,
OF

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 23 (key). —Timberlake, 1971. Calif.
Univ. Pubs. Ent. 66: 9 (geogr. and floral record).
mandibularis Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Unknown, but
visits flowers of Bebbia juncea, Chaenactis carphoclinia, Dalea, Geraea canescens.
Perdita mandibularis Timberlake, 1954. calif. Univ. Pubs. Ent. 9: 411, figs. 107, 108, 169.
ORGr
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 23 (key).
melanochlora Cockerell. Ariz. (Santa Rosa Valley and Tucson), south. Calif. (Colorado and
Mojave Deserts), Nev. (Arden and Charleston Mts.). Pollen: Unknown, but visits flowers
of Baileya multiradiata, Bebbia juncea, Chrysothamnus, Encelia farinosa, Sphaeralcea,

Viguiera deltoides, V. nevadensis.
Perdita melanochlora Cockerell, 1922. Amer. Mus. Novitates 33: 7. 2.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 405, 406, 408, figs. 97, 98, 164. 2, ¢.
—Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 329 (key). —Timberlake, 1958. Calif. Univ.
Pubs. Ent. 14: 385 (key). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 23 (key).
—Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 9 (geogr. records).

nigroviridis Timberlake. Tex. Pollen: Unknown, but visits flowers of Gaillardia pulchella,
Helenium, Helianthus annuus.
Perdita nigroviridis Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 407, figs. 95, 96, 163. 2, ¢.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 386 (geogr. and floral records).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 23, 26 (key, geogr. and floral record).

Genus PERDITA Subgenus PERDITA Smith

Perdita Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 128.
Type-species: Perdita halictoides Smith. Monotypic.
Neoperdita Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 85.

Superfamily APOIDEA 1889

Type-species: Perdita zebrata Cresson. Monotypic and orig. desig.

Perdita subg. Geoperdita Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 415.
Type-species: Perdita chamaesarachae Cockerell. Monotypic.

Perdita subg. Tetraperdita Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 415.
Type-species: Perdita sexmaculata Cockerell. Monotypic and orig. desig.

Zaperdita Robertson, 1918. Ent. News 29: 91.
Type-species: Perdita maura Cockerell. Monotypic and orig. desig.

This is the largest subgenus of Perdita. Six groups of species are currently recognized and,
although some of these groups have been divided into subgroups in the existing classification, no
attempt has been made to reflect these subsidiary units in this catalog. The six recognized spe-
cies groups are arranged in accordance with the current classification of this subgenus.

Revision: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 303-365 (zonalis group).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 365-374 (halictoides group). —Timberlake,
1960. Calif. Univ. Pubs. Ent. 17: 1-121 (octomaculata group). —Timberlake, 1960. Calif.
Univ. Pubs. Ent. 17: 135 (halictoides group). —Timberlake, 1962. Calif. Univ. Pubs. Ent.
28: 1-86 (ventralis group). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1, 97-101 (zonalis
group). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1, 101-102 (halictoides group).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1, 102-106 (octomaculata group).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 106-107 (valida group). —Timberlake, 1964.
Calif. Univ. Pubs. Ent. 28: 1, 125-362 (sphaeralceae group). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 40-57 (zonalis group). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49:
57-62 (halictoides group). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 62-95
(octomaculata group). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 95-112 (ventralis
group). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 112-161 (sphaeralceae group).
—Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 11-56 (suppl. studies on the species of the
subgenus Perdita).

SPECIES GROUP ZONALIS

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 303-315 (tax. characters, key to
included spp.). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1, 97-101 (tax. characters,
modified key to included spp.). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 40-43 (suppl.
key to included spp.). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 11-13 (suppl. key to
included spp.).

accepta Timberlake. Oreg. (Crook County).
Perdita accepta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 340. @.
adjuncta Timberlake. Nev. (Austin), Calif. (Modoc County), Oreg. (Lake County), Idaho
(Bingham, Cassia and Elmore Counties). Pollen: Unknown, but visits flowers of

Chrysothamnus.
Perdita adjuncta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 327, figs. 390, 391, 479. 9,
3.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 97 (key).
aemula aemula Timberlake. Oreg., Utah. Pollen: Unknown, but visits flowers of
Chrysothamnus, Solidago.
Perdita aemula Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 330, figs. 396, 397, 482. d, 9.
Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 98 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 42 (key).
aemula quadrifasciata Timberlake. Calif. (Mono County). Pollen: Unknown, but visits flowers

of Chrysothamnus.
Perdita aemula quadrifasciata Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 43. 2, ¢.

affecta Timberlake. Ariz. (Tuba City). Pollen: Unknown, but visits flowers of Artemisia
filifolia.
Perdita affecta Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 14. °.
albopicta Timberlake. Utah (Millard County). Pollen: Unknown, but visits flowers of

Chrysothamnus.
Perdita albopicta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 341. °.

1890 Hymenoptera in America North of Mexico

ampla Timberlake. N. Mex. (McKinley and San Juan Counties). Pollen: Unknown, but visits
flowers of Chrysothamnus nauseosus, Senecio.
Perdita ampla Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 14. °.
baccharidis Cockerell. Ariz. (Maricopa and Pima Counties), Calif. (Riverside and Imperial
Counties). Pollen: Apparently an oligolege of Baccharis including B. emoryi, B.
sarothroides.
Perdita baccharidis Cockerell, 1900. Entomologist 33: 61. °.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 309, 313, 352-353, figs. 428, 429, 498.
2, 6 (redescription). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 44 (geogr. and floral
records).

basinicola Timberlake. Nev. (Churchill County), Calif. (Inyo County).
Perdita basinicola Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 44, figs. 1213, 1214, 1310.
Ohne
chrysothamni Timberlake. Calif. (Mojave Desert). Pollen: Apparently an oligolege of
Chrysothamnus nauseosus.
Perdita chrysothamni Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 319, figs. 374, 375, 473.
Omron
ciliata Timberlake. North. Calif., Oreg., Idaho. Pollen: Unknown, but visits flowers of Aster,
Chrysothamnus nauseosus, Eriogonum, Gutierrezia, Haplopappus, Heterotheca
grandiflora, Lessingia glandulifera, Melilotus alba, Senecio douglasii, Solidago
occidentalis. Predator: Philanthus pacificus arizonae Dunning.
Perdita interserta ciliata Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 334, figs. 404, 405,
486. 2, 3.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 43, 45 (key, tax. status, geogr.
records, variation).
confinis Timberlake. Calif. (Mono County).
Perdita confinis Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 15. 9.
crassihirta Timberlake. Wash. (Chelan County).
Perdita crassihirta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 46. °.
craterognatha Timberlake. Calif. (Mono County).
Perdita craterognatha Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 46, figs. 1215, 1216,
1Stils Gad
depressa Timberlake. Utah (Monument Valley), Ariz. (Havasu Canyon). Pollen: Unknown, but
visits flowers of Cleome jonesii.
Perdita depressa Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 48, figs. 7, 1217, 1218, 1312.
OnaGr
dicksoni Timberlake. Calif. (Inyo County). Pollen: Unknown, but visits flowers of

Chrysothamnus albidus.
Perdita dicksoni Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 328, figs. 392, 393, 480. 2, d.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 97 (key).
dubia dubia Cockerell. Colo. (Garfield and Pitkin Counties), Ariz. (Yavapai County). Pollen:
Unknown, but visits flowers of Gutierrezia microcephala, G. sarothrae.
Perdita dubia Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 75. 6.
Perdita fraterna Timberlake, 1929. N. Y. Ent. Soc., Jour. 39: 123. 3.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 306, 312, 329, figs. 394, 395, 481. 9,
6 (redescription, key, as fraterna). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 43, 50
(key, tax. status). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 16-17 (tax. status,
synonymy).
dubia parilis Timberlake. Ariz. (Coconino and Mohave Counties), Utah, Nev. (Clark County),
Idaho (Cassia and Franklin Counties), Wyo. (Franklin County). Pollen: Unknown, but
visits flowers of Aplopappus, Chrysothamnus, Eriogonum, Gutierrezia.
Perdita parilis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 342, figs. 414, 415, 491. 2, 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 42, 43, 52 (key, geogr. and floral
records). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 17-18 (tax. status).

Superfamily APOIDEA 1891

ericameriae Timberlake. South. Calif. (Los Angeles and Riverside Counties). Pollen: Unknown,
but visits flowers of Aplopappus palmeri var. pachylepis, Gutierrezia californica,

Lepidospartum squamatum.
Perdita ericameriae Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 317, figs. 372, 373, 472.

Qe
festiva Timberlake. Utah (Emery County).
Perdita festiva Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 321. 2.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 98 (key).

foleyi Timberlake. Calif. (Kern and Tulare Counties). Pollen: Unknown, but visits flowers of

Chrysothamnus nauseosus.
Perdita foleyi Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 347, figs. 418, 419, 493. 9, d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 50 (geogr. and floral record).
haigi Timberlake. Nev. (Churchill County).
Perdita haigi Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 100, figs. 833, 834, 895. 2, d.

impunctifrons Timberlake. Calif. (Inyo and San Bernardino Counties).
Perdita impunctifrons Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 355. 2.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 18 (geogr. record).
interserta Cockerell. South. Calif. (cismontane and Mojave Desert). Pollen: Unknown, but
visits flowers of Aplopappus palmeri var. pachylepis, A. veneta var. vernonioides,
Baccharis emoryi, Chrysothamnus nauseosus, Eschscholzia californica, Lepidospartum
squamatum, Polygonum lopathifolium, Rhus diversiloba, Satureja occidentalis,
Solidago occidentalis.
Perdita interserta Cockerell, 1922. U. S. Natl. Mus., Proc. 60 (18): 20. °.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 307, 311, 312, 333-334, figs. 402, 403,
485. 6, 2 (key, tax. characters, tax. status, geogr. and floral records). —Timberlake, 1968.
Calif. Univ. Pubs. Ent. 49: 45 (tax. status).

irregularis Timberlake. N. Mex. (McKinley County). Pollen: Unknown, but visits flowers of

Chrysothamnus nauseosus.
Perdita irregularis Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 18, figs. 1867, 1368, 1402.

Onn
isocomae Timberlake. Calif. (Riverside). Pollen: Unknown, but visits flowers of Aplopappus
venetus var. vernonioides, Baccharis emoryi, Polygonum lopanthifolium, Solidago

occidentalis.
Perdita isocomae Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 335, figs. 408, 409, 488. d,

OF
lepidosparti lepidosparti Timberlake. Calif. (Mojave Desert and Lassen County), Nev.
(Washoe County), Idaho (Bingham, Cassia, Fremont and Owyhee Counties). Pollen:
Unknown, but visits flowers of Chrysothamnus, Eriogonum heermannii, Gutierrezia
lucida, Lepidospartum squamatum, Tetradymia comosa.
Perdita lepidosparti lepidosparti Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 325, figs.
386, 387, 478. 5, °.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 98 (key). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 50 (geogr. and floral records).
lepidosparti novella Timberlake. South. Calif. (San Bernardino and San Gabriel Mts.).
Perdita lepidosparti novella Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 326, figs. 388,
389. 2, d.
Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 98 (key).

lompocensis Timberlake. Calif. (Lompoc).
Perdita lompocensis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 337, figs. 410, 411, 489.

Gras

melanderi Timberlake. Calif. (Morro Bay). é
Perdita melanderi Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 50, fig. 8. 2.

munda Timberlake. Utah, Ariz. (Apache County).
Perdita munda Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 340. °.

1892 Hymenoptera in America North of Mexico

nigrocincta Timberlake. Calif. (Mono County).
Perdita nigrocincta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 351. °.

obispoensis Timberlake. Calif. (San Luis Obispo County).
Perdita obispoensis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 335, figs. 406, 407, 487. d.
oregonensis expleta Timberlake. Calif. (Sierra Nevada Mts.). Pollen: Unknown, but visits
flowers of Chrysothamnus.
Perdita oregonensis expleta Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 99. °.
oregonensis oregonensis Timberlake. Wash. and Idaho, south to Calif. and Nev. Pollen:
Unknown, but visits flowers of Chrysothamnus viscidiflorus, Gutierrezia.
Perdita oregonensis Timberlake, 1929. Pan-Pacifie Ent. 6: 52. 3.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 306, 311, 320-321, figs. 378, 379, 475.
2, S (key, redescription, geogr. and floral records). —Timberlake, 1962. Calif. Univ. Pubs.
Ent. 28: 98 (key, tax. status). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 41, 52 (geogr.
and floral records).

pallidiventris Timberlake. Calif. (Fresno County). Pollen: Unknown, but visits flowers of
Eriogonum fasciculatum.
Perdita pallidiventris Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 99. °.
placida Timberlake. Calif. (Inyo County).
Perdita placida Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 339. 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 42 (key).
polita Timberlake. Calif. (Kings, Tulare and Stanislaus Counties). Pollen: Unknown, but visits
flowers of Hemizonia pungens.
Perdita polita Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 332, figs. 400, 401, 484. 2, do.
primula Timberlake. Calif. (Indio). Pollen: Unknown, but visits flowers of Melilotus alba.
Perdita primula Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 347, figs. 420, 421, 494. ¢.
proxima Timberlake. Calif. (Mohave Desert). Pollen: Unknown, but visits flowers of
Chrysothamnus nauseosis, C. n. consimilis, Cleomella obtusifolia, Haplopappus

acradenius, Solidago occidentalis.
Perdita proxima Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 322, figs. 380, 381, 476. °, 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 52 (geogr. and floral records).
punctifrons Timberlake. Calif. (Los Angeles and San Bernardino Counties). Pollen: Unknown,

but visits flowers of Lessingia germanorum var. glandulifera.
Perdita punctifrons Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 350, figs. 426, 427, 497. °,
3

repens Timberlake. Calif. (Mt. Hamilton). Pollen: Unknown, but visits flowers of Haplopappus
arborescens.
Perdita repens Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 19, figs. 1369, 1870, 1403. d.
rivalis Timberlake. Calif. (San Bernardino, San Gabriel and Sierra Nevada Mts.), Nev.
(Daggett Pass). Pollen: Unknown, but visits flowers of Aster adscendens, A. canescens,
Erigeron divergens, E. foliolosus var. stenophyllus, Gayophytum heterozygos, G.

nuttallii.
Perdita rivalis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 349, figs. 424, 425, 496. °, ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 43, 52 (key, geogr. and floral
records).
scotti Timberlake. Calif. (Inyo, Los Angeles and Kern Counties). Pollen: Unknown, but visits

flowers of Chrysothamnus nauseosus.
Perdita scotti Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 338, figs. 412, 413, 490. @, ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 53 (geogr. and floral records).

similis pascoensis Timberlake. Wash. (Pasco).
Perdita similis pascoensis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 324, figs. 384, 385.
Cae
similis similis Timberlake. Oreg., Idaho, and Wyo., south to N. Mex., Ariz. and south. Calif.
(Inyo, Riverside and Imperial Counties). Pollen: Unknown, but visits flowers of

Superfamily APOIDEA 1893

Chrysothamnus nauseosus, C. viscidiflorus, Cleome serrulata, Erigeron, Gutierrezia
sarothrae, Haplopappus heterophyllus, Prosopis juliflora.
Perdita similis similis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 323, figs. 382, 383, 477.

,

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 22 (collection site). —Timberlake,
1968. Calif. Univ. Pubs. Ent. 49: 41, 53 (key, geogr. records, floral preferences).
ocia Timberlake. Calif. (Inyo County).
Perdita socia Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 42, 43, 53-54, plate figs. 1219,
1220, 13138. 2, d.
itottleri Cockerell. Sask., south to N. Mex. west to Idaho, Nev. and Calif. (Mono County).
Pollen: Unknown, but visits flowers of Aster, Chrysothamnus graveolens, C. g. var.
glabrata, C. nauseosus, Gutierrezia sarothrae, Solidago missouriensis.
Perdita stottleri Cockerell, 1896. N. Y. Ent. Soc., Jour. 4: 205. 9.
Perdita stottleri flavida Swenk and Cockerell, 1907. Ent. News 18: 58. °, 3.

Taxonomy: Cockerell, 1904. Entomologist 37: 6. ¢. —Timberlake, 1958. Calif. Univ. Pubs.
Ent. 14: 304, 310, 315 (tax. characters, key, geogr. and floral records). —Timberlake, 1958.
Calif. Univ. Pubs. Ent. 14: 304, 306, 310, 315-316, figs. 368, 369, 470 (tax. characters, key,
geogr. and floral records, as stottleri flavida). —Timberlake, 1962. Calif. Univ. Pubs. Ent.
28: 98 (key, as stottleri flavida). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 54
(synonymy, geogr. and floral records).

subvestita Timberlake. N. Mex. (Citron County), Ariz. (Coconino County), Utah (Kane County).
Pollen: Unknown, but visits flowers of Aplopappus gracilis, Chrysothamnus
viscidiflorus, Gutierrezia microcephala.
Perdita subvestita Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 55, figs. 1221, 1222, 1314.
OaTOr
swezeyi Timberlake. Calif., Nev. (Ormsby County). Pollen: Unknown, but visits flowers
Chrysothamnus, Erigeron ramosus.
Perdita swezeyi Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 348, figs. 422, 423, 495. 9, 3.
taeniata Timberlake. N. Mex. (McKinley and San Juan Counties). Pollen: Unknown, but visits
flowers of Chrysothamnus nauseosus, Senecio longi.
Perdita taeniata Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 20, figs. 1871, 1872, 1404. ?,
é.
toschiae Timberlake. Nev. (Douglas County).
Perdita toschiae Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 56. 9.
townsendi Cockerell. N. Mex. (Otero and Rio Arriba Counties), Ariz. (Apache County). Pollen:
Unknown, but visits flowers of Aster, Chrysothamnus, Haplopappus heterophyllus.
Perdita townsendi Cockerell, 1896. N. Y. Ent. Soc., Jour. 4: 204. 2, 6.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 305, 310, 320, figs. 376, 377, 474 (tax.
characters, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 56
(geogr. and floral records).

vestita Timberlake. Utah (Grand and Kane Counties), Ariz. (Coconino County). Pollen:
Unknown, but visits flowers of Chrysothamnus, Gutierrezia.
Perdita vestita Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 316, figs. 370, 371, 471. 2, 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 41, 42, 57 (key, geogr. and floral
records).
zonalis aequalis Timberlake. Oreg., northern Calif., ?Nev., ?7Mont. Pollen: Unknown, but visits
flowers of Chrysothamnus including C. nawseosus speciosus, Solidago.
Perdita zonalis aequalis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 344. 2, d.
zonalis bernardina Timberlake. Calif. (Bear Valley and Santa Ana Canyon in San Bernardino
Mts.). Pollen: Unknown, but visits flowers of Aster canescens, Chrysothamnus
nauseosus, C. viridulus, C. viscidiflorus, Eriogonum molestum var. davidsonii, E.
wrightii var. subscaposum, Gutierrezia californica, G. lucida, Solidago californica.
Perdita zonalis bernardina Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 345, figs. 416, 417,
BOZO RIGk

1894. Hymenoptera in America North of Mexico

zonalis monticola Timberlake. Calif. (Sierra, Shasta, Nevada, Plumas and Placer Counties).
Pollen: Unknown, but visits flowers of Chrysothamnus, Solidago.
Perdita zonalis monticola Timberlake, 1958 Calif. Univ. Pubs. Ent. 14: 345. 9, 6.
zonalis pallescens Timberlake. Calif. Pollen: Unknown, but visits flowers of Gutierrezia
lucida, Haplopappus acradenius.
Perdita zonalis pallescens Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 344. 2, d.
zonalis zonalis Cresson. Calif. (Riverside and Kern Counties). Pollen: Unknown, but visits
flowers of Eriogonum, Gutierrezia californica, Lepidospartum squamatum.
Perdita zonalis Cresson, 1879. Amer. Ent. Soc. Trans. 7: 202. 9.

Taxonomy: Timberlake, 1929. Pan-Pacific Ent. 6: 49. 2, 6. —Timberlake, 1958. Calif. Univ.
Pubs. Ent. 14: 308, 313, 343-344 (key, redescription, geogr. and floral records).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 41, 43 (key).

SPECIES GROUP HALICTOIDES

The species of this group collect pollen from flowers of the solanaceous genera Physalis and
Chamaesaracha, but visit these and other flowers for nectar.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 365-367 (tax. characters, key to
included spp.). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1 (key). —Timberlake, 1968.
Calif. Univ. Pubs. Ent. 49: 57-59 (key to included spp.).

beameri Timberlake. Tex. Pollen: Collects pollen from the flowers of Chamaesaracha including
C. conioides, but visits other flowers for nectar including Astragalus, Gilia acerosa,

Physalis lobata.
Perdita beameri Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 276, figs. 56, 1041, 1042,

1164. 2, 6.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 57, 59 (key, geogr. and floral
records).

chamaesarachae Cockerell. N. Mex. (Albuquerque, High Rolls, Las Vegas, Roswell and Santa
Fe), Ariz. (Cochise and Yavapai Counties); Mexico (Durango). Pollen: Apparently
collects pollen from the flowers of Chamaesaracha coronopus, but visits other flowers
including Physalis.
Perdita chamaesarachae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 65. 6, °.
Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 367, 371, figs. 450, 451, 509 (key,
redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 58,
59 (key).
fidissima Timberlake. Tex. (Kenedy County). Pollen: Unknown, but visits flowers of
Parkinsonia.
Perdita fidissima Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 59. 9.
halictoides Smith. Wis., Mich. and Ind., west to N. Dak., Colo., and Tex.; Fla. Pollen: Unknown,
but visits flowers of Euphorbia nuttallii, Gaillardia, Monarda punctata var.
occidentalis, Physalis heterophylla, P. lanceolata, Solidago.
Perdita halictoides Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 128. 9.
Perdita seemaculata var. punctata Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 71. 9.
Perdita maura Cockerell, 1901. Entomologist 34: 191. dg, °.
Perdita bisignata Cockerell, 1922. Amer. Mus. Novitates 33: 11. 9.

Taxonomy: Cockerell, 1904. Canad. Ent. 36: 303. —Timberlake, 1928. Hawaii. Ent. Soc., Proc.
7: 155. —Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 268 (tax. characters, as maura).
—Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 366, 367, 370-371, figs. 448, 449, 508
(redescription, key, geogr. and floral records, as maura). —Timberlake, 1958. Calif. Univ.
Pubs. Ent. 14: 366, 371. 2 (key, redescription). —Timberlake, 1960. Calif. Univ. Pubs. Ent.
17: 135 (synonymy, geogr. records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
318-319, figs. 76-79, table 9 (redescription, synonymy). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 57, 58 (key).

Superfamily APOIDEA 1895

lenis Timberlake. Tex., N. Mex., Ariz., Calif. (Indio). Pollen: Unknown, but visits flowers of
Baileya multiradiata, Chamaesaracha conioides, Physalis heterophylla, Verbesina

encelioides.
Perdita lenis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 372, figs. 452, 453, 510. 2, d.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 19, figs. 47-49 (larva). —Timberlake,
1968. Calif. Univ. Pubs. Ent. 49: 58, 59 (key). —Timberlake, 1971. Calif. Univ. Pubs. Ent.
66: 22 (geogr. and floral records).
levigata Timberlake. Colo. (Fremont County). Pollen: Unknown, but visits flowers of Physalis.
Perdita levigata Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 60, figs. 1225, 1226, 1316. ¢.
missionis Timberlake. Tex.; Mexico (Tamaulipas, San Luis Potosi). Pollen: Unknown, but visits

flowers of Chamaesaracha conioides, Lesquerella, Physalis lobata.
Perdita missionis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 369, figs. 446, 447, 507. d,°.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 58 (key).
physalidis Timberlake. Calif. (Riverside County), Ariz. (Continental). Pollen: Unknown, but

visits flowers of Physalis, including P. crassifolia.
Perdita physalidis Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 374, figs. 454, 455, 511. °,¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 58, 59, 61 (key, geogr. and floral
records).
rozeni Timberlake. Calif. (Colorado Desert); Mexico (Baja California and Sonora). Pollen:
Unknown, but visits flowers of Boerhaavia, Dalea mollis.
Perdita rozeni Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 61, figs. 1227, 1228, 1317. 2, d.
sexmaculata sexmaculata Cockerell. Ariz., N. Mex., Tex., Kans., Colo.; Mexico (Chihuahua,
Durango, Hidalgo). Pollen: Unknown, but visits flowers of Astragalus, Chamaesaracha
conioides, C. coronopus, Phacelia piperi, Physalis lobata, Prosopis, Scutellaria,
Sphaeralcea albicaulis, Thelesperma gracile. Another subspecies, Perdita sexmaculata
octonaria Timberlake, occurs in Baja California, Mexico.
Perdita sermaculata Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 12. 2.
Perdita anograe Cockerell, 1902. Amer. Nat. 36: 812. d.

Taxonomy: Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 416. 9, 6.
—Timberlake, 1929. N. Y. Ent. Soc., Jour. 37: 118. ¢. —Timberlake, 1951. In Muesebeck,
Krombein and Townes, U. S. Dept. Agr., Agr. Monog. 2: 1095 (synonymy). —Timberlake,
1958. Calif. Univ. Pubs. Ent. 14: 366, 367-369, figs. 444, 445, 506 (key, redescription).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 101 (tax. status). —Rozen, 1966. Amer.
Mus. Novitates 2259: 20-21, fig. 51 (larva, as sexmaculata?). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 58 (key).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 38-40, figs. 16, 18, table 1 (nest
architecture, life history).

SPECIES GROUP OCTOMACULATA

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 1-22 (tax. characters, key to
included spp.). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 62-67 (suppl. key to included
spp.). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 22-25 (suppl. key to included spp.).

abdominalis Timberlake. South. Calif. (Colorado Desert); Mexico (Baja California). Pollen:
Unknown, but visits flowers of Aplopappus, Heliotropium cu rassavicum, Pectis

papposa.
Perdita abdominalis Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 38, figs. 550, 551, 667. 2,6.
Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 26 (geogr. and floral records).
affinis Cresson. Colo., N. Mex., Ariz. Pollen: Unknown, but visits flowers of Aplopappus
gracilis, Grindelia inornata, G. squarrosa, Helianthus petiolaris, Solidago.
Perdita affinis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 69. 2, d.

1896 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 11, 21, 108-109, figs. 636, 637, 720
(redescription, geogr. and floral records).
albipes Timberlake. Tex. (Big Bend National Park).
Perdita albipes Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 102, figs. 837, 838, 897. d.
apacheorum Timberlake. N. Mex. (Hidalgo County), Wyo. (Johnson County), Ariz. (Cochise
and Mohave Counties). Pollen: Unknown, but visits flowers of Asclepias, Baccharis,

Euphorbia, Grindelia squarrosa, Gutierrezia sarothrae, Heterotheca.
Perdita apacheorum Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 99. 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 67-68. d (geogr. and floral records).
aperta Timberlake. N. Mex. (Lincoln County), Ariz. (Navajo County). Pollen: Unknown, but
visits flowers of Gutierrezia microcephala, Lepidium montanum.
Perdita aperta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 68, figs. 9, 1231, 1232, 1319. 3.
aplopappi Timberlake. Ariz., Utah (Beaver and Washington Counties). Pollen: Unknown, but
visits flowers of Aplopappus gracilis, Eriogonum, Gutierrezia microcephala.
Perdita aplopappi Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 98, figs. 626, 627, 715. °,
3.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 69 (geogr. and floral record).
aridella Timberlake. Utah, Ariz. (Navajo County). Pollen: Unknown, but visits flowers of
Gutierrezia sarothrae, Helianthus.
Perdita aridella Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 81, figs. 600, 601, 702. 2, ¢.
atriventris Timberlake. Tex. (Calvert). Pollen: Unknown, but visits flowers of Heterotheca
subaxillaris.
Perdita atriventris Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 111, figs. 640, 641, 722. 3.
beatula Timberlake. Nev. (Churchill, Humboldt and Washoe Counties). Pollen: Unknown, but
visits flowers of Sphaeralcea.
Perdita beatula Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 22, figs. 522, 523, 662. d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 62, 69-70. 2 (geogr. and floral
records).
bigeloviae Cockerell. Tex. (E] Paso County), N. Mex., Colo. (Bent and Logan Counties). Pollen:
Unknown, but visits flowers of Aplopappus heterophyllus, A. pleuriflorus.
Perdita bigeloviae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 87. 6, 2.
Perdita fasciata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 57. 2.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 7, 15, 71-72, figs. 588, 589, 696
(redescription). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 70 (synonymy, geogr. and
floral records).

biparticeps Cockerell. N. Mex. Pollen: Unknown, but visits flowers of Pectis papposa.
Perdita biparticeps Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 84. d.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 121 (tax. characters). —Timberlake,
1968. Calif. Univ. Pubs. Ent. 49: 66, 70. ¢ (type, redescription).
bridwelli Timberlake. Ariz. (Yuma County), south. Calif. (Colorado Desert). Pollen: Unknown,
but visits flowers of Sphaeralcea including S. emoryi, S. orcuttii.
Perdita bridwelli Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 34, figs. 542, 548, 678. 2, d.
bruneri Cockerell. Tex. to Man. and Alta. east of Rocky Mts. Pollen: Unknown, but visits
flowers of Aplopappus divaricatus, Boltonia asteroides, Cleome serrulata, Grindelia
squarrosa, Solidago altissima, S. rigida.
Perdita bruneri Cockerell, 1897. Ent. News 8: 23. 6 (? misdet.).
Perdita Cockerelli Crawford, 1906. Canad. Ent. 38: 282. 6, 9.
Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 11, 21, 101-108, figs. 628, 629, 716
(redescription, geogr. and floral records).
butleri Timberlake. Ariz. (Gila Bend and Salome). Pollen: Unknown, but visits flowers of

Hymenothrix wislizeni, Pectis papposa.
Perdita butleri Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 96, figs. 624, 625, 714. @, o.

Superfamily APOIDEA 1897

chloris Timberlake. Ariz. (Yuma County), Nev. (Churchill and Lyon Counties), Calif. (Imperial
and Inyo Counties). Pollen: Unknown, but visits flowers of Dalea polyadenia,
Sphaeralcea.
Perdita chloris Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 64, figs. 576, 577, 690. 2, d.
claripennis Timberlake. N. Mex. (Dona Ana County).
Perdita claripennis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 71. 9.
cognata Timberlake. Wyo. (Sweetwater County), Utah (Grand County). Pollen: Unknown, but
visits flowers of Chrysothamnus.
Perdita cognata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 93, figs. 616, 617, 710. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 67, 71-72. 2, d (redescription, key,
tax. status, geogr. and floral records).
congrua Timberlake. Tex. (Marathon). Pollen: Unknown, but visits flowers of Gutierrezia.
Perdita congrua Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 26, figs. 1373, 1374, 1405. ¢.
consobrina consobrina Timberlake. N. C., S. C. Pollen: Unknown, but visits flowers of
Chrysopsis, Gerardia, Haplopappus, Heterotheca, Kuhnistera.
Perdita consobrina Timberlake, 1928. Amer. Mus. Novitates 321: 3. 2.

Taxonomy: Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 202-203. d. —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 12, 21, 109-111, figs. 638, 639, 721. 2, 5 (redescription).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 315, figs. 76-79, table 9
(redescription).

consobrina lepida Timberlake. Fla. (Dunedin, Lake Worth).
Perdita consobrina lepida Timberlake, 1952. Ent. Soc. Wash., Proce. 54: 203. 2, d.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 21, 111. —Mitchell, 1960. N. C.
Agr. Expt. Sta. Tech. Bul. 141: 315-316 (redescription).
croceipes Timberlake. N. Mex. (Eddy and Socorro Counties), Ariz. (Mohave and Yavapai
Counties), Utah (Kane County). Pollen: Unknown, but visits flowers of Baileya,

Gutierrezia including G. microcephala.
Perdita croceipes Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 72, figs. 590, 591, 697. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 64, 66, 72-73. 2 (key, geogr. and
floral records).
crotonis caerulea Timberlake. Utah (St. George), Nev. (Glendale). Pollen: Apparently an
oligolege of Croton including C. longipes.
Perdita crotonis caerulea Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 47. 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 73 (key).
crotonis crotonis Cockerell. N. Mex., Ariz. (Maricopa County); Mexico (Chihuahua). Pollen:
Apparently an oligolege of Croton.
Perdita crotonis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 64. 2, d.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 5, 17, 43-44, figs. 558, 559, 681
(redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 73
(key).

crotonis cucullata Timberlake. Calif. (Anza). Pollen: Apparently an oligolege of Croton
including C. californicus.
Perdito crotonis cucullata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 46. 2, d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 73 (key).

crotonis decipiens Timberlake. Tex. (Victoria).
Perdita crotonis decipiens Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 48. 2, d.

crotonis dilucida Timberlake. Ariz., possibly Kans. and Nebr. Pollen: Presumably an oligolege
of Croton, but visits flowers of Euphorbia.
Perdita crotonis dilucida Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 47. 2, d.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 73 (key).
crotonis juabensis Timberlake. Utah (Juab County). Pollen: Apparently an oligolege of Croton,

visits flowers of C. texensis.
Perdita crotonis juabensis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 103. 2, d.

1898 Hymenoptera in America North of Mexico

crotonis leucoptera Timberlake. South. Calif. (Riverside, San Diego and San Bernardino
Counties). Pollen: Apparently an oligolege of Croton californicus, but also visits flowers
of Hoffmanseggia presumably for nectar.
Perdita crotonis leucoptera Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 45. 2, ¢.
crotonis perpicta Timberlake. Nebr., N. Mex. (Arriba County). Pollen: Presumably an
oligolege of Croton, but visits flowers of Eriogonum, Euphorbia marginata.
Perdita crotonis perpicta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 74. 2, ¢.
crotonis subnitens Timberlake. Kans. (Reno County). Pollen: Apparently an oligolege of
Croton, visits flowers of C. monanthogynus.
Perdita crotonis subnitens Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 48. 9.
crotonis titusi Timberlake. Calif. (Chino).
Perdita crotonis titusi Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 46. 2, 6.
crotonis undecimalis Cockerell. N. Mex. (San Miguel County). Pollen: Presumably an oligolege
of Croton, visits flowers of Croton sp.
Perdita crotonis undecimalis Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 281. 2.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 5, 45. 2° (key, redescription).
dalyi Timberlake. Tex. (E] Paso). Pollen: Unknown, but visits flowers of Haplopappus
heterophyllus.
Perdita dalyi Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 119. 9.
dilecta Timberlake. Utah (Kane County), Tex. (E] Paso).
Perdita dilecta Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 40, figs. 554, 555, 679. 2, 3.
discreta Timberlake. N. C. (coastal plains and sand dunes). Pollen: Unknown, but visits flowers
of Haplopappus.
Perdita discreta Timberlake, 1954. Ent. News 65: 14. °.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 22, 115-116, figs. 646, 647, 725. 6,
2 (key, redescription, geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 76, 77 table 9 (redescription).
dolichocephala Swenk and Cockerell. Nebr., Kans., Colo. (Weld County). Pollen: Unknown, but
visits flowers of Amorpha canescens, Boltonia asteroides, Euphorbia, Helianthus
petiolaris, H. subrhomboideus, Heterotheca subaxillaris.
Perdita dolichocephala Swenk and Cockerell, 1907. Ent. News 18: 54. @.
Perdita wunderi Cockerell, 1922. Amer. Mus. Novitates 33: 10. 2.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 19, 116-117, figs. 648, 649, 726. °,
6 (redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49:
74 (synonymy, tax. characters).
electa Timberlake. Wyo (Sweetwater County). Pollen: Unknown, but visits flowers of
Chrysothamnus nauseosus.
Perdita electa Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 86. 2.
elegans Timberlake. Calif. (Colorado Desert), Ariz. (Yuma). Pollen: Unknown, but visits
flowers of Abronia, Larrea tridentata, Palafoxia linearis, Pectis papposa.
Perdita elegans Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 55, figs. 568, 569, 686. 2, 3.
fallax Cockerell. Tex. to Ariz., north to Nebr. and Mont. Pollen: Unknown, but visits flowers of
Baileya, Eriogonum, Grindelia, Gutierrezia, Helianthus including H. petiolaris,
Heterotheca, Verbesina encelioides. Predator: Philanthus crabroniformis Smith, P.
pacificus arizonae Dunning.
Perdita fallax Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 90. 9.
Perdita erigeronis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 398. ¢.
Perdita fallax fontis Cockerell, 1922. Amer. Mus. Novitates 33: 9. °.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 11, 20, 106-108, figs. 634, 635, 719
(key, redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent.
49: 65, 74-75 (key, synonymy, variation, geogr. and floral records).
flavicauda Timberlake. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers of Larrea
tridentata, Melilotus, Nama hispidum, Sphaeralcea.
Perdita flavicauda flavicauda Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 39. 2
(Misspelled flavicanda flavicanda in key, p. 4).

Superfamily APOIDEA _ 1899

Perdita flavicauda formosa Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 40. 2 (Misspelled
flavicanda formosa in key, p. 4).

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63, 66, 75-76, figs. 1233, 1234, 1320.
3, 2 (key, synonymy, tax. characters, geogr. and floral records).
flaviceps Timberlake. Nev. (Las Vegas).
Perdita flaviceps Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 37, figs. 548, 549, 676. ¢.
flavifrons Timberlake. N. Mex. (Carrizozo), Utah (Lucia), south. Calif. (Colorado Desert).

Pollen: Unknown, but visits flowers of Chrysothamnus, Gutierrezia microcephala.
Perdita flavifrons Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 76, figs. 1235, 1236, 1321.
3

fuscipes Timberlake. N. Mex. (Arch in Roosevelt County).
Perdita fuscipes Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 77. °.
gerardiae Crawford. N. C., Fla., Miss. Pollen: Apparently an oligolege of Gerardia including G.
fasciculata, G. purpurea.
Perdita gerardiae Crawford, 1932. Ent. Soc. Wash., Proc. 34: 75. 6, 9.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 13, 18, 68, figs. 582, 583, 693. 2, 3
(redescription). — Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 317, figs. 76-79, table
9 (redescription).
gerhardi dallasiana Cockerell. Tex. Pollen: Unknown, but visits flowers of Aster
tanacetifolius, Dalea aurea, Helianthus annuus, Monarda citriodora, M. punctata.
Perdita dallasiana Cockerell, 1906. Entomologist 39: 178. ¢.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 13, 14, 15, 28-29. 2, d
(redescription, tax. status, geogr. and floral records).
gerhardi gerhardi Viereck. Wis., Ill., Ind. Pollen: Unknown, but visits flowers of Monarda

punctata.
Perdita gerhardi Viereck, 1904. Ent. News 15: 21. 9, 6.
Perdita gerhardi arenicola Timberlake, 1929. N. Y. Ent. Soc., Jour. 37: 119. 2, 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 2, 3, 18, 27-28 (redescription,
synonymy, geogr. range). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 317-318
(redescription, as gerhardi gerhardi and g. arenicola).

gerhardi monardae Viereck. N. J., Va., Fla. Pollen: Unknown, but visits flowers of Monarda
punctata.
Perdita monardae Viereck, 1904. Ent. News 15: 22. 2, 6.

Taxonomy: Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 204 (tax. status). —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 7, 15, 28 (redescription). — Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 318, figs. 76, 77, table 9 (redescription).

gutierreziae Cockerell. Colo., Tex., N. Mex., Ariz., Calif. (Mojave Desert). Pollen: Unknown,
but visits flowers of Baileya pleniradiata, Chrysothamnus, Gutierrezia lucida, G.
microcephala, Haplopappus acradenius, H. heterophyllus, Solidago confinis,
Sphaeralcea.
Perdita gutierreziae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 85. d.

Taxonomy: Swenk and Cockerell, 1907. Ent. News 18: 56. d, ?. —Timberlake, 1960. Calif.
Univ. Pubs. Ent. 17: 8, 16, 74-76, figs. 594, 595, 699. 2°, d (key, redescription, geogr. and
floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 65, 77-78 (tax. status, geogr.
and floral records).

halli Timberlake. Calif. (Palm Springs).
Perdita halli Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 89, figs. 612, 613, 708. 2, d.
hirsuta Cockerell. N. Mex., Ariz. (Willcox). Pollen: Unknown, but visits flowers of Dalea
including D. scoparia.
Perdita hirsuta Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 79. ¢.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 8, 18, 66, figs. 580, 581, 692. 2, 3

(redescription).

1900 Hymenoptera in America North of Mexico

hirtella Timberlake. Utah (Wasatch County).
Perdita hirtella Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 78, fig. 10. 9°.
hirticeps apicata Timberlake. Calif. (Colorado and Mojave Deserts), Ariz. (Mohave County),
Utah (Cornish and Oak City). Pollen: Collects pollen from flowers of Stephanomeria
including S. exigua, S. pauciflora, but visits these and other flowers for nectar including
Malacothrix glabrata.
Perdita hirticeps apicata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 59. 2, 6.
Perdita hirticeps candidipennis Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 60. 2.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 79 (tax. characters, as hirticeps
apicata and h. candidipennis). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 28
(synonymy, geogr. and floral record).

hirticeps hirticeps Timberlake. Calif. (cismontane). Pollen: Collects pollen from the flowers of
Stephanomeria including S. exigua, S. e. var. coronaria, S. virgata, but visits these and
other flowers for nectar including Coreopsis lanceolata, Hemizonia.
Perdita hirticeps hirticeps Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 58, figs. 570, 571,
GSienericr

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 79 (tax. characters, variation).
hirticeps luteocincta Timberlake. Calif. (Antioch). Pollen: Unknown, but visits flowers of
Gutierrezia californica.
Perdita hirticeps luteocincta Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 60. °.
indioensis Timberlake. Calif. (Indio). Parasite: Neolarra vigilans (Ckll.). Pollen: Unknown, but
visits flowers of Haplopappus acradenius.
Perdita indioensis Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 54, figs. 566, 567, 685. 2,
3.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63 (key).
idonea Timberlake. Utah, Ariz. (Coconino County). Pollen: Unknown, but visits flowers of
Gutierrezia.
Perdita idonea Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 79, figs. 1237, 1238, 1322. °,
é.
jonesi Cockerell. Tex., Kans. Pollen: Unknown, but visits flowers of Dalea aurea, Monarda
citriodora, M. punctata, Petalostemon multiflorwm.
Perdita jonesi Cockerell, 1906. Entomologist 39: 177. 2, 3.
Perdita birkmanni Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 282. °.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 22, 114-115, figs. 644, 645, 724
(redescription, synonymy, geogr. and floral records).
knowltoni Timberlake. Utah (Moab). Pollen: Unknown, but visits flowers of Chrysothamnus.
Perdita knowltoni Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 51, figs. 562, 563, 683. °
(3 misdet.).

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63, 80 (key, tax. status, geogr. and
floral record).
labergei Timberlake. Kans. (Hutchinson). Pollen: Unknown, but visits flowers of Euphorbia.
Perdita labergei Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 41, figs. 556, 557, 680. 2, d.
lasiogastra Timberlake. Tex. (Cameron County), N. Mex. (San Miguel County), Colo. (Moffat
County). Pollen: Unknown, but visits flowers of Pectis papposa.
Perdita lasiogastra Timberlake, 1929. N. Y. Ent. Soc., Jour. 37: 115. ¢.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 9, 20, 84-86, figs. 606, 607, 705. 2, 3
(key, redescription, geogr. and floral records).
laticincta Swenk and Cockerell. N. Dak., Nebr., Kans., Wyo., N. Mex. Pollen: Unknown, but
visits flowers of Amorpha canescens, Haplopappus ciliatus, Helianthus petiolaris,
Melilotus.
Perdita laticincta Swenk and Cockerell, 1907. Ent. News 18: 52. 2, 6.
Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 8, 20, 76, figs. 596, 597, 700. 2, 3
(key, redescription, geogr. and floral records).

Superfamily APOIDEA 1901

luteiceps Cockerell. Colo. (Conejos, Garfield and Routt Counties), Wyo. (Fremont County).
Pollen: Unknown, but visits flowers of Gutierrezia.
Perdita luteiceps Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 74. d.

Taxonomy: Timberlake, 1928. N. Y. Ent. Soc., Jour. 37: 112. d, 2. —Timberlake, 1960. Calif.
Univ. Pubs. Ent. 17: 3, 14, 36-37, figs. 544, 545, 674. 2, ¢ (key, redescription, geogr
records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 81 (geogr. and floral records).

luteola Cockerell. Nebr., Colo., N. Mex., Ariz. Pollen: Unknown, but visits flowers of

Chrysothamnus, Gutierrezia sarothrae, Haplopappus heterophyllus.
Perdita luteola Cockerell, 1894. Ent. News 5: 328. 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 2, 18, 23-24, figs. 526, 527, 664 (key,
geogr. and floral records).

maculigera var. bilineata Timberlake. IIl., Mo., Kans., Nebr., S. Dak.
Perdita maculipennis var. bilineata Timberlake, 1929. N. Y. Ent. Soc., Jour. 37: 121. 2, d.

Taxonomy: Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 204 (tax. status). —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 3, 14, 33 (tax. status, tax. characters, key, geogr. records).
—Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 320, figs. 76, 77, table 9 (tax.
characters).

maculigera maculigera Cockerell. Tex. (Southmost), N. Mex. (Las Cruces). Pollen: Unknown,
but visits flowers of Salix.
Perdita maculigera Cockerell, 1896. Ent. News 7: 255. d.

Taxonomy: Timberlake, 1952. Ent. Soc. Wash., Proc. 54: 204 (tax. status). —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 3, 14, 32-33 (key, redescription, geogr. and floral records).
maculigera maculipennis Graenicher. Minn., Wis., Mich., Ill., lowa, Kans., Tex. (Anderson and

Robertson Counties). Parasite: Ewphyto pollinaris Reinhard. Pollen: Collects pollen
from flowers of Salix nigra although has been listed as collecting pollen from flowers of
Melilotus alba, visits other flowers presumably for nectar including Erigeron
philadelphicus.

Perdita maculipennis Graenicher, 1910. Canad. Ent. 42: 102. 2.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 9, 14, 33-34, figs. 540, 541, 671, 672.
2, 3 (key, redescription, geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 320-321, figs. 75, 77, table 9 (redescription). —Rozen, 1966. Amer. Mus.
Novitates 2259: 19-20 (larva). —Yager and Rozen, 1966. Amer. Mus. Novitates 2265: 12
(pupa).

Biology: Michener and Ordway, 1963. Kans. Ent. Soc., Jour. 36: 34-45, 10 figs. (nest
architecture, life history, immature stages, flight behavior, pollen source, parasite).

maculipes Cockerell. N. Mex. (Las Cruces). Pollen: Unknown, but visits flowers of
Haplopappus heterophyllus.
Perdita maculipes Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 87. d.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 121 (tax. characters). —Timberlake,
1968. Calif. Univ. Pubs. Ent. 49: 65, 81-82 (key, redescription of type).

media Timberlake. Calif. (Colorado and Mojave Deserts). Pollen: Unknown, but visits flowers
of Gutierrezia lucida, Haplopappus acradenius, Solidago confinis.
Perdita media Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 78, figs. 592, 598, 698. 2, d.
melanostoma albocincta Timberlake. N. Mex., Ariz. Pollen: Unknown, but visits flowers of
Baileya, Gutierrezia, Haplopappus spinulosus.
Perdita albocincta Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 78. 2.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63, 83-84 (geogr. and floral records).
melanostoma melanostoma Swenk and Cockerell. Idaho, Utah, Colo., Nebr., N. Mex., Ariz.
Pollen: Unknown, but visits flowers of Chrysothamnus, Gutierrezia including G.

sarothrae.
Perdita melanostoma Swenk and Cockerell, 1907. Ent. News 18: 57. 2.

1902 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 8, 78 (key, redescription, geogr. and
floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63, 65, 82-83, figs. 11, 1241,
1242, 1324. 3 (key, redescription, geogr. and floral records).

mesillensis Timberlake. N. Mex. (Mesilla Park). Pollen: Unknown, but visits flowers of
Gutierrezia lucida, Pectis papposa.
Perdita mesillensis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 84, figs. 1243, 1244, 1325.
é.
Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 28 (tax. characters, synonymy).
microsticta Timberlake. Kans. (Scott County).
Perdita microsticta Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 89. 2.

mimula Timberlake. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers of

Haplopappus acradenius.
Perdita mimula Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 53. 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63 (key).
nuda Cockerell. N. Mex., Ariz., Colo., Utah, Idaho (Franklin County). Parasite: Megaselia,
Sphecodes sp. near fragariae Ckll. Pollen: Apparently collects pollen only from vernal
and autumnal flowering Compositae including Chrysothamnus, Erigeron, Grindelia

including G. squarrosa.
Perdita nuda Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 93. 9°.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 119. 2 (redescription, key).
—Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 104-105, figs. 839, 840, 898. d, 2 (geogr.
and floral records). —Torchio, 1975. Kans. Ent. Soc., Jour. 48: 266-275, 277-278, figs. 1-18,
26, 27 (immature stages).

Biology: Torchio, 1975. Kans. Ent. Soc., Jour. 48: 257-279, 27 figs. (nest architecture, life
history, nest associates).
numerata hesperia Timberlake. South. Calif. (Colorado Desert); Mexico (Baja California).
Pollen: Unknown, but visits flowers of Geraea canescens, Larrea tridentata, Melilotus,
Prosopis, Tamarix.
Perdita numerata hesperia Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 62. 2, 6.
numerata numerata Cockerell. Tex., N. Mex., Ariz. (Phoenix), ?Calif. (Needles). Pollen:
Unknown, but visits flowers of Actinea, Prosopis, Salix.
Perdita numerata Cockerell, 1895. Amer. Ent. Soc., Trans. 22: 296. 9.

Taxonomy: Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 281. 6. —Timberlake, 1960.
Calif. Univ. Pubs. Ent. 17: 6, 15, 60-62, figs. 572, 573, 688 (key, redescription, tax. status,
geogr. and floral records).

occidua Timberlake. Ariz. (Oak Creek Canyon).
Perdita occidua Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 120. 9.
occlusa Timberlake. Ariz. (Walnut Canyon near Flagstaff).
Perdita occlusa Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 85, figs. 1245, 1246, 13826. 3.
octomaculata octomaculata (Say). N. B. south to Ga., west to Minn., Ill. and Miss. Pollen:
Unknown, but visits flowers of Aster ericoides, A. puniceus, Bidens, Boltonia,
Eupatorium, Euthamia, Lycopus, Monarda, Solidago.
Panurgus 8-maculatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 350. d, 2.
Perdita octomaculata Dalla Torre, 1896. Cat. Hym., v. 10, p. 173. Emend.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 22, 111-118, figs. 642, 643, 723
(key, redescription, geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech.
Bul. 141: 321-322, figs. 76-79, table 9 (redescription, geogr. range, floral records).

octomaculata terminata Cockerell. N. Dak., Nebr., Kans., Iowa, IIl., Ark., Miss. Pollen:
Unknown, but visits flowers of Aster azwreus, A. multiflorus, A. saliciflorus, Solidago
canadensis, S. rigida.
Perdita octomaculata terminata Cockerell, 1922. Amer. Mus. Novitates 33: 8. 6, @.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 12, 22, 1138-114 (key, tax. characters,
geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 322 (tax.
characters, geogr. range).

Superfamily APOIDEA 1903

ordinata Timberlake. Ariz. (Cochise and Navajo Counties), N. Mex. (Hidalgo and Socorro
Counties), Utah (Cache and Washington Counties). Pollen: Unknown, but visits flowers
of Baileya pleniradiata, Dithyrea including D. wislizenii, Heterotheca subazillaris,
Pectis papposa.
Perdita ordinata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 105, figs. 841, 842, 899. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 67, 86, fig. 12 (key, geogr. and floral
records). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 28-29. ¢, 2 (geogr. and floral
records).

paroselae Timberlake. South. Calif. (Colorado Desert). Pollen: Possibly an oligolege of Dalea,
visits flowers of Dalea californica, D. emoryi.
Perdita paroselae Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 65, figs. 578, 579, 691. 2, d.
pectidis Cockerell. N. Mex., Ariz. (Salome). Pollen: Unknown, but visits flowers of
Kallstroemia, Pectis papposa, Sesuvium verrucosum, Wedeliella.
Perdita pectidis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 83. 2, 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 9, 19, 91-92, figs. 614, 615, 709. 2, d
(key, tax. characters, geogr. and floral records).
percincta Timberlake. Utah (Skull Rock Pass, 6,100ft.), Ariz. (Coconino County). Pollen:
Unknown, but visits flowers of Aster, Gutierrezia microcephala, Wislizenia refracta.
Perdita percincta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 86. ?.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 30-31, figs. 1379, 1380, 1408. d, °
(key, redescription, geogr. and floral records).
perixantha Timberlake. Ariz. (Navajo County).
Perdita perixantha Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 23, figs. 524, 525, 668. d.
perpallida citrinella Graenicher. Wis., Minn., N. Dak., Nebr., Alta. Pollen: Unknown, but visits
flowers of Petalostemon purpureum, P. villosum.
Perdita citrinella Graenicher, 1910. Canad. Ent. 42: 103. °.

Taxonomy: Crawford, 1912. Canad. Ent. 44: 359. d, 2. —Timberlake, 1952. Ent. Soc. Wash.,
Proc. 54: 204 (tax. status). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 3, 14, 26-27 (key,
tax. characters, geogr. and floral records).

perpallida perpallida Cockerell. Man., Minn., N. Dak., Nebr., Kans., N. Mex. Pollen:
Apparently an oligolege of vernal and autumnal flowering Compositae, visits flowers of
Amorpha canescens, Helenium autumnale, Petalostemon candidum, P. oligophyllum,
P. villosum, Solidago occidentalis.
Perdita perpallida Cockerell, 1901. Entomologist 34: 190. °, d.

Taxonomy: Swenk and Cockerell, 1907. Ent. News 18: 57. —Stevens, 1919. Canad. Ent. 51:

207. 2, 3 (tax. characters, floral records). —Timberlake, 1952. Ent. Soc. Wash., Proc. 54:

204 (tax. status). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 2, 13, 25-26, figs. 532, 533,

667. 2, 3 (key, redescription, geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt.

Sta. Tech. Bul. 141: 322 (tax. status).

phymatae Cockerell. N. Mex., Ariz., Utah. Pollen: Unknown, but visits flowers of Bahia

absinthifolia, Baileya pleniradiata, Gutierrezia lucida, G. microcephala, Haplopappus
heterophyllus, Pectis papposa. Predator: Phymata sp.

Perdita phymatae Cockerell, 1895. Acad. Nat. Sci. Phila:. Proce. 472 12..9.

Perdita nitidella Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 16. d.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 9, 92-93 (key, redescription,
geographic, floral and predator records). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17:
70-71, figs. 586, 587, 695. d (as nitidella). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49:
64, 85. ° (key, redescription, as nitidella). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49:
65 (key). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 31 (synonymy, geogr. range, tax.
characters).

picturata Timberlake. Tex. (Victoria and Aransas Counties).
Perdita picturata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 50, figs. 560, 561, 682. 2, ¢.

1904 Hymenoptera in America North of Mexico

plucheae Timberlake. Calif. (Colorado Desert), Ariz.; Mexico (Baja California). Pollen:
Unknown, but visits flowers of Pluchea sericea, Tamarix gallica.
Perdita plucheae Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 69, figs. 584, 585, 694. 2, d.
polygonellae Timberlake. N. C. (Coastal Plain). Pollen: Unknown, but visits flowers of
Polygonella polygama.
Perdita polygonellae Timberlake, 1954. Ent. News 65: 12. 2, 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 4, 16, 40, figs. 552, 553, 678 (key, tax.
characters). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 323, figs. 76-79, table 9
(redescription).

prionopsidis Timberlake. Kans. (Reno and Montgomery Counties). Pollen: Unknown, but visits
flowers of Boltonia asteriodes, Haplopappus ciliatus, Helianthus petiolaris, Heterotheca
subaxillaris.
Perdita prionopsidis Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 104, figs. 632, 638, 718.
OOr
rectangulata Cockerell. Colo., Idaho, Ariz. (Holbrook). Pollen: Unknown, but visits flowers of
Chrysothamnus, Gutierrezia sarothrae, Solidago.
Perdita rectangulata Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 72. 2 (6 misdet.).

Taxonomy: Stevens, 1919. Canad. Ent. 51: 207. d, 2 (as martini). —Timberlake, 1960. Calif.
Univ. Pubs. Ent. 17: 3, 9, 16, 82-83, figs. 602, 603, 703. 2, d (key, redescription, geogr. and
floral records, in part martini). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 64, 65,
87-88 (key, tax. characters and tax. status).

reperta Timberlake. N. Mex. (Quemado), Ariz. (Navajo County), Colo. (Alamosa County).
Pollen: Unknown, but visits flowers of Chrysothamnus including C. viscidiflorus.
Predator: Philanthus sp.
Perdita reperta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 88, figs. 18, 1247, 1248, 1327.
Or
retusa Timberlake. Ariz. (Mohave and Yuma Counties), Calif. (Colorado and Mojave Deserts).
Pollen: Unknown, but visits flowers of Chrysothamnus paniculatus, Haplopappus.
Perdita retusa Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 86, figs. 608, 609, 706. 2, d.
rhodura Cockerell. Ariz., N. Mex., Nebr. (Sioux County). Pollen: Unknown, but visits flowers
of Gutierrezia sarothrae, Haplopappus, Zinnia grandiflora.
Perdita rhodura Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 511. 2, 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 6, 15, 52-58, figs. 564, 565, 684. 2, 3
(key, redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent.
49: 63, 66 (key).
scitula antiochensis Timberlake. Calif. (Antioch, Oakley). Pollen: Unknown, but visits flowers
of Eriogonum, Gutierrezia californica, Heterotheca grandiflora, Lessingia glandulifera.
Perdita scitula antiochensis Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 80, figs. 598, 599,
701. 2, do.
scitula scitula Timberlake. Calif. (Inyo County). Pollen: Unknown, but visits flowers of
Chrysothamnus nauseosus var. gnaphalodes.
Perdita scitula scitula Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 79. 2, 6.
sedulosa Timberlake. N. Mex. (Socorro County). Pollen: Unknown, but visits flowers of
Baileya pleniradiata.
Perdita sedulosa Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 31. 9.
sejuncta Timberlake. N. Mex. (Socorro County), Utah (Wah Wah Mts.); Mexico (Chihuahua).
Pollen: Unknown, but visits flowers of Gutierrezia microcephala.
Perdita sejuncta Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 90. 2.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 32 (geogr. record).
senecionis Cockerell. N. Mex. (Las Cruces). Pollen: Unknown, but visits flowers of Senecio
douglasii.
Perdita senecionis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 94. °.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 11, 100-101. 2 (redescription, key).

Superfamily APOIDEA 1905

separata Timberlake. Tex. (Zapata County).
Perdita separata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 106. °.
snowii Cockerell. Colo. (Boulder, Costilla and Moffat Counties), Ariz. (Holbrook). Pollen:
Unknown, but visits flowers of Aster, Grindelia, Gutierrezia.
Perdita snowti Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 73. 2.

Taxonomy: Cockerell, 1910. Psyche 17: 244. 6. —Timberlake, 1960. Calif. Univ. Pubs. Ent.
17: 9, 20, 83-84, figs. 604, 605, 704. 2, ¢ (key, redescription, geogr. and floral records).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 64, 91 (tax. characters, key).

solidaginis Cockerell. Colo. (White Rocks, near Boulder). Pollen: Unknown, but visits flowers
of Solidago.
Perdita solidaginis Cockerell, 1922. Amer. Mus. Novitates 33: 12. 6.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 316-317 (tax. characters).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 66, 91-92. ¢ (key, redescription of type
specimen).

stepheni Timberlake. Tex. (Dimmit County). Pollen: Unknown, but visits flowers of Monarda
punctata.
Perdita stepheni Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 29, figs. 536, 537, 669. 3.
submedia Timberlake. Ariz. (Apache County). Pollen: Unknown, but visits flowers of
Chrysothamnus.
Perdita submedia Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 32. 9.
swenki Crawford. N. Y. (Long Island), Mich., Ill., Wis., Minn., N. Dak., Nebr., Alta. Pollen:
Unknown, but visits flowers of Chrysopsis, Grindelia, Helianthus maximillianii,
Liatris, Solidago juncea, S. rigida.
Perdita swenki Crawford, 1915. Insecutor Inscitiae Menstruus 3: 109. d, °.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 10, 19, 103-104, figs. 630, 631, 717. 2,
6 (key, redescription, geogr. and floral records). —Mitchell, 1960. N. C. Agr. Expt. Sta.
Tech. Bul. 141: 324, figs. 76-78, table 9 (redescription). —Timberlake, 1971. Calif. Univ.
Pubs. Ent. 66: 33 (geogr. records).

tacita Timberlake. Idaho (Owyhee County).
Perdita tacita Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 93. 9°.
translineata Timberlake. Tex. (Brazos and Van Zandt Counties), N. Mex. (Socorro County).
Pollen: Unknown, but visits flowers of Baileya pleniradiata, Eriogonum multiflorum,
Haplopappus divaricatus, Heterotheca subaxillaris.
Perdita translineata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 93, figs. 618, 619, 711. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 65, 67, 94. °, ¢ (key, redescription,
geogr. and floral records).
tridentata Stevens. N. Dak., Nebr., Kans. Pollen: Unknown, but visits flowers of Haplopappus,
Helianthus petiolaris.
Perdita tridentata Stevens, 1919. Canad. Ent. 51: 206. d.
Taxonomy: Stevens, 1921. Canad. Ent. 53: 66. 9. —Timberlake, 1960. Calif. Univ. Pubs. Ent.
17: 12, 21, 118-119, figs. 650, 651, 727. 2, d (key, redescription, geogr. and floral records).
trifida Timberlake. Calif. (Mojave Desert). Pollen: Unknown, but visits flowers of Baccharis

emoryi, Haplopappus acradenius.
Perdita trifida Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 37, figs. 546, 547, 675. d.

trimaculata Timberlake. South. Calif. (Colorado and Mojave Deserts), Ariz. (Mohave County).
Pollen: Unknown, but visits flowers of Chrysothamnus nauseosus, C. paniculatus,

Gutierrezia lucida.
Perdita trimaculata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 62, figs. 574, 575, 689. °,

3.

truncatella Timberlake. Ariz. (Snowflake).
Perdita truncatella Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 94, figs. 1251, 1252, 1329.

é.

1906 Hymenoptera in America North of Mexico

variegata pura Timberlake. Tex., Kans. (Garden City). Pollen: Unknown, but visits flowers of
Monarda punctata var. occidentalis.
Perdita variegata pura Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 31. 9, 6.
variegata variegata Timberlake. Kans. Pollen: Unknown, but so far has been collected
exclusively at the flowers of Monarda including M. punctata var. occidentalis.
Perdita variegata variegata Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 30, figs. 538, 539,
670. 2, d.
versuta Timberlake. Tex. (Kingsville).
Perdita versuta Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 105, figs. 848, 844, 900. ¢.
xanthochroa Timberlake. Colo. (Moffat County), Utah (Juab and Uinta Counties), Idaho
(Owyhee County). Pollen: Unknown, but visits flowers of Sphaeralcea ambigua,

Stanleya pinnata.
Perdita xanthochroa Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 24, figs. 530, 531, 666. 9,
3

xanthodes Timberlake. Calif. (Mohave Desert). Pollen: Unknown, but visits flowers of
Gutierrezia lucida, Haplopappus cooperi.
Perdita xanthodes Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 24, figs. 528, 529, 665. °,
é.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 63 (key). —Timberlake, 1971. Calif.
Univ. Pubs. Ent. 66: 33 (geogr. and floral records).
xanthops Timberlake. Wyo. (Carbon County).
Perdita xanthops Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 33, figs. 1381, 1382, 1409. @,
Ore

SPECIES GROUP VENTRALIS

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1, 2-14 (key, tax. characters, key to
included spp.). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 95-97 (suppl. key to included
spp.).

acaciae Timberlake. South. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers of
Acacia greggti, Agave consociata, A. deserti.
Perdita acaciae Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 61, figs. 799, 800, 878. 2, ¢.
albata Timberlake. Ariz. (Navajo County). Pollen: Unknown, but visits flowers of Mentzelia
mila.
Perdita albata Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 97. 2.
amplipennis Timberlake. Calif. mts. (Inyo and Mono Counties). Pollen: Unknown, but visits

flowers of Eriogonum fasciculatum, Stanleya pinnata, and presumably Dalea.
Perdita amplipennis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 30, figs. 751, 752, 854. ?,
é.

austini Cockerell. N. Mex. Pollen: Unknown, but visits flowers of Baileya pleniradiata,
Gutierrezia microcephala, G. sarothrae, Haplopap pus.
Perdita austini Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 13. d.
Perdita aeneifrons Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 91. ?.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 10, 40-41, figs. 769, 770, 863. 3
(redescription). —Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 125. d, 2 (synonymy).
—Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 96, 97, 98-99 2, ¢ (key, redescription,
geogr. and floral records).

bicuspidariae Timberlake. South. Calif. (Colorado Desert). Pollen: Collects pollen from the
flowers of Mentzelia involucrata, but also visits flowers of Hyptis emoryi presumably
for nectar.
Perdita bicuspidariae Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 20, figs. 741, 742, 849.
On oe
brevihirta Timberlake. Ariz. (Maricopa County), Wyo. (Fremont County). Pollen: Unknown,
but visits flowers of Chrysothamnus nauseosus, Pectis papposa.
Perdita brevihirta Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 41, figs. 771, 772, 864. 9,
3.

Superfamily APOIDEA 1907

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 34 (geogr. and floral records).
claypolei australior Timberlake. Calif. (San Diego and Los Angeles Counties). Pollen:
Unknown, but visits flowers of Adenostoma fasciculatum, Eriogonum fasciculatum,
Hemizonia fasciculata, Heteromeles arbutifolia, Opuntia.
Perdita claypolei australior Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 68. 2, é.
claypolei claypolei Cockerell. South. Calif. (cismontane). Pollen: Collects pollen from the
flowers of Eriogonum including E. elongatum, E. fasciculatum, E. gracile, E. wrightii
var subscaposum, E. umbellatum, but visits these and other flowers for nectar
including Eriastrum virgatum.
Perdita claypolei Cockerell, 1901. Canad. Ent. 33: 281. 2.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 6, 13, 66-68, figs. 805, 806, 881. 2, ¢
(key, redescription, geogr. and floral records).
claypolei limatula Timberlake. Calif. (Kern County) to Oreg., and Nev. (Douglas County).
Pollen: Collects pollen from the flowers of Eriogonum including E. elatwm, E.
fasciculatum, E. gracile, E. heermannii, E. inflatum, E. latifolium, E. nudum, E.
vimineum, E. virgatum, E. wrightii var. subscaposum, but visits these and other
flowers for nectar including Achillea millefolium, Brickellia, Cirsium, Solidago.
Perdita claypolei limatula Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 68. 2, ¢.
clypeata clypeata Timberlake. South. Calif. (Colorado Desert). Pollen: Unknown, but visits
flowers of Eriogonum inflatwm.
Perdita clypeata clypeata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 79, figs. 817, 818,
887. 2, do.
clypeata immaculata Timberlake. Ariz. (near Mesa), south. Calif. (Desert Center and
Hayfield). Pollen: Unknown, but visits flowers of Eriogonum inflatum.
Perdita clypeata immaculata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 80. 2, ¢.
colei Timberlake. Calif. (San Bernardino Mts. and vicinity). Pollen: Unknown, but visits
flowers of Chorizanthe staticoides, Chrysothamnus nauseosus, Gutierrezia lucida,
Lepidospartum squamatum, Solidago confinis.
Perdita colei Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 48, figs. 783, 784, 870. 2°, d.
crandalli Timberlake. Ariz. (Pima and Pinal Counties). Pollen: Unknown, but visits flowers of
Petalonyx thurberi.
Perdita crandalli Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 38. °.
dasylirii Cockerell. Western Tex. to Ariz. Pollen: Unknown, but visits flowers of Acacia,
Agave palmeri, Amorpha fruticosa, Asclepias galioides, Ceanothus, Dasylirion wheeleri,
Euphorbia albomarginata, Mertensia franciscana, Nolina microcarpa, Rudbeckia
laciniata, Salix taxifolia, Sapindus saponaria.
Perdita dasylirii Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 5389. “2” = ¢ (6
misdet.).
Perdita nolinae Cockerell, 1922. Amer. Mus. Novitates 33: 11. °.
Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 5, 12, 55-57, figs. 798, 794, 875. 2, d
(key, redescription, geogr. and floral records).
dimidiata Timberlake. Ariz. (Pinal County); Mexico (Baja California). Pollen: Unknown, but
visits flowers of Seswviwm verrucosum.
Perdita dimidiata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 39, figs. 767, 768, 862. ¢.
distans Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Unknown, but visits
flowers of Eriogonum including E. reniforme, E. thomasii.
Perdita distans Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 73, figs. 811, 812, 884. 2, d.
erythropyga Timberlake. South. Calif. (Colorado Desert). Pollen: Collects pollen of Dalea
including D. californica, D. emoryi, but also visits other flowers for nectar including
Cryptantha angustifolia, Isomeris arborea, Larrea tridentata.
Perdita erythropyga Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 42, figs. 773, 774, 865. 2,
3.

exilis Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Collects pollen from

Petalonyx thurberi.
Perdita exilis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 36, figs. 7638, 764, 860. d, °@.

1908 Hymenoptera in America North of Mexico

fulvicauda Timberlake. South. Calif. (Colorado Desert); Mexico (Baja California). Pollen:
Unknown, but visits flowers of Baileya pleniradiata, Coldenia plicata, Dalea mollis,
Larrea tridentata, Melilotus, Nama hispidum, Phacelia distans, Prosopis juliflora,
Verbena.
Perdita fulvicauda Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 43, figs. 775, 776, 866. °,
3.
gentilis Timberlake. Nev. (Churchill and Washoe Counties), Ariz. (Vermillion Cliffs). Pollen:
Unknown, but visits flowers of Eriogonum.
Perdita gentilis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 78. 2.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 96, 97, 109-110. 2°, 3 (key, geogr.
and floral record).
glabrella Timberlake. Ariz. (Continental).
Perdita glabrella Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 104. 9.
glabrescens Timberlake. Calif. (Inyo County).
Perdita glabrescens Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 38, figs. 765, 766, 861. 2,
3.
grandiceps Cockerell. N. Mex. (Alamogordo, La Cueva and Las Cruces). Pollen: Unknown, but
visits flowers of Fallugia paradoxa, Solidago.
Perdita grandiceps Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 96. 6.

Taxonomy: Cockerell, 1899. Canad. Ent. 31: 256. 2. —Timberlake, 1962. Calif. Univ. Pubs.
Ent. 28: 3, 10, 32-33, figs. 757, 758, 857. 2, d (key, redescription, geogr. and floral records).
holoxantha Timberlake. Utah (Grand and Washington Counties), Ariz. (Holbrook). Pollen:

Unknown, but visits flowers of Mentzelia pumila, Poleomintha incana.
Perdita holoxantha Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 17, figs. 737, 738, 847. d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 95, 96, 100. 2, 3d (key, geogr. and
floral records).
inornata Timberlake. Calif. (Clark Mt. and Panamint Mts.), Nev. (Charleston Mts.), Ariz.
(Grand Canyon, south rim). Pollen: Unknown, but visits flowers of Agave deserti,

Cowania stansburiana, Eriodictyon.
Perdita inornata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 49, figs. 787, 788, 872. 2, ¢.

jucunda Timberlake. Calif., Nev. (Churchill County), Ariz. (Painted Desert). Pollen: Collects
pollen from the flowers of Eriogonwm, including E. aureum, E. fasciculatum, E. gracile,
E. inflatum, E. molestum var. davidsonii, E. nudum, but also visits other flowers for
nectar including Chorizanthe staticoides, Gutierrezia lucida, Lepidospartum
squamatum.
Perdita jucunda Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 64, figs. 803, 804, 880. 2, ¢.
kanabensis Timberlake. Utah (Kanab), Ariz. (Coconino County). Pollen: Unknown, but visits
flowers of cleomella, Eriogonum corymbosum.
Perdita kanabensis Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 34, figs. 1383, 1384, 1410.
labrata Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Evidently collects
pollen from the flowers of Eriogonwm including E. fasciculatum var. polifolium, but
also visits other flowers presumably for nectar including Larrea tridentata.
Perdita labrata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 81, figs. 821, 822, 889, 901. 2,
é.
lateralis daleae Timberlake. Calif. (Inyo and Panamint Mts.). Pollen: Apparently an oligolege
of Larrea tridentata, but visits other flowers presumably for nectar including
Chaenactis, D. fremontii, D. f. var. johnsonii, Eriogonum fasciculatum, E. inflatum.
Perdita lateralis daleae Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 29. 2, 6.
lateralis lateralis Timberlake. Calif. (Colorado Desert), Ariz. (Maricopa, Pinal and Yuma
Counties). Pollen: Apparently an oligolege of Larrea tridentata, but also visits other
flowers presumably for nectar including Chaenactis, Dalea californica, Eriogonum
fasciculatum, Hyptis emoryi, Prosopis juliflora.
Perdita lateralis lateralis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 27, figs. 749, 750,
853. 2, od.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 25 (floral relationships).

Superfamily APOIDEA 1909

lucens Timberlake. South. Calif. (Mojave Desert), Nev. (Churchill County). Pollen: Possibly an
oligolege of Eriogonum, visits flowers of E. fasciculatum, E. inflatum, E. mohavense,
E. nodosum.
Perdita lucens Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 71, figs. 809, 810, 883. 2, ¢.
lunulata Timberlake. Ariz. (Apache and Navajo Counties). Pollen: Collects pollen from the
flowers of Mentzelia pumila, but also visits the flowers of Eriogonwm aureum
presumably for nectar.
Perdita lunulata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 22. 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 96, 101. 2, ¢ (tax. status, key,
geogr. and floral relationships).
melanops Timberlake. South. Calif. (Santa Rosa Mts.). Pollen: Unknown, but visits flowers of
Physalis crassifolia.
Perdita melanops Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 105. 2.
melanura Timberlake. Ariz. (Grand Canyon).
Perdita melanura Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 52. °.
mentzeliae Cockerell. Colo. (Antonita and Trinidad), N. Mex., Ariz., (Cochise, Greenlee and
Navajo Counties). Pollen: Collects pollen from the flowers of Mentzelia pumila.
Perdita mentzeliae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 60. d, °.
Perdita pallidior Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 61. d, @.
Perdita pulchrior Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 62. 6.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 3, 10, 24-26, figs. 745, 746, 851. 2, 3
(key, redescription, synonymy, geogr. and floral records). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 101 (geogr. and floral records).

Biology: Linsley and Hurd, 1959. Ent. News 70: 64, 65, 67 (floral relationships).
mentzeliarum Cockerell. West. Tex. to Ariz.; Mexico (Baja California and Chihuahua). Pollen:
Apparently an oligolege of Mentzelia pumila, but visits flowers of Bahia absinthifolia

for nectar.
Perdita mentzeliarum Cockerell, 1897. N. Mex. Agr. Expt. Sta., Bul. 24: 43. 2, d.
Perdita mentzeliarum var. lauta Cockerell, 1905. Entomologist 38: 145. 9, d.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 3, 9, 23-24, figs. 748, 744, 850. 2, d
(key, redescription, synonymy, geogr. and floral records). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 101 (geogr. and floral records).

michelbacheri Timberlake. Ariz. (Maricopa County); Mexico (Baja California). Pollen:
Unknown, but visits flowers of Acacia, Carnegiea gigantea.
Perdita michelbacheri Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 60, figs. 797, 798, 877.
OFTOL-
nasuta galacticoptera Timberlake. Calif. (Inyo and Panamint Mts.). Pollen: Apparently an
oligolege of Eriogonum, visits flowers of E. inflatum.
Perdita nasuta galacticoptera Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 76. 2, d.
nasuta nasuta Timberlake. South. Calif. (Colorado and Mojave Deserts), Nev. (Clark County),
Ariz. (Mohave, Pinal and Yuma Counties). Pollen: Apparently an oligolege of
Eriogonum, visits flowers of E. trichopes.
Perdita nasuta nasuta Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 74, figs. 813, 814, 885.
ORT.
nasuta obscurescens Timberlake. Ariz., N. Mex. (Las Cruces). Pollen: Apparently an oligolege
of Eriogonum, visits flowers of E. thomasii, E. trichopes.
Perdita nasuta obscurescens Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 76. 2, d.
nigridia Timberlake. Calif. (Colorado and Mojave Deserts), Ariz. (Mohave County); Mexico
(Baja California). Pollen: Apparently collects pollen only from the flowers of Mentzelia
including M. albicaulis, M. involucrata, M. pumila, M. tricuspis, but visits these and
apparently other flowers for nectar including Encelia farinosa, Eriogonum

fasciculatum, Eucnide urens. :
Perdita nigridia Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 18, figs. 777, 778, 867. d, °.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 101 (geogr. and floral record).

1910 Hymenoptera in America North of Mexico

nodosicornia Timberlake. Calif. (Coalinga).
Perdita nodosicornia Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 110, fig. 1360. ¢.
pectoralis Timberlake. Ariz. (Vermillion Cliffs). Pollen: Unknown, but visits flowers of
Eriogonum.
Perdita pectoralis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 111, figs. 1265, 1266, 1336.
3.
perplexa Timberlake. N. Mex. (Catron County). Pollen: Unknown, but visits flowers of
Mentzelia.
Perdita perplexa Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 19, figs. 739, 740, 848. 2 (¢
misdet.).

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 101 (tax. status).
polytropica obsoleta Timberlake. Ariz. (Maricopa, Pima and Pinal Counties). Pollen: Polylectic,
visits flowers of Acacia, Carnegiea gigantea, Cercidium, Echinocactus.
Perdita polytropica obsoleta Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 59. 2, ¢.
polytropica polytropica Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen:
Polylectic, visits a wide variety of flowers including Acacia greggti, Agave deserti, _
Asclepias subulata, Carnegiea gigantea, Croton californicus, Dalea spinosa, Encelia
farinosa, Echinocactus, Eriogonum inflatum, Ferrocactus acanthodes, Hyptis emoryi,
Larrea tridentata, Olneya tesota, Opuntia, Prosopis juliflora.
Perdita polytropica polytropica Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 57, figs. 795,
TSB, GH. 5 Cie
punctifera Cockerell. Ariz., south. Calif. (Blythe); Mexico (Baja California). Pollen: Collects
pollen from the flowers of Mentzelia pumila, but also visits the flowers of Gossypium
thurberi presumably for nectar.
Perdita punctifera Cockerell, 1914. Ent. Soc. Wash., Proc. 16: 32. 9.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 3, 9, 26-27, figs. 747, 748, 852. 2, ¢
(key, redescription, geogr. and floral records).
rehni praeclara Timberlake. Ariz. (Southwestern Research Station).
Perdita rehni praeclara Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 106, fig. 14. °.
rehni rehni Cockerell. N. Mex., western Tex.; Mexico (Chihuahua). Pollen: Unknown, but visits

flowers of Dasylirion wheeleri, Nolina.
Perdita Rehni Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 131. 2 (= male).

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 8, 12, 50-52, figs. 789, 790, 873. 2, ¢
(key, redescription, geogr. and floral records).
rhois reducta Cockerell. Calif. (South Coast Ranges). Pollen: Apparently polylectic, visits a
wide variety of flowers including Adenostoma fasciculatum, Eriodictyon parryi,
Eriogonum fasciculatum, Heteromeles arbutifolia, Rhus lawrina.
Perdita rhois mut. reducta Cockerell, 1901. Canad. Ent. 33: 283. °.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 6, 54-55. 2, d (key, redescription,
geogr. and floral records).
rhois rhois Cockerell. South. Calif. (chiefly cismontane); Mexico (Baja California). Pollen:

Polylectic, visits a wide variety of flowers for pollen and nectar including Acacia
greggut, Adenostoma fasciculatum, A. sparsifolium, Agave deserti, Alyssum
maritimum, Baccharis emoryi, B. viminea, Chrysanthemum, Eriogonum fasciculatum,
E. f. var. polifolium, E. gracile, Euphorbia albomarginata, Gutierrezia californica,
Haplopappus linearifolius, Heliotropium oculatum, Heteromeles arbutifolia, Lotus
scoparius, Mentha, Nolina parryi, Polygonum lapanthifolium, Prunus ilicifolia,
Rhamnus californicus, Rhus laurina, Rhus ovata, Schinus molle.

Perdita rhois Cockerell, 1901. Canad. Ent. 33: 282. 9.

Perdita hypoxantha Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 13: 424. ¢.

Taxonomy: Timberlake, 1951. Jn Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr.
Monog. 2: 1094 (synonymy). —Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 6, 12, 53-54,
figs. 791, 792, 874. 2, 6 (key, redescription, geogr. and floral records).

Superfamily APOIDEA 1911

semicrocea Cockerell. N. Mex. (Dona Ana, Otero and San Miguel Counties), Ariz. (Willcox).
Pollen: Unknown, but visits flowers of Aster, Bahia absinthifolia, Gutierrezia sarothrae,
Haplopappus, Solidago.
Perdita semicrocea Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 13. 9.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 4, 9, 35-36, figs. 761, 762, 859. 2, 6
(key, redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent.
49: 96, 101 (key, geogr. and floral record).
semilutea Timberlake. Calif. (Box Canyon in Riverside County). Pollen: Unknown, but, visits
flowers of Eriogonum thomasii.

Perdita semilutea Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 81, figs. 819, 820, 888. 3.
snellingi Timberlake. South. Calif. (Kern and Inyo Counties). Pollen: Presumably an oligolege
of vernal and autumnal flowering Compositae, visits flowers of Chrysothamnus

nauseosus consimilis, Cleome, Eriogonum heermannii, Euphorbia albomarginata,
Gutierrezia lucida, G. microcephala, Rosa californica, Solidago spectabilis.
Perdita snellingi Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 46, figs. 781, 782, 869. 2, 3.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 106-107 (tax. characters, geogr. and
floral records).
sodalis Timberlake. South. Calif. (Santa Rosa Mts.). Pollen: Unknown, but visits flowers of
Hyptis emoryi.
Perdita sodalis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 101, figs. 1255, 1256, 1331. ¢.
stabilis Timberlake. Ariz. (Douglas).
Perdita stabilis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 102, figs. 1257, 1258, 1332. d.

subfasciata Cockerell. N. Mex. Ariz., Utah (Juab County), Nev. (White Pine County), Calif.
(Inyo County Mts.). Pollen: Apparently an oligolege of vernal and autumnal flowering
Compositae, visits flowers of Chrysothamnus nauseosus, C. viscidiflorus, Gutierrezia
lucida, G. microcephala.
Perdita subfasciata Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 512. @.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 4, 11, 45-46, figs. 779, 780, 868. 2, 3
(key, redescription, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent.
49: 107 (geogr. and floral records).
thermophila thermophila Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen:
Possibly an oligolege of Eriogonum, visits flowers of E. deserticola, E. hirtella, E.
inflatum, E. reniforme, E. thomasii.
Perdita thermophila thermophila Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 77, figs.
815, 816, 886. 2, d.
thermophila trilobata Timberlake. Ariz., south. Calif. (Mojave Desert). Pollen: Possibly an
oligolege of Eriogonum, visits flowers of Baileya, Eriogonum deflexum, E. densum, E.
deserticola, E. trichopes.
Perdita thermophila trilobata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 78. 2, d.
timberlakei Cockerell. South. Calif. (cismontane and adjacent desert areas). Pollen: Evidently
depends mostly upon the pollen and nectar of Eriogonum gracile, but also visits flowers
of Brassica, Eriogonum fasciculatum, E. mohavense, E. virgatum.
Perdita timberlakei Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 194. 9, 6.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 6, 13, 70-71, figs. 807, 808, 882. 2, 3
(key, redescription, geogr. and floral records).
varleyi niveipennis Timberlake. Calif. (San Benito County). Pollen: Apparently an oligolege of
Eriogonum including EL. inflatum.
Perdita varleyi niveipennis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 86. 2, d.
varleyi varleyi Timberlake. South. Calif. (Mojave Desert, western margin). Pollen: Apparently
an oligolege of Eriogonum, visits flowers of E. fasciculatum var. polifolium, E.
inflatum.
Perdita varleyi varleyi Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 85, figs. 825, 826, 891,
904. 3, @.

1912 Hymenoptera in America North of Mexico

vicina Timberlake. Nev. (Charleston Mts.). Pollen: Unknown, but visits flowers of Fallugia
paradoxa.
Perdita vicina Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 31, figs. 753, 754, 855. 6.
viridinotata Timberlake. N. Mex. (Alamogordo).
Perdita viridinotata Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 16, figs. 735, 736, 846. °,
3.
wheeleri Timberlake. Ariz. (Grand Canyon). Pollen: Unknown, but possibly may visit flowers
of Mentzelia.
Perdita wheeleri Timberlake, 1928. Amer. Mus. Novitates 321: 5. o.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 10, 32, figs. 755, 756, 856. 3
(redescription, possible pollen source).
wootonae Cockerell. N. Mex., Colo., Kans., Nebr. Pollen: Apparently an oligolege of Mentzelia
including M. decapetala, M. nuda, but visits other flowers presumably for nectar
including Eriogonum effusum, Tragopogon porrifolius.
Perdita wootonae Cockerell, 1898. Ent. News 9: 215. 9, 6.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 2, 8, 15-16, figs. 733, 734, 845. 9, 6
(key, redescription, geogr. and floral records).
xanthoxyli Timberlake. Tex. (Corpus Christi State Park); Mexico (Puebla). Pollen: Unknown,
but visits flowers of Xanthoxylon.
Perdita xanthoxyli Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 107, figs. 15, 1261, 1262,
1334. 2, 3.
xerophila discrepans Timberlake. Nev. (Clark County). Pollen: Possibly an oligolege of
Eriogonum, visits flowers of E. fasciculatum, E. trichopes.
Perdita xerophila discrepans Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 84, figs. 823,
824, 890. 2, d.
xerophila fuscicornis Timberlake. Calif. (Inyo County Mts.). Pollen: Possibly an oligolege of
Eriogonum, visits flowers of E. inflatum, Larrea tridentata.
Perdita xerophila fuscicornis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 84, fig. 903. 2,
é.

xerophila xerophila Timberlake. Ariz. (Yuma County), et Calif. (Colorado Desert). Pollen:
Possibly an oligolege of Eriogonum, visits flowers of E. inflatum, E. trichopes.
Perdita xerophila xerophila Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 82, fig. 902. 2, 3.
yosemitensis Timberlake. Calif. (Sierra Nevada Mts.). Pollen: Possibly an oligolege of
Eriogonum, visits flowers of E. nudum, E. wrightii.
Perdita yosemitensis Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 63, figs. 801, 802, 879.
Ose

SPECIES GROUP SPHAERALCEAE

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 1 (key). —Timberlake, 1964. Calif.
Univ. Pubs. Ent. 28: 126-158 (key to included spp.). —Timberlake, 1971. Calif. Univ. Pubs.
Ent. 66: 38-42 (suppl. key to included spp.).

adustiventris Timberlake. South. Calif. (Parker Dam); Mexico (Baja California: Angel de
la Guardia Island). Pollen: Unknown, but visits flowers of Eucnide urens, Mentzelia

hirsutissima.
Perdita adustiventris Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 166. @.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 42-43, figs. 5, 1885, 1386, 1411. 3
(geogr. and floral records).
albiventris Timberlake. Tex. (Pecos County). Pollen: Unknown, but the type specimen bears a
load of fine, mealy fulvous pollen, probably leguminaceous.
Perdita albiventris Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 294, fig. 60. ?.
albofasciata Timberlake. N. Mex. (Santa Fe). Pollen: Unknown, but visits flowers of Tamarix

gallica.
Perdita albofasciata Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 152. 9.

Superfamily APOIDEA 1913

ambigua Timberlake. Nev. (Lander and Washoe Counties), Calif. (Siskiyou County). Pollen:
Unknown, but visits flowers of Chrysothamnus nauseosus var. consimilis, Cirsium
lanceolatum.
Perdita ambigua Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 311. 2.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 54-55. ¢ (geogr. and floral records).
a naar Ariz. (Willcox). Pollen: Unknown, but visits flowers of Aplopappus
artwegi.

Perdita nish Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 295. 9.
ancoralis Timberlake. South. Calif. (Boyd Desert Research Center).

Perdita ancoralis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 155. 9.
ashmeadi ashmeadi Cockerell. Tex. (Brownsville); Mexico (Sonora). Pollen: Unknown, but

visits flowers of Prosopis. :
Perdita ashmeadi Cockerell, 1899. Ann. and Mag. Nat. Hist. (7) 8: 492. 2.
Perdita exclamans atramentata Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 96. ¢.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 141, 158, 324-325, figs. 1069, 1070,
1178. 2, ¢ (key, redescription, synonymy, geogr. and floral records).
ashmeadi simulans Timberlake. South. Calif. (Colorado Desert), ?Ariz. (Sentinel); Mexico
(Baja California). Pollen: Unknown, but visits flowers of Geraea canescens, Larrea
tridentata, Prosopis juliflora.
Perdita ashmeadi simulans Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 326, figs. 1071,
LOTZIIUTO TS.
ashmeadi vierecki Cockerell. West. Tex., N. Mex. (Alamogordo), Ariz. Pollen: Unknown, but
visits flowers of Mimosa, Prosopis.
Perdita Vierecki Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 129. 9, 6.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 141, 158, 325-326. 2, d (key,
redescription, tax. status, geogr. and floral records).
assimilis Timberlake. Calif. (Fresno County). Pollen: Unknown, but visits flowers of Melilotus.

Perdita assimilis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 228, figs. 37, 987, 988, 1137.
oe
atrata Timberlake. Calif. (Colorado and Mojave Deserts). Pollen: Unknown, but visits flowers
of Camissonia brevipes, C. dentata var. parishii, Chaenactis stevioides, Mentzelia
affinis, M. albicaulis, M. veatchiana, Sphaeralcea ambigua.
Perdita atrata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 163, figs. 3, 911, 912, 1009. °,
é.
barri Timberlake. Idaho (Midvale). Pollen: Unknown, but visits flowers of Phacelia.
Perdita barri Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 251, figs. 1018, 1014, 1150. ¢.

blanda Timberlake. N. Mex. (Albuquerque).
Perdita blanda Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 358. 2.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 137, 262 (key, tax. status).
calloleuca calloleuca Cockerell. N. Mex., Colo., Utah, Ariz. Pollen: Unknown, but visits
flowers of Artemisia filifolia, Chrysothamnus, Cleome lutea, Eriogonum, Salsola kali,
Tamarix, Trifolium, Wislizenia refracta.
Perdita calloleuca Cockerell, 1922. Amer. Mus. Novitates 33: 1220On
Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 140, 154, 298-300, figs. 63, 1059,
1060, 1173. 2, ¢ (key, redescription, geogr. and floral records). —Timberlake, 1966. Calif.
Univ. Pubs. Ent. 66: 43 (geogr. and floral records).
calloleuca convergens Timberlake. N. Mex. (Virden), Ariz. (Graham and Yuma Counties), Nev.
(Clark County). Pollen: Unknown, but visits flowers of Atriplex canescens,
Chenopodium oblongifolium, Euphorbia, Suaeda, Wislizenia refracta.
Perdita calloleuca convergens Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 300. 2, ¢.
chihuahua Timberlake. N. Mex. (Rodeo); Mexico (Chihuahua and Zacatecas).
Perdita chihuahua Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 203, figs. 955, 956, 1121.
OF Gi

1914 Hymenoptera in America North of Mexico

chionostoma Timberlake. Calif. (Fresno County). Pollen: Unknown, but visits flowers of
Melilotus.
Perdita chionostoma Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 240, figs. 43, 1001, 1002,
1144. 9, 6.
cleomellae Cockerell. Calif. (Mojave Desert). Pollen: Apparently an oligolege of Cleomella
including C. obtusifolia.
Perdita cleomellae Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 193. 2, 6.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 309, 315, 353-354, figs. 430, 431, 499.
2, 3 (key, redescription, geogr. and floral records).
cochiseana Timberlake. Ariz. (Douglas). Pollen: Unknown, but visits flowers of Euphorbia.
Perdita cochiseana Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 43. °.
compta Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Unknown, but most
likely source is Eriastrum; visits flowers of Cryptantha intermedia, Eriastrum
virgatum, Eriogonum reniforme, E. trichopes, Lepidium fremontii, Malacothrix
glabrata, Stillingia paucidentata.
Perdita compta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 236, figs. 41, 997, 998, 1142.
25 Sc
confusa Timberlake. West. Tex., N. Mex., Ariz.; Mexico (Chihuahua). Pollen: Apparently an
oligolege of Lepidiwm including L. alyssoides, L. montanum, L. thurberi.
Perdita confusa Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 247, figs. 46, 1007, 1008, 1147.
REG

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2259: 19, fig. 50 (larva, as questionably this
species).

Biology: Rozen, 1967. Amer. Mus. Novitates 2297: 38, fig. 15, table 1 (nest architecture, life
history, floral association).
covilleae Timberlake. Ariz., Utah (St. George), Nev. (Las Vegas), south. Calif. (Colorado and
Mojave Deserts); Mexico (Baja California). Pollen: Oligolege of Larrea tridentata, also
visits other flowers for nectar including Heliotropium curassavicum, Prosopis.
Perdita covilleae Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 361, figs. 438, 489, 503. 9,
3.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 139, 262 (key).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 24, fig. 5 (intrafloral ecology).
cruciferarum Timberlake. Nev. (Battle Mountain). Pollen: Unknown, but visits flowers of an
unidentified Cruciferae.
Perdita cruciferarum Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 140, figs. 24, 25, 1291,
1292, 1349. 9, 6.
cushmani Timberlake. Tex. (Brewster and Hudspeth Counties); Mexico (Chihuahua). Pollen:
Unknown, but visits flowers of Gutierrezia, Prosopis juliflora.
Perdita cushmani Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 222, figs. 979, 980, 1133. °,
3.
cuspidata Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Collects pollen
from the flowers of Nama demissum, Phacelia distans, but visits other flowers for

nectar including Oenothera.
Perdita cuspidata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 231, figs. 39, 991, 992, 1139.
OMOE
dammersi Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Collects pollen
from flowers of Malacothrix including M. glabrata, but visits these and other flowers
for nectar including Chaenactis fremontii, Stephanomeria exigua.
Perdita dammersi Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 169, figs. 7, 919, 920, 1103.
OR
davidsoni Timberlake. South. Calif.
Perdita davidsoni Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 235, figs. 995, 996, 1141. ¢.

Superfamily APOIDEA 1915

deltophora Timberlake. Tex. Pollen: Unknown, but visists flowers of Chamaesaracha
conioides, Physalis lobata, Prosopis.
Perdita deltophora Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 207, figs. 25, 959, 960,
23 ee
dentata Timberlake. South. Calif. (Colorado Desert). Pollen: Possibly an oligolege of Phacelia
including P. distans, but visits other flowers presumably for nectar including Acacia
greggui, Geraea canescens, Krameria canescens, Larrea tridentata.
Perdita dentata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 165, figs. 4, 913, 914, 1100. °,
3

difficilis Timberlake. Tex. (Brewster County), N. Mex. (Dona Ana and Eddy Counties), Ariz.,
south. Calif. (Colorado and Mojave Deserts); Mexico (Baja California). Pollen:
Apparently an oligolege of Prosopis including P. juliflora, but visits flowers of other
desert shrubs presumably for nectar including Cercidium floridum, Larrea tridentata,
Mimosa and also has been collected at the flowers of Melilotus.
Perdita difficilis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 323, figs. 1067, 1068, 1177.
digna Timberlake. Calif. (Quincy).
Perdita digna Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 310. °.
discors Timberlake. South. Calif. (Thousand Palms).
Perdita discors Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 361. °.
dispar Timberlake. Tex. (Starr and Webb Counties); Mexico (Chihuahua). Pollen: Unknown,
but visits flowers of Condalia lycioides, Croton, Prosopis juliflora.
Perdita dispar Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 220, figs. 33, 997, 998, 1132. 2,
Sr

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 43: 136 (geogr. and floral record).
distincta Timberlake. Oreg. (Baker County), Calif. (Sierra County). Pollen: Unknown, but

visits flowers of Eriogonum.
Perdita distincta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 174, figs. 10, 925, 926, 1106.

OE
Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 45-46. 9.

drymariae Timberlake. N. Mex. (Dona Ana and Hidalgo Counties), Mich. (Baraga County);
Mexico (San Luis Potosi). Pollen: Unknown, but visits flowers of Drymaria holosteoides,

Tidestromia lanuginosa.
Perdita drymariae Timberlake, 1960. Jn Mitchell, N. C. Agr. Expt. Sta. Tech. Bul. 141: 324.

ORIGe ;

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 129, 148, 201-202, figs. 22, 951, 952,
1119. 2, 3 (key, tax. characters, geogr. and floral records). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 112, 134 (key, geogr. and floral record).

emarginata Timberlake. Ariz. (Tucson), south. Calif. (Desert Center). Pollen: Unknown, but
visits flowers of Dalea.
Perdita emarginata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 304. 2.
eremica Timberlake. Ariz. (Maricopa and Pinal Counties), south. Calif. (Colorado and Mojave
Deserts). Pollen: Apparently an oligolege of Larrea tridentata, but visits other flowers
including Dalea californica, D. fremontii var. johnsonii, D. f. var. saundersii, D. schottii,
Eriogonum inflatum, Funastrum hirtellum, Hyptis emoryi, Nama demissum.
Perdita eremica Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 268, figs. 54, 1033, 1034, 1160.
OeOre
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 25 (floral relationships).
eremophila Timberlake. South. Calif. (Mojave Desert). Pollen: Possibly an oligolege of
Phacelia including P. distans, P. fremontii, but visits other flowers including
Haplopappus cooperi, Lepidium fremontii.
Perdita eremophila Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 238, figs. 42, 999, 1000,
1143. 2, 6.

1916 Hymenoptera in America North of Mexico

eriastri Timberlake. South. Calif. (cismontane). Pollen: Collects pollen from the flowers of
Eriastrum including E. virgatum, but visits other flowers for nectar including
Cryptantha intermedia, Eriogonum fasciculatum, Gilia.
Perdita eriastri Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 233, figs. 40, 993, 994, 1140.
2,6.
eriastri var. fusciventris Timberlake. South. Calif. (Palm Springs). Pollen: Collects pollen from
flowers of Eriastrum virgatum. A specimen similar to the holotype has been collected
in Arroyo Seco, Los Angeles County at the flowers of Eriodictyon parryi.
Perdita eriastri var. fusciventris Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 235. 9.
eucnides eucnides Timberlake. Calif. (Inyo County), Nev. (Churchill County). Pollen:
Polylectic, collects pollen from the flowers of Cowania mexicana var. stansburiana,
Dalea polyadenia, Eucnide wrens, Larrea tridentata, but also visits other flowers
including Eriogonum inflatum possibly for nectar.
Perdita eucnides eucnides Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 301, figs. 64, 1061,
1062, 1174. 6, 9.
eucnides platyzona Timberlake. Ariz. (Yuma County), south. Calif. (Colorado Desert); Mexico
(Baja California). Pollen: Polylectic, visits flowers of Acacia greggii, Larrea tridentata,
Pluchea sericea, Sesuvium verrucosum.
Perdita eucnides platyzona Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 304. 2, ¢.
euzonata Timberlake. Ariz. (Painted Desert). Pollen: Unknown, but visits flowers of
Eriogonum aureum.
Perdita euzonata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 295. 2.
exclamans Cockerell. West. Tex. to south. Calif. (Colorado and Mojave Deserts) and south
Nev. (Clark County); Mexico (Baja California and Sonora). Pollen: Collects pollen from
the flowers of Prosopis, but also visits other flowers presumably for nectar including
Baccharis, Cercidium floridum, Dithyrea wislizeni, Larrea tridentata, Melilotus,
Mimosa, Phoradendron.
Perdita nitidella var. ecclamans Cockerell, 1895. Psyche 1 (sup.): 5. 6.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 141, 157, 158, 320-3238, figs. 70, 1065,
1066, 1176. 5, 2 (key, redescription, geogr. and floral records).
exigua becki Timberlake. Nev. (Nye and White Pine Counties).
Perdita exigua becki Timberlake, 1971 Calif. Univ. Pubs. Ent. 66: 44. 2, 6.
exigua exigua Timberlake. Calif. (Hallelujah Junction).
Perdita exigua Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 278, figs. 1043, 1044, 1165. °,
3.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 44 (tax. status).

eximia Timberlake. Ariz. (Maricopa and Yuma Counties), south. Calif. (Colorado and Mojave
Deserts), Nev. (Esmeralda County). Pollen: Possibly an oligolege of Phacelia including
P. crenulata, but visits a variety of other flowers including Acacia greggii, Camissonia
clavaeformis, Cryptantha angustifolia, Cercidium floridum, Dalea schottu, Encelia,
Eriogonum inflatum, E. thomasii, Heliotropium curassavicum, Isomeris arborea,
Larrea tridentata, Prosopis juliflora.

Perdita eximia Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 255, figs. 49, 1019, 1020, 1153.

Onn

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 145 (key).
eysenhardtiae Timberlake. Tex. (Alpine), Ariz. (Cochise and Santa Cruz Counties); Mexico
(Distrito Federal, Durango and Hidalgo). Pollen: Unknown, but visits flowers of
Baccharis glutinosa, Eysenhardtia polystachya.
Perdita eysenhardtiae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 215. @.
falcata Timberlake. Calif. (Inyo County). Pollen: Unknown, but visits flowers of Dalea
fremontii, Encelia farinosa, Gilia, Sphaeralcea.
Perdita falcata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 161, figs. 2, 909, 910, 1098. °,
é.

Superfamily APOIDEA 1917

flavipes Timberlake. South. Calif. (Colorado and Mojave Deserts); Mexico (Baja California).
Pollen: Apparently an oligolege of Larrea tridentata, but visits other flowers including
Heliotropium curassavicum, Prosopis juliflora.
Rae flavipes Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 259, figs. 50, 1023, 1024, 1155.
Opes

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 24 (floral relationships).
florissantella Cockerell. Colo., N. Mex. (Taos County). Pollen: Collects pollen from the flowers
of Lepidium jonesi, but visits other flowers presumably for nectar including
Chenopodium album, Eriogonum umbellatum, Phacelia.
Perdita florissantella Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 440. ¢, 9.
Perdita lepidii Cockerell, 1907. Entomologist 40: 266. °.
Perdita opacifrons Timberlake, 1929. N. Y. Ent. Soc., Jour. 37: 113. 6.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 137, 151, 242-244, figs. 44, 1003,
1004, 1145. 2, 3 (key, redescription, synonymy, geogr. and floral records). —Timberlake,
1968. Calif. Univ. Pubs. Ent. 49: 139 (key).

foxi Cockerell. N. Mex. (Santa Fe).
Perdita foxi Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 18. 3.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 152, 317. 6 (key, redescription).
gemella Timberlake. Calif. (Apple Valley and Deep Creek on Mojave Desert). Pollen:
Unknown, but visits flowers of Eriodictyon trichocalyx.
Perdita gemella Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 200, figs. 949, 950, 1118. ¢.
geminata Timberlake. N. Mex. (Otero County). Pollen: Unknown, but visits flowers of Larrea
tridentata.
Perdita geminata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 251, figs. 1015, 1016, 1151.
ONor
genalis genalis Timberlake. South. Calif. (Colorado and Mojave Deserts). Pollen: Unknown,
but visits flowers of Acacia greggii, Cercidium floridum, Larrea tridentata, Prosopis
juliflora.
Perdita genalis genalis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 359, figs. 76, 1093,
1094, 1190. 2, 6.
genalis panamintensis Timberlake. Calif. (Panamint Mts.). Pollen: Unknown, but visits flowers
of Larrea tridentata, Prosopis juliflora, Stanleya pinnata.
Perdita genalis panamintensis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 360. 2, ¢.
gracilis Timberlake. Calif. (Lone Pine). Pollen: Unknown, but visits flowers of Malacothriz.
Perdita gracilis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 309. °.
greggiae Timberlake. Tex. (Reeves County), N. Mex. (Eddy County). Pollen: Unknown, but
visits flowers of Lepidium montanum, Lesquerella gordonii, Nerisyrenia camporum.
Perdita greggiae Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 142, figs. 1293, 1294, 1350. °,
3.

heliotropii heliotropii Cockerell. Tex. (E] Paso) west to south. Calif. (Colorado and Mojave
Deserts), Nev. (Lyon County), Utah (St. George).; Mexico (Baja California, Sonora,
Chihuahua). Pollen: Collects pollen from the flowers of Heliotropium including H.
curassavicum, but visits these and other flowers for nectar including Larrea tridentata,
Salix, Tamarix.
Perdita heliotropii Cockerell, 1900. Entomologist 33: 63. °, d.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 309, 314, 364-365, figs. 442, 448, 505,
521. 9, 6 (key, tax. characters, tax. status, geogr. and floral records). —Timberlake, 1964.
Calif. Univ. Pubs. Ent. 28: 139, 262 (key, tax. characters). —Timberlake, 1968. Calif. Univ.
Pubs. Ent. 49: 116, 148 (key, tax. characters, geogr. record).

heliotropii perducta Timberlake. Central Calif. Pollen: Collects pollen from the flowers of
Heliotropium curassavicum, but also visits flowers of Stanleya pinnata presumably for

nectar.
Perdita heliotropii perducta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 365. 2, 6.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 139, 262 (key, geogr. and floral
record).

1918 Hymenoptera in America North of Mexico

humilis Timberlake. N. Mex. (Mesilla Park). Pollen: Unknown, but visits flowers of Dithyrea
wislizent.
Perdita humilis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 156. °.
imberbis Timberlake. Calif. (Sierra and Siskiyou Counties). Pollen: Unknown, but visits

flowers of Erigeron.
Perdita imberbis Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 139, fig. 23. 2.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 46-47, figs. 1387, 1388, 1412. 6, 2
(key, geogr. and floral record).
impigra Timberlake. Tex. (San Jacinto County).
Perdita impigra Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 157, fig. 31. 2.
impressa Timberlake. Ariz. (Apache County). Pollen: Unknown, but visits flowers of
Eriogonum aureum.
Perdita impressa Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 1538. 9.
incompta Timberlake. Tex. (Southmost). Pollen: Unknown, but visits flowers of Lippia.
Perdita incompta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 208, figs. 26, 961, 962, 1124.
ORE
infelix Timberlake. Nebr. (Harrison).
Perdita infelix Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 314. °.
innotata Timberlake. Ariz. (Maricopa and Yuma Counties), south. Calif. (Colorado and Mojave
Deserts). Pollen: Possibly an oligolege of Prosopis including P. juliflora, but also visits
flowers of Nama hispidum.
Perdita innotata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 333, figs. 1081, 1082, 1184.
5 Gc
insequens Timberlake. South. Calif. (Piute Butte in Mojave Desert). Pollen: Unknown, but
visits flowers of Chaenactis brachypappa.
Perdita insequens Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 146, fig. 27. @.
koebelei concinna Timberlake. South. Calif. (Colorado and Mojave Deserts), Ariz. (Mohave
and Yuma Counties). Pollen: Apparently an oligolege of Mentzelia including M.
imvolucrata, M. tricuspis, but also visits flowers of Eucnide urens, Hyptis emoryi.
Perdita koebelei concinna Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 161, figs. 907, 908.
Onn
Perdita masoni Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 306. 9°.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 47 (synonymy, geogr. and floral
records).
koebelei koebelei Timberlake. Calif. (Inyo County). Pollen: Collects pollen from the flowers of
Eucnide urens, Mentzelia, but also visits other flowers presumably for nectar including
Argemone platyceras, Encelia farinosa, Eriogonum inflatum, Phacelia calthifolia,
Stephanomeria.
Perdita koebelei koebelei Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 159, figs. 1, 905, 906,
LOS fencien or
krombeini Timberlake. Fla. (Lee County).
Perdita krombeini Timberlake, 1960. In Mitchell, N. C. Agr. Expt. Sta. Tech. Bul. 141: 327.
3.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 144, 167, figs. 915, 916, 1101. ¢ (key,
tax. characters).
leucogastra Timberlake. Tex. (Terrell County). Pollen: Unknown, but visits flowers of Gilia
acerosa.
Perdita leucogastra Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 280, figs. 1045, 1046,
1166. 2, 6d.
leucosticta Timberlake. South. Calif. (San Jacinto Mts. and environs). Pollen: Possibly an
oligolege of Eriastrum virgatum, but visits other flowers including Eriogonum
fasciculatum, Gilia exilis, Layia platyglossa.
Perdita leucosticta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 224, figs. 34, 981, 982,
Sa CAN)

Superfamily APOIDEA 1919

linsleyi Timberlake. South. Calif. (Los Angeles and Ventura Counties). Pollen: Unknown, but
visits flowers of Chaenactis, Euphorbia albomarginata, Lasthenia chrysostoma.
Perdita linsleyi Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 199, figs. 21, 947, 948, 1117.
OOr
luciae decora Timberlake. Ariz. (Maricopa, Pima and Yuma Counties), Nev. (Clark County),
south. Calif. (Colorado Desert); Mexico (Baja California and Sonora). Pollen: Polylectic,
especially flowers of Larrea tridentata and Prosopis juliflora, but also visits a wide
variety of other flowers including Acacia greggii, Baileya pleniradiata, Cercidium
floridum, Cryptantha angustifolia, C. intermedia, Nama hispidum, Psilostrophe
cooperi, Sphaeralcea emoryi.
Perdita luciae decora Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 336, fig. 73. 2, d.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 25 (floral relationships).
luciae luciae Cockerell. Tex. (Pecos County), N. Mex. (Las Cruces), Ariz. (Cochise, Maricopa
and Pima Counties); Mexico (Sonora). Pollen: Unknown, but visits flowers of Medicago
sativa, Mimosa, Prosopis.
Perdita luciae Cockerell, 1899. Ann. and Mag. Nat. Hist. (7) 3: 494. d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 141, 156, 157, 334-336, figs. 72, 1083,
1084, 1185 (key, tax. status, redescription, geogr. and floral records).
lycii Timberlake. Ariz. (Pima County). Pollen: Unknown, but visits flowers of Lyciwm.
Perdita lycii Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 305, fig. 65. 2.
macneilli Timberlake. Calif. (Santa Clara County).
Perdita macneilli Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 312. °.
macswaini Timberlake. Tex. (Cameron and Harligen Counties), N. Mex. (Eddy County).
Pollen: Unknown, but visits flowers of Coreopsis douglasii, Gaillardia, Monarda
citriodora, Prosopis.
Perdita macswaini Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 289, figs. 1053, 1054, 1170.
25 We
macswaini var. flavolineata Timberlake. Tex. (Hidalgo County).
Perdita macswaini var. flavolineata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 290. d.
martini Cockerell. West. Tex., N. Mex. (Dona Ana County). Pollen: Unknown, but visits
flowers of Sesuvium verrucosum.
Perdita martini Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 14. d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 140, 155, 287, figs. 58, 1051, 1052,
1169. 2, 3 (key, tax. status, redescription, geogr. and floral records).
melanogastra Timberlake. Calif. (Tulare County).
Perdita melanogastra Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 313, fig. 67. °.
mimosae Timberlake. Tex., N. Mex. (Eddy County). Pollen: Unknown, but visits flowers of
Mimosa, Phacelia popei, Prosopis.
Perdita mimosae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 270, figs. 1035, 1036, 1161.
Cr Oe
modestissima Timberlake. Nev. (Battle Mountain). Pollen: Unknown, but holotype is labeled
from an undetermined species of Cruciferae.
Perdita modestissima Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 147, fig. 28. 2.
munita: Timberlake. Ariz. (Cochise County); Mexico (Durango). Pollen: Unknown, but visits
flowers of Chamaesaracha coronopus, Eriogonum, Physalis.
Perdita munita Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 205, figs. 24, 957, 958, 1122.
Oe
nigricornis Timberlake. Ariz. (Cochise and Pima Counties); Mexico (Sonora). Pollen:
Apparently may collect pollen from the flowers of Prosopis.
Perdita nigricornis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 331, figs. 1077, 1078, 1182.
é.
nigroaenea Timberlake. Utah (Kane County). Pollen: Unknown, but visits flowers of

Gutierrezia microcephala.
Perdita nigroaenea Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 158. °.

1920 Hymenoptera in America North of Mexico

nigroclypeata Timberlake. Colo. (Alamosa and Rio Grande Counties).
Perdita nigroclypeata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 314, fig. 69. 9°.
obliqua Timberlake. Ark. (Garland County), Ariz., south. Calif. (Colorado Desert); Mexico
(Baja California and Sonora). Pollen: Apparently polylectic, visits flowers of Acacia,
Carnegiea gigantea, Cercidium, Dalea spinosa, Eriogonum trichopes, Medicago sativa,
Prosopis, Wislizenia refracta.
Perdita obliqua Timberlake, 1928. Pan-Pacifie Ent. 5: 26. °.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 134, 144, 148, 218-219, figs. 975, 976,
1131. 2, 5 (key, redescription, geogr. and floral records). —Timberlake, 1971. Calif. Univ.
Pubs. Ent. 66: 48 (geogr. record).

obscurella Timberlake. N. Mex. (Hidalgo and Socorro Counties), Ariz. (Cochise and Pima
Counties); Mexico (Chihuahua). Pollen: Unknown, but visits flowers of Baileya
pleniradiata, Boerhaavia, Euphorbia albomarginata.
Perdita obscurella Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 202, figs. 23, 593, 594,
1120 OMe
omani Timberlake. Calif. (Bakersfield).
Perdita omani Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 296, fig. 62. 2.
oreophila Timberlake. Calif. (Tuolumne County). Pollen: Unknown, but visits flowers of a
Compositae.
Perdita oreophila Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 244. °.
ovaliceps Timberlake. N. Mex. (Eddy County), Ariz. (Maricopa and Mohave Counties), Nev.
(Clark County), south. Calif. (Colorado and Mojave Deserts). Pollen: Unknown, but visits
flowers of Chaenactis carphoclinia, C. stevioides var. brachypappa, Excelia farinosa,
Gerrea canescens, Larrea tridentata.
Perdita ovaliceps Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 265, figs. 52, 1029, 1030,
1158. 3.
Perdita luctuosa Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 266, fig. 53. 2.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 150-151 (synonymy, geogr. records).
pallidipes Timberlake. Ariz. (Maricopa and Pima Counties), south. Calif. (Colorado Desert);
Mexico (Sonora). Pollen: Unknown, but visits flowers of Prosopis juliflora.
Perdita pallidipes Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 332, figs. 1079, 1080, 1183.
é.

panocheana Timberlake. Calif. (Fresno County). Pollen: Unknown, but visits flowers of
Heliotropium oculatum.
Perdita panocheana Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 274, figs. 1039, 1040,
1163. 2, o.
parryellae Timberlake. Ariz. (Navajo County). Pollen: Unknown, but visits flowers of
Parryella filifolia.
Perdita parryellae Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 48, figs. 1889, 1390, 1413.
placens Timberlake. Ariz. (Willcox).
Perdita placens Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 124, figs. 1273, 1274, 1340. 6.
planifrons Timberlake. Calif. (Mono County).
Perdita planifrons Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 148. 9.
propinqua Timberlake. Calif. (Ventura County). Pollen: Unknown, but visits flowers of
Linanthus aureus, Phacelia douglasit.
Perdita propinqua Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 226, figs. 35, 983, 984,
1135. 2, do.
prosopidis Timberlake. Ariz. (Yuma County), south. Calif. (Colorado and Mojave Deserts);
Mexico (Baja California). Pollen: Possibly an oligolege of Prosopis juliflora, but visits
other flowers for nectar including Melilotus.
Perdita prosopidis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 318, figs. 69, 1063, 1064,
17 5e Oeoe
puncticeps Timberlake. Calif. (Palm Springs).
Perdita puncticeps Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 49. °.

Superfamily APOIDEA 1921

punctosignata flava Timberlake. Tex. west. to Ariz.; Mexico (Baja California, Chihuahua,
Coahuila, Sonora and Sinaloa). Pollen: Apparently an oligolege of Prosopis, but also
visits other flowers presumably for nectar including Gutierrezia, Medicago, Sapindus
saponaria.
Perdita punctosignata flava Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 354. 2, d.
punctosignata punctosignata Cockerell. N. Mex. (Dona Ana and Otero Counties), Ariz., Calif.
(Colorado Desert); Mexico (Baja California and Sonora). Pollen: Apparently an oligolege
of Prosopis.
Perdita punctosignata Cockerell, 1895. Psyche 7 (supp.): 6. 3.
Perdita howardi Cockerell, 1899. Ann. and Mag. Nat. Hist. (7) 3: 492. 9, d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 142, 157, 351-353, figs. 75, 1087,
1088, 1187. 2, d (key, redescription, synonymy, geogr. and floral records).
punctosignata sulphurea Timberlake. Ariz., south. Calif. (Colorado and Mojave Deserts, also
San Benito County), Nev. (Clark County), Utah (Washington County); Mexico (Baja
California). Pollen: Apparently an oligolege of Prosopis, but also visits other flowers
presumably for nectar including Acacia greggii, Cercidium floridum, Larrea tridentata,
Melilotus.
Perdita punctosignata sulphurea Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 353. 2, d.
punctulata Timberlake. South. Calif. (Colorado and Mojave Deserts), Ariz. (Yuma); Mexico
(Baja California and Sonora). Pollen: Oligolege of Larrea tridentata, but also visits other
flowers for nectar including Baccharis emoryi, Cercidium floridum, Lasthenia
chrysostoma, Melilotus, Prosopis juliflora.
Perdita punctulata Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 362, figs. 440, 441, 504. d,
he

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 139, 262 (key).

Biology: Hurd and Linsley, 1975. Calif. Univ. Pubs. Ent. 193: 24-25, fig. 6 (floral
relationships).

pusilla Timberlake. South. Calif. (western margin of Mojave Desert).
Perdita pusilla Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 257, figs. 1021, 1022, 1154. 9,
3

quadraticeps Timberlake. Ariz. (Patagonia).
Perdita quadraticeps Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 213, figs. 29, 969, 970,
1128. 6.
replicans Timberlake. Calif. (San Luis Obispo County).
Perdita replicans Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 149, fig. 29. 2.
richardsi Timberlake. California (Apple Valley in the Mojave Desert).
Perdita richardsi Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 227. 6.
salicis coloradana Timberlake. Colo. (Delta and Garfield Counties).
Perdita salicis coloradana Timberlake, 1929. N. Y. Ent. Soc., Jour. 37: 111. °.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 340, 341. ¢ (key, redescription,
geogr. records).
salicis euxantha Timberlake. Idaho (Idaho and Nez Perce Counties), Oreg. (near Corvallis).
Perdita salicis euxantha Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 349. 2, d.
salicis hirsutior Timberlake. Calif. (Olancha). Pollen: Presumably an oligolege of Saliz, visits
the flowers of S. exigua.
Perdita salicis hirsutior Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 346. 2, d.
salicis imperialis Cockerell. Wash. (Asotin, Spokane, Walla Walla and Whitman Counties),
Idaho (Blaine, Canyon, Nez Perce and Twin Falls Counties), Oreg. (Umatilla and
Walloun Counties), Utah (Cache, Iron, Juab and Washington Counties), south. Calif.
(Colorado and Mojave Deserts), Ariz. (Maricopa and Pima Counties), Colo. (Archuleta
County). Pollen: Presumably an oligolege of Salia, including S. gooddingii, but visits
other flowers apparently for nectar including Acacia greggii, Asparagus, Daucus carota,
Prosopis, Sisymbriwm.
Perdita exclamans imperialis Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 193. °.

1922 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 340, 342-343. 2, 5 (key,
redescription, geogr. and floral records).
salicis laeta Timberlake. South. Calif. (Needles). Pollen: Presumably an oligolege of Salix,
visits flowers of Prosopis juliflora.
Perdita salicis laeta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 347. 9, d.
salicis monoensis Timberlake. Calif. (Eldorado and Mono Counties). Pollen: Presumably an
oligolege of Salix, visits flowers of S. sessilifolia.
Perdita salicis monoensis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 345. 2, d.
salicis occidentalis Timberlake. South. Calif. (cismontane, except one population at Helendale
on Mojave Desert). Pollen: Apparently an oligolege of Salix including S. exigua, S.
gooddingii, S. nigra, but visits other flowers presumably for nectar including Alyssum
maritimum, Cryptantha microphylla var. lepida, Funastrum heterophyllum,
Ligustrum, Pyracantha.
Perdita salicis occidentalis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 348. 2, ¢.
salicis personata Timberlake. South. Calif. (San Diego County). Pollen: Presumably an
oligolege of Salix, visits flowers of S. laevigata.
Perdita salicis personata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 344. 9°, d.
salicis salicis Cockerell. Colo., N. Mex. (Las Cruces); Mexico (Chihuahua). Pollen: Presumably
an oligolege of Salix.
Perdita salicis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 80. °, d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 138, 141, 157, 338-341, figs. 1085,
1086, 1186. 2, 3d (key, redescription, geogr. and floral records).
salicis sublaeta Timberlake. Oreg. (Hood River and The Dalles).
Perdita salicis sublaeta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 348. 2, 6.
salicis subtristis Cockerell. Colo. (Archuleta and Chaffee Counties), Utah (Grand County).
Pollen: Apparently an oligolege of Salix.
Perdita subtristis Cockerell, 1933. Canad. Ent. 65: 234. 9.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 135, 339, 341-342. 2, d (key,
redescription, geogr. and floral records).
salicis tristis Timberlake. Calif. (San Joaquin Valley). Pollen: Apparently an oligolege of Saliz,
visits flowers of S. hindsiana and also Populus.
Perdita salicis tristis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 345, fig. 74. 2, 6.
salviae Timberlake. South. Calif. (Colorado Desert). Pollen: Collects pollen from the flowers of
Salvia vaseyi.
Perdita salviae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 184, figs. 935, 936, 1111. 9, 6.
sandhouseae Timberlake. Ariz. (Apache County). Pollen: Unknown, but visits flowers of
Gutierrezia.
Perdita sandhouseae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 171, figs. 8, 921, 922,
1104. 6.
schlingeri Timberlake. Nev. (Emigrant Pass).
Perdita schlingeri Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 258. °.
semicaerulea Cockerell. Tex. to Ariz.; Mexico (Chihuahua and Coahuila). Pollen: Oligolege of
Larrea tridentata, visits both the spring and late summer and fall blooms of this plant
from which it obtains pollen and nectar, but also visits flowers of Prosopis presumably
for nectar only.
Perdita semicaerulea Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 64. °.
Perdita quadrangularis Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 129. d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 131, 150, 186-188, figs. 14, 937, 938,
1112. 2, 3 (key, redescription, synonymy, geogr. and floral records).
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 25 (floral relationships).

sexnotata Timberlake. South. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers of
Coldenia palmeri, Eschscholzia parishii.
Perdita sexnotata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 307, fig. 66. °.

Superfamily APOIDEA 1923

sidae Cockerell. N. Mex. (Mesilla); Mexico (Durango). Pollen: Unknown, but visits flowers of

Sida hederacea.
Perdita sidae Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 353. 9, d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 128, 132, 148, 263-264, figs. 51, 1025,
1026, 1156. ¢ (key, redescription). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 50. 2
(redescription, geogr. record).

sonorensis Cockerell. Ariz. (Yuma County), south. Calif. (Colorado and Mojave Deserts);
Mexico (Baja California and Sonora). Pollen: Oligolege of Prosopis including P. juliflora,
P. pubescens, but also visits flowers of Encelia farinosa, Eucnide urens, Heliotropium
curassavicum, Melilotus, Peucephyllum schottii, Pluchea sericea.
Perdita sonorensis Cockerell, 1899. Ann. and Mag. Nat. Hist. (7) 3: 493. °.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 136, 138, 152, 272-274, figs. 55, 1037,
1038, 1162. 2, 6 (key, redescription, geogr. and floral records).
speciosa Timberlake. Ariz. (Coconino County). Pollen: Unknown, but visits flowers of
Wislizenia refracta.
Perdita speciosa Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 51, figs. 1891, 1392, 1414. d.
sphaeralceae alticola Cockerell. N. Mex., Ariz. (Cochise and Coconino Counties). Pollen:
Possibly an oligolege of Sphaeralcea, visits flowers of S. lobata, but also visits flowers
of Heterotheca subaxillaris.
Perdita sphaeralceae race alticola Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 68. °,

3.
Perdita sphaeralceae race alticola mut. suffusa Cockerell, 1896. Acad. Nat. Sci. Phila.,
Proc. 48: 68. d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 154, 178-179, fig. 11. 2, 3d (key,
geogr. and floral records).
sphaeralceae sphaeralceae Cockerell. Tex. (Hudspeth County), N. Mex. (Dona Ana and Luna
Counties); Mexico (Chihuahua, San Luis Potosi and Zacatecas). Pollen: Possibly an
oligolege of Sphaeralcea including S. angustifolia, but visits other flowers including
Cologania, Gutierrezia lucida, Haplopappus heterophyllus, Mentzelia, Solidago

canadensis.
Perdita sphaeralceae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 66. 2, d.

Perdita pellucida Cockerell, 1896. Acad. Nat. Sci. Phila., Proce. 48: 88. ¢.
Perdita sphaeralceae var. ridens Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 547. 6.

Taxonomy: Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 121 (tax. status, as pellucida).
—Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 127, 154, 176-178. 2, 3 (key, redescription,
synonymy, geogr. and floral records).

stathamae eluta Timberlake. Ariz. (Salt River Mts.).
Perdita stathamae eluta Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 358. 2, d.
stathamae stathamae Timberlake. N. Mex. (Alamogordo), Ariz. (Chihuahua and Sonora
Deserts), south. Calif. (Colorado Desert); Mexico (Baja California and Sonora). Pollen:
Unknown, but visits flowers of Acacia greggii, Prosopis juliflora.
Perdita stathamae stathamae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 365, figs. 1091,
1092, 1189. 2, do.
stenopyga Timberlake. Calif. (Inyo County). Pollen: Unknown, but visits flowers of
Chaenactis, Mentzelia affinis.
Perdita stenopyga Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 229, figs. 38, 989, 990, 1138.
REG)
sternalis Timberlake. South. Tex. (Cameron and Galveston Counties). Pollen: Unknown, but
visits flowers of Coreopsis douglasii, Heliotropium, Rubus.
Perdita sternalis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 291, figs. 1055, 1056, 1171.
Omar
sulphuripes Timberlake. Calif. (near Palm Springs, also Slate Mts., Inyo County). Pollen:
Unknown, but visits flowers of Malacothriz.
Perdita sulphuripes Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 267, figs. 1031, 1082,

1159. 6.

1924 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 151. ¢ (geogr. and floral record).
tarda Cockerell. N. Mex. (Las Cruces). Pollen: Unknown, but visits flowers of Gutierrezia

microcephala.
Perdita tarda Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 85. d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 146, 151, 279-280 (redescription of
type specimen).
tenebrosa Timberlake. Ariz. (Chiricahua Mts.).
Perdita tenebrosa Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 131. 2.
tessellata Timberlake. Idaho (Elmore County), Nev. (Battle Mountain). Pollen: Unknown, but
visits flowers of Chaenactis, Tetradymia.
Perdita tessellata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 264, figs. 1027, 1028, 1157.
Gr

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 118, 117, 151-152, fig. 30. 2, d (key,
redescription, geogr. and floral records).
thelypodii Timberlake. Calif. (Inyo and Los Angeles Counties). Pollen: Unknown, but visits
flowers of Cleomella obtusifolia, Thelypodium brachycarpum.
Perdita thelypodii Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 354, figs. 432, 433, 500. ?,
é.

tortifoliae fremonti Timberlake. South. Calif. (Mohave Desert), Nev. (Clark County), Utah
(Washington County). Pollen: Collects pollen from the flowers of Lepidiwm fremontii.
Perdita fremonti Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 250, figs. 1011, 1012, 1149.
OG.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 143 (tax. status, geogr. and floral
records).
tortifoliae tortifoliae Cockerell. Colo. (Florissant and Wilkerson Pass), N. Mex. (Rio Arriba
County). Pollen: Presumably collects pollen from the flowers of Lepidiwm jonesii, but
also visits flowers of Bigelovia tortifoliae, Eriogonwm umbellatum, Ranunculus

eremogenes.
Perdita tortifoliae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 440. 9.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 132, 144, 245-247, figs. 45, 1005,
1006, 1146. 2, 3 (key, redescription, geogr. and floral records). —Timberlake, 1968. Calif.
Univ. Pubs. Ent. 49: 143 (tax. status, geogr. and floral record).

triangulifera Timberlake. West. Tex. to south. Calif., deserts. Pollen: Apparently an oligolege
of Prosopis, but visits other flowers presumably for nectar including Acacia greggii,
Cercidium floridum, Dithyrea wislizeni, Melilotus, Mimosa, Phoradendron, Tamarix.
Perdita triangulifera Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 328, figs. 71, 1075-1076,
iblebl, Se ee
trinotata Timberlake. Tex. Pollen: Unknown, but visits flowers of Lesquerella gordonii, Nama
hispidum, Nerisyrenia camporum.
Perdita trinotata Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 249, figs. 47, 1009, 1010,
1148. 3.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 115, 116, 144-145, fig. 26. 2, ¢ (key,
redescription, geogr. and floral records).
veris Timberlake. Ariz. (Pima County). Pollen: Collects pollen from the flowers of Nama
hispidum, but visits other flowers for nectar including Teucrium cubense depressum.
Perdita veris Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 187, figs. 21, 1289, 1290, 1348. 2
3.
vidua Timberlake. Tex. (E] Paso County). Pollen: Unknown, but visits flowers of Gossypium.
Perdita vidua Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 309. 2.
vittata conformis Timberlake. Nev. (Lincoln County).
Perdita vittata conformis Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 284. 2, ¢.

Superfamily APOIDEA = 1925

vittata tricolor Timberlake. South. Calif. (Mojave Desert and south. San Joaquin Valley).
Pollen: Unknown, but visits flowers of Cleomella obtusifolia, Wislizenia refracta. The
typical subspecies, Perdita vittata vittata Cockerell, occurs in Mexico (Baja California).
Perdita vittata tricolor Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 283. ¢, @.
werneri Timberlake. Ariz. (Springerville).
Perdita werneri Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 154, figs. 1297, 1298, 1352. 3.

wilmattae miricornis Cockerell. Wyo. (Green River), Colo. (Mesa and Moffat Counties), Utah
(Carbon County), Ariz. (Grand Canyon). Pollen: Unknown, but visits flowers of Cleome,
Engelmannia pinnatifida, Phacelia, Stanleya pinnata.
Perdita miricornis Cockerell, 1922. Amer. Mus. Novitates 33: 9. 6, 2.
Perdita miricornis var. leucorhina Cockerell, 1922. Amer. Mus. Novitates 33: 10. 9.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 310, 314, 360. 3, 2 (key,
redescription, tax. status, geogr. and floral records). —Timberlake, 1964. Calif. Univ. Pubs.
Ent. 28: 245 (tax. status). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 145 (key).

wilmattae stanleyae Timberlake. Utah (Grand, Uintah and Washington Counties), Nev., Calif.
(Inyo County). Pollen: Unknown, but visits flowers of Asclepias, Cleome lutea,
Mentzelia, Prunus, Stanleya pinnata, Thelypodium laciniatum.
Perdita stanleyae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 253, figs. 48, 1017, 1018,
152 One

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 145 (key, tax. status, geogr. record).

wilmattae wilmattae Cockerell. Colo., Utah, Nev. (Humboldt County). Pollen: Unknown, but
visits flowers of Cleome lutea, Engelmannia, Phacelia alba, Stanleya pinnata.
Perdita wilmattae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 441. d, 2.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 310, 314, 359-360, figs. 436, 437, 502.
3, 2 (key, redescription, tax. status, geogr. and floral records). —Timberlake, 1964. Calif.
Univ. Pubs. Ent. 28: 137, 245 (key, floral records). —Timberlake, 1968. Calif. Univ. Pubs.
Ent. 49: 145 (key).

wislizeniae Timberlake. N. Mex. (Luna County), Ariz. (Cochise County); Mexico (Chihuahua).
Pollen: Apparently an oligolege of Wislizenia refracta, but also visits flowers of Cleome,
Eriogonum thomasii, Lepidium.
Perdita wislizeniae Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 284, figs. 57, 1049, 1050,
1168. 2, 6. ;

zebrata flavens Timberlake. Nev., Oreg., Idaho, Wyo., Utah, Colo., Calif. (Inyo County). Pollen:
Apparently collects pollen only from the flowers of Cleome including C. lutea, C.
serrulata, but visits other flowers for nectar including Engelmannia pinnatifida,
Phacelia, Salsola kali.
Perdita zebrata flavens Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 357. 9, d.

Taxonomy: Timberlake, 1964. Calif. Univ. Pubs. Ent. 28: 138, 153, 262 (key, geogr. and floral
record).
zebrata zebrata Cresson. N. Dak., Nebr. and N. Mex., west to Idaho, Nev. and Ariz. Parasite:
Neolarra pruinosa Ashmead. Pollen: Collects pollen from the flowers of Cleome
including C. serrulata, but visits other flowers presumably for nectar including
Haplopappus heterophyllus, Helianthus petiolaris, Nolina microcarpa, Solidago.
Perdita zebrata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 69. 2 (part).
Perdita canina Cockerell, 1895. Acad. Nat. Sci. Phila., Proc. 47: 17. d.
Perdita bakerae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 69. d, °.

Taxonomy: Cockerell, 1903. In Viereck, Amer. Ent. Soc., Trans. 29: 51. —Timberlake, 1958.
Calif. Univ. Pubs. Ent. 14: 310, 313, 356-357, figs. 434, 435, 501 (key, tax. characters, tax.
status, synonymy, geogr. and floral records). —Timberlake, 1964. Calif. Univ. Pubs. Ent.
28: 138, 153, 262 (key, geogr. range). —Rozen, 1966. Amer. Mus. Novitates 2259: 17-19, figs.
41-46 (larva). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 118, 150, figs. 1295, 1296,
1351. 3 (key, variation, geogr. records).

Biology: Custer, 1929. Canad. Ent. 61: 49 (nesting habits with description of larva). —Rozen,
1967. Amer. Mus. Novitates 2297: 34-38, figs. 2, 4, 11, 14, tables 1-2 (nest architecture, life
history, pollen source, parasite).

1926 Hymenoptera in America North of Mexico

SPECIES GROUP VALIDA
This species may be better placed in a new genus related to Perdita.

Taxonomy: Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 106 (tax. characters).

valida Timberlake. N. Mex. (Mesilla).
Perdita valida Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 106. °.

Genus PERDITA Subgenus PERDITELLA Cockerell

Perdita subg. Perditella Cockerell, 1899. Psyche 8: 312.
Type-species: Perdita larreae Cockerell. Orig. desig. (=Perdita laneae Cockerell).

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 348 (key). —Timberlake, 1956. Calif.
Univ. Pubs. Ent. 11: 267-268 (redescription, key to included spp.).
cladothricis Cockerell. West. Tex., N. Mex., Ariz.; Mexico (Baja California, San Luis Potosi,
Sonora and Zacatecas). Pollen: Possibly an oligolege of Tidestromia including T.
lanuginosa, but visits other flowers presumably for nectar including Gutierrezia
microcephala, Haplopappus wrightii, Medicago sativa, Pectis papposa.
Perdita cladothricis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 82. 2, 6.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 268, 270, figs. 207, 208, 305 (male
genitalia, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 30
(geogr. records).

larreae Cockerell. N. Mex., Ariz., South. Calif.; Mexico (Baja California, Chihuahua and

Coahuila). Pollen: Collects pollen from the flowers of Larrea tridentata, but visits these
and other flowers for nectar including Aplopappus heterophyllus, Asclepias, Baccharis
glutinosa, Croton californicus, Dalea, Dicraurus, Eriogonum reniforme, Euphorbia
polycarpa, Pectis papposa, Tamarix.

Perdita larreae Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 62. ¢.

Perdita larrearum Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 63. °.

Perdita larreae var. modesta Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 355. 6.

Perdita laneae Cockerell, 1899. Psyche 8: 312. Error for larreae.

Taxonomy: Timberlake, 1928. Amer. Mus. Novitates 321: 11. 2. —Timberlake, 1956. Calif.
Univ. Pubs. Ent. 11: 268, figs. 203, 204, 304 (tax. characters, synonymy, geogr. and floral
records). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17: 134 (variation, geogr. and floral
records).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 25-26, fig. 7 (floral
relationships).
marcialis Cockerell. West. Tex. to south. Calif., Nev.; Mexico (Coahuila, Sonora). Pollen:

Collects pollen from the flowers of Larrea tridentata and possibly also from other
flowers including Aloysia wrightii, Chrysothamnus paniculatus, Eriogonum
fasciculatum var. polifolium, Phacelia, Salvia vaseyi.

Perdita marcialis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 62. ¢.

Perdita phaceliae Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 450. 2.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 268, 269-270, figs. 205, 206 (key,
redescription, geogr. and floral records). —Timberlake, 1960. Calif. Univ. Pubs. Ent. 17:
134-135 (synonymy).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 26 (floral relationships).

minima Cockerell. Ariz., South. Calif.; Mexico (Baja California). Pollen: Collects pollen from
the flowers of Euphorbia including E. albomarginata, E. polycarpa, E. p. var. hirtella,
but visits these and other flowers for nectar including Croton californicus, Eriogonum
gracile, Gutierrezia.
Perdita minima Cockerell, 1923. Amer. Mus. Novitates 66: 4. 9.

Taxonomy: Cockerell, 1925. Pan-Pacific Ent. 1: 179. 6. —Timberlake, 1956. Calif. Univ. Pubs.
Ent. 11: 268, 270, figs. 209, 210 (key, redescription, geogr. and floral records).

Superfamily APOIDEA 1927

Genus PERDITA Subgenus PROCOCKERELLIA Timberlake

Perdita subg. Procockerellia Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 402.
Type-species: Perdita albonotata Timberlake. Orig. desig.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 5-7 (key to spp. of subgenera
Procockerellia, Allomacrotera).
albonotata Timberlake. South. Calif. (Colorado and Mojave Deserts), Ariz. (Mohave County),
Utah (Cache and Washington Counties); Mexico (Sonora). Pollen: Collects pollen from
the flowers of Stephanomeria including S. exigua, S. pauciflora, but visits other flowers
presumably for nectar including Agave deserti, Chaenactis glabriuscula, Eriogonwm
plumatella.
Perdita albonotata Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 403, figs. 91, 92, 161. 2, d.
brachyglossa Timberlake. Ariz. (Coconino County). Pollen: Unknown, but visits flowers of
Thelesperma.
Perdita brachyglossa Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 6. 2.
excellens Timberlake. Utah (Grand and Washington Counties), Ariz. (Coconino County).
Pollen: Unknown, but visits flowers of Cleome lutea.
Perdita excellens Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 384. 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 22. d.

moabensis Timberlake. Utah (Moab).
Perdita moabensis Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 7. 9.

Genus PERDITA Subgenus PPEUDOMACROTERA Timberlake

Perdita subg. Pseudomacrotera Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 349.
Type-species: Perdita turgiceps Timberlake. Orig. desig.
turgiceps Timberlake. Calif. (Colorado and Mojave Deserts), Ariz., (Maricopa Mts.); Mexico

(Baja California). Pollen: Apparently an oligolege of Larrea tridentata, but visits a wide
variety of other flowers for nectar including Acacia greggii, Cercidium floridum,
Cowania stansburiana, Cryptantha angustifolia, C. barbigera, Dalea californica, D.
fremontii, D. schottii, Datura meteloides, Encelia farinosa, Eriodictyon crassifolium,
Eriogonum inflatum, E. trichopes, Eschscholzia, Eucnide urens, Hyptis emoryi, Larrea
tridentata, Lasthenia, Nama hispida, Prosopis juliflora.

Perdita turgiceps Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 350, figs. 3, 4, 116, 117. 3, ¢:

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 26 (floral relationships).
Genus PERDITA Subgenus PYGOPERDITA Timberlake

Perdita subg. Pygoperdita Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 275.
Type-species: Perdita interrupta Cresson. Orig. desig.

Although several of the species visit flowers of the Compositae, especially Layia and
Malacothrix, most members of this subgenus visit flowers of other families, such as the
Rosaceae, Papaveraceae, Rhamnaceae and Liliaceae.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 275-284 (key to included spp.).
SPECIES GROUP INTERRUPTA
associata Timberlake. Calif. (San Benito County).
Perdita associata Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 32. °.
aureovittata aureovittata Cockerell. South. Calif. (cismontane, south of San Gabriel Mts.).

Pollen: Unknown, but visits flowers of Layia, Malacothrix californica var. glabrata.
Perdita (Cockerellia) aureovittata Cockerell, 1916. Canad. Ent. 48: 391. OF

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 391 (tax. status). —Timberlake, 1956.
Calif. Univ. Pubs. Ent. 11: 276, 281, 285, figs. 221, 222, 311, 387. 2, d (redescription, new
status, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 33 (geogr.
range).

1928 Hymenoptera in America North of Mexico

aureovittata maderensis Timberlake. Calif. (Madera County). Pollen: Unknown, but
presumably visits flowers of Malacothrix californica.
Perdita aureovittata maderensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 286. 2, 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 33 (geogr. range).
aureovittata soluta Timberlake. Calif. (South Coast and Peninsular Ranges). Pollen: Unknown,
but visits flowers of Agoseris auriantiaca, A. heterophylla, Encelia actoni, Linanthus

aureus, Malacothrix californica.
Perdita soluta Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 394, fig. 520. ¢.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 33 (tax. status, geogr. range).
aureovittata stenozona Timberlake. Calif. (San Benito and San Luis Obispo Counties). Pollen:
Unknown, but visits flowers of Chaenactis glabriuscula, Malacothrix californica.
Perdita aureovittata stenozona Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 286. 2, 6.
fieldi Timberlake. South. Calif. (Monterey to San Diego County). Pollen: Unknown, but visits

flowers of Adenostoma fasciculatum, Calochortus splendens, Eriogonum fasciculatum.
Perdita fieldi Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 291, figs. 227, 228, 315, 340. °,
Oe
inflexa Timberlake. Calif. (Riverside County). Pollen: Unknown, but visits flowers of
Eschscholzia multiflora var. darwinensis.
Perdita inflexa Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 298, figs. 237, 238, 345, 346. d.
interrupta interrupta Cresson. South. Calif. (cismontane). Pollen: Collects pollen from the
flowers of Eschscholzia californica, but visits other flowers for nectar including
Baccharis emoryi, Coreopsis californica, Cryptantha intermedia, Lasthenia
chrysostoma, Lepidospartum squamatum, Sisymbrium irio, Solidago occidentalis,
Tamarix gallica; also has been taken at the flowers of Yucca, but since these flowers
are not attractive to bees, the plant in question was presumably Nolina parryi.
Perdita interrupta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 70. 3.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 277, 282, 292, figs. 229, 230, 316, 341.
9, 3 (redescription, tax. status, geogr. and floral records).
interrupta kernensis Timberlake. Calif. (Kern County). Pollen: Unknown, but visits flowers of
Cryptantha.
Perdita interrupta kernensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 293. 9, ¢.
interrupta vernalis Timberlake. Calif. (San Diego County).
Perdita interrupta vernalis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 298. 9, ¢.
layiae excisa Timberlake. South. Calif. (cismontane). Pollen: Unknown, but visits flowers of
Chaenactis glabriuscula, Layia platyglossa.
Perdita layiae excisa Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 288. 2, d.
layiae layiae Cockerell. Calif. (San Miguel Island). Pollen: Unknown, but visits flowers of
Layia platyglossa.
Perdita layiae Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 152. 9, 6.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 277, 281, 288, figs. 223, 224, 318, 338
(redescription).
malacothricis Timberlake. South. Calif. deserts, Nev. (Clark County), Ariz. (Tucson). Pollen:

Unknown, but visits flowers of Aster, Baileya multiradiata, B. pleniradiata,
Calycoseris parryi, Chaenactis fremontii, Hymenoclea salsola, Larrea tridentata,
Malacothrix californica var. glabrata, Stephanomeria exigua, S. pauciflora.

Perdita malacothricis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 289, figs. 225, 226, 314,
Boo Neel

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 36 (geogr. and floral records).
malacothricis var. unica Timberlake. Calif. (Riverside County).
Perdita malacothricis var. unica Timberlake, 1962. Calif. Univ. Pubs. Ent. 28: 96. °.
micheneri micheneri Timberlake. South. Calif. (cismontane); Mexico (Baja California). Pollen:
Unknown, but visits flowers of Ceanothus cordulatus, C. greggii, C. leucodermis, C.

Superfamily APOIDEA — 1929

orcuttii, Cryptantha intermedia, C. micrantha, C. muricata, Eriophyllum
confertiflorum, Eschscholzia californica, Rhamnus crocea, Rhus ovata.

Perdita micheneri micheneri Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 294, figs. 231,
232, 317, 342. 2, d.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 36 (geogr. record).
micheneri rhamnophila Timberlake. Calif. (Rock Creek in San Gabriel Mts.). Pollen:
Unknown, but visits flowers of Rhamnus crocea.
Perdita micheneri rhamnophila Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 295. d.
mucronata Timberlake. Calif. (Riverside County); desert. Pollen: Unknown, but visits flowers
of Camissonia brevipes, Eschscholzia minutiflora var. darwinensis.
Perdita mucronata Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 297, figs. 235, 236, 318,
344. 6.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 36. 9.
nitens Timberlake. Calif. (Mendocino, San Benito, Santa Barbara and Stanislaus Counties).
Pollen: Unknown, but visits flowers of Eriogonum, Eschscholzia californica, Mimulus
fremontii, Phacelia.
Perdita nitens Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 295, figs. 233, 234, 343. 2, 6.
quadrisignata Timberlake. Calif. Pollen: Unknown, but visits flowers of Baccharis, Ceanothus,
Clarkia, Coreopsis bigelovii, Cryptantha muricata, Eriophyllum confertiflorum,
Eschscholzia californica, Layia platyglossa, Plagiobothrys nothofulvus, Prunus
subcordata, Rhamnus crocea, Salix.
Perdita quadrisignata Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 296. °.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 39. d.
transversa Timberlake. Oreg. (Jackson County).
Perdita transversa Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 287, figs. 312. 6.

vandykei Timberlake. Calif. (Monterey County).
Perdita vandykei Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 286. 2, d.

SPECIES GROUP CALIFORNICA

Biology: Linsley, 1958. Hilgardia 27: 562, table 5 (oligolecty).
argemones Timberlake. Ariz. (Maricopa, Pima and Pinal Counties). Pollen: Unknown, but visits
flowers of Argemone.
Perdita argemones Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 309, figs. 261, 262, 327,
SDS Or
arizonica Timberlake. Ariz. (Oak Creek Canyon). Pollen: Apparently collects pollen from the
flowers of Calochortus.
Perdita arizonica Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 310, figs. 263, 264, 328, 359.
Omron
bilobata Timberlake. Calif. (Inyo and San Bernardino Counties). Pollen: Unknown, but visits
flowers of Calochortus aureus, C. kennedyi, Haplopappus cooperi.
Perdita bilobata Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 305, figs. 253, 254, 354. 2, d.
bispinata Timberlake. Calif. (San Bernardino and Riverside Counties). Pollen: Unknown, but
visits flowers of Calochortus kennedyi, Haplopappus cooperi.
Perdita bispinata Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 308, figs. 249, 250, 3238, 352.
ORIG s
bohartorum Timberlake. Calif. (Mono County).
Perdita bohartorum Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 319, figs. 275, 276, 365.
OPO
californica californica (Cresson). Coastal Calif. (Mt. Diablo, southward); Mexico (Baja
California). Pollen: Apparently collects pollen from the flowers of Calochortus including
C. concolor, C. kennedyi, C. luteus, C. plummerae, C. splendens, C. venustus, C. weedii,
but visits other flowers for nectar including Cryptantha, Eriogonum fasciculatum,
Eriophyllum confertiflorum, Grindelia, Opuntia.
Macrotera californica Cresson, 1878. Amer. Ent. Soe., Trans. 7: 71. 6.

1930 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 280, 282, 301-302. 9. —Timberlake,
1968. Calif. Univ. Pubs. Ent. 49: 34 (tax. status, geogr. range).
californica inopina Timberlake. Calif. (Inyo County).
Perdita californica inopina Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 33. 9°.
calochorti Timberlake. Calif. (San Bernardino and Inyo Counties). Pollen: Unknown, but visits
flowers of Calochortus nuttallii.
Perdita calochorti Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 299, figs. 239, 240, 320, 347.
25 8.
coalingensis Timberlake. Calif. (Coalinga). Pollen: Unknown, but visits flowers of E'schscholzia
californica.
Perdita coalingensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 313. °.
cowaniae Timberlake. Calif. (Inyo County). Pollen: Unknown, but visits flowers of Calochortus
kennedyi var. munzti, Cowania stansburiana.
Perdita cowaniae Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 315, figs. 269, 270, 331, 362.
ORG
digressa Timberlake. Ariz. (Tucson), Calif. (Granite Mts., San Bernardino County). Pollen:
Unknown, but visits flowers of Baileya multiradiata, Chaenactis.
Perdita digressa Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 34, figs. 1211, 1212, 1309,
1354. 6, @.
distropica Timberlake. Calif. (South Coast Ranges). Pollen: Collects pollen from the flowers of
Calochortus including C. clavatus, C. luteus, C. splendens, C. venustus and Eschscholzia
californica, but visits other flowers for nectar including Adenostoma, Eriogonum.
Perdita distropica Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 312, figs. 265, 266, 329, 360.
OPEGr
duplonotata Timberlake. Utah (Garfield County), Nev. (Charleston Mts.), Calif. (San
Bernardino County). Pollen: Unknown, but visits flowers of Fallugia paradoxa,
Isomeris arborea.
Perdita duplonotata Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 307, figs. 257, 258, 356.
ORaGle
eriogoni Cockerell. Colo. Pollen: Unknown, but visits flowers of Eriogonum umbellatum.
Perdita eriogoni Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 621. 9, ¢.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 279, 284, 322-323, figs. 279, 280, 367
(tax. characters). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 35-36. 3 (color variation).
fallugiae Timberlake. Nev. (Charleston Mts.). Pollen: Unknown, but visits flowers of
Calochortus flecuosus, Fallugia paradoxa, Hymenoxys cooperi.
Perdita fallugiae Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 314, figs. 267, 268, 330, 361.
OAC
leucostoma Timberlake. Calif. (Kern and Inyo Counties). Pollen: Unknown, but visits flowers
of Calochortus leichtlinii, Haplopappus linearifolius.
Perdita leucostoma Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 300, figs. 241, 242, 321,
348. 2, d.
macrostoma Cockerell. Calif. Pollen: Apparently collects pollen from the flowers of
Calochortus.
Perdita macrostoma Cockerell, 1922. U. S. Natl. Mus., Proc. 60 (18): 18. d, @.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 278, 283, 304, figs. 251, 252, 324, 353
(tax. characters).
mohavensis mohavensis Timberlake. South. Calif. deserts. Pollen: Probably collects pollen

mainly from the flowers of E'schscholzia, but visits flowers of Amsinckia intermedia,
Argemone platyceras, Calochortus aureus, Calycoseris wrightii, Chaenactis stevioides
var. brachypappa, Encelia farinosa, Eschscholzia californica, E. glyptosperma, E.
minutiflora var. darwinensis, E. parishii, Haplopappus cooper, Hyptis emoryi, Larrea
tridentata, Monardella exilis, Salix, Salvia.

Perdita mohavensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 306, figs. 255, 256, 325,
355. 2, d.

Superfamily APOIDEA 1931

mohavensis pimana Timberlake. Ariz. (Pima County). Pollen: Unknown, but visits flowers of
Baileya.
Perdita mohavensis pimana Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 36. 9.
montereyensis Timberlake. Calif. (Monterey, San Benito and San Luis Obispo Counties).
Pollen: Collects pollen from the flowers of Calochortus splendens, C. venustus and
Eschscholzia californica, but visits other flowers for nectar including Eriogonum,
Rhamnus crocea.
Perdita montereyensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 302, figs. 245, 246, 350.
ONE
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 36 (color variation, geogr. and floral
record).
mormonica Timberlake. Utah (Garfield and Tooele Counties). Pollen: Unknown, but visits
flowers of Sphaeralcea.
Perdita mormonica Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 311. @.

Taxonomy: Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 10 (color variation).

nebrascensis Swenk and Cockerell. Nebr., (Neligh, Antelope County). Pollen: Unknown, but
visits flowers of Helianthus.
Perdita nebrascensis Swenk and Cockerell, 1907. Ent. News 18: 53. 2, 6.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 395-396 (key, tax. status).
nevadensis culbertsoni Timberlake. Calif. (Tulare, Santa Cruz Counties, Yosemite Natl. Pk.).
Perdita nevadensis culbertsoni Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 318. 2, 6.
nevadensis molina Timberlake. Calif. (San Bernardino Mts.).
Perdita nevadensis molina Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 318, figs. 273, 274,
364. 2, d.

Taxonomy: Timberlake, 1958. Calif. Univ. Pubs. Ent. 14: 396 (key).
nevadensis nevadensis Cockerell. B. C. to north. Calif. Pollen: Unknown, but visits flowers of

Eriogonum.
Perdita nevadensis Cockerell, 1896. Acad. Nat. Sci. Phila., Proc. 48: 58. @.
Perdita nigricollis Timberlake, 1929. Pan-Pacific Ent. 6: 54. 9.

Taxonomy: Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 279, 284, 316-318, figs. 271, 272, 332,
363. 2, d (redescription, synonymy).
obtusa Timberlake. Calif. (Tulare County).
Perdita obtusa Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 37. 2.
pulliventris Timberlake. Calif. (Tuolumne County).
Perdita pulliventris Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 38. °.
robustula Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Acacia greggii,
Agave deserti, Argemone corymbosa, A. platyceras, Cryptantha utahensis, Encelia
farinosa, Eriogonum inflatum, Eschscholzia minutiflora var. darwinensis, Prosopis

juliflora, Stanleya pinnata.
Perdita robustula Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 308, figs. 259, 260, 326, 357.

Oe
santaclarensis Timberlake. Calif. (Santa Clara County).
Perdita santaclarensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 314. 9°.

tularensis Timberlake. Calif. (Alpine, Tuolumne, and Mariposa Counties, south to Tehachapi
Pass). Pollen: Unknown, but visits flowers of Calochortus luteus, C. venustus.
Perdita tularensis Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 303, figs. 247, 248, 351. °,
Or

wyomingensis sculleni Timberlake. Oreg., Wash.
Perdita sculleni sculleni Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 320, figs. 277, 278,

333, 366. 2, do.
Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 40 (tax. status, key).
wyomingensis segona Timberlake. Utah, Wyo., Nev. Pollen: Unknown, but visits flowers of

Calochortus nuttallir.
Perdita sculleni segona Timberlake, 1956. Calif. Univ. Pubs. Ent. 11: 322. 2, ¢.

1932 Hymenoptera in America North of Mexico

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 40 (tax. status, key).
wyomingensis wyomingensis Cockerell. Wyo. (Teton County). Predator: Philanthus pulcher
Dalla Torre.
Perdita wyomingensis Cockerell, 1922. Amer. Mus. Novitates 33: 13. 9.

Taxonomy: Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 40 (type, tax. status, key).

Genus PERDITA Subgenus XEROMACROTERA Timberlake

Perdita subg. Xeromacrotera Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 412.
Type-species: Macrotera cephalotes Cresson. Orig. desig.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 347 (key).
cephalotes (Cresson). Nev., Calif. (Kern and San Bernardino Counties), Ariz. (Coconino
County). Pollen: Unknown, but visits flowers of Chrysothamnus, Gutierrezia lucida.
Macrotera cephalotes Cresson, 1878. Amer. Ent. Soc., Trans. 7: 71. 6.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 413, figs. 109, 110, 170 (male
genitalia, geogr. and floral records). —Timberlake, 1968. Calif. Univ. Pubs. Ent. 49: 22
(geogr. and floral record). —Timberlake, 1971. Calif. Univ. Pubs. Ent. 66: 7 (geogr. and
floral record).

Genus PERDITA Subgenus XEROPHASMA Cockerell

Xerophasma Cockerell, 1923. Amer. Mus. Novitates 66: 1.
Type-species: Perdita bequaertiana Cockerell. Monotypic and orig. desig.
(=Xerophasma bequaerti Cockerell).

These species are apparently crepuscular and obtain pollen only from the flowers of
Oenothera.

Taxonomy: Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 346, 348-349 (redescription, key to
included spp.).
bequaertiana Cockerell. Tex. (E] Paso and Howard Counties), N. Mex. (Albuquerque). Pollen:
Collects pollen from the flowers of Oenothera.
Xerophasma bequaerti Cockerell, 1923. Amer. Mus. Novitates 66: 2. 2. Preocce.
Perdita bequaertiana Cockerell, 1951. In Michener, Jn Muesebeck, Krombein and Townes,
U.S. Dept. Agr., Agr. Monog. 2: 1089. N. name.

Taxonomy: Timberlake, 1953. Kans. Univ. Sci. Bul. 35: 962-963. ¢. —Timberlake, 1954. Calif.
Univ. Pubs. Ent. 9: 349, figs. 1, 2, 115 (key). —Timberlake, 1958. Calif. Univ. Pubs. Ent. 14:
375 (geogr. record).

pallida Timberlake. Calif. (Colorado Desert), Ariz. (Yuma). Ecology: Attracted to artificial
lights near dusk through early evening. This species was readily observed in flight at
Borrego and Hopkins Well on the Colorado Desert by shining automobile headlights on
the patches of Oenothera being visited by this species. Commencing about nightfall
many males and some females were attracted to the lights of gasoline lanterns. Activity
at the flowers continued well into the darkness of early evening. These observations
were made in sand dune areas and suggest the possibility that this species may be
arenophilous in its choice of nesting sites. Pollen: Collects pollen from the flowers of
Oenothera deltoides, O. trichocalyx, but visits these and other flowers for nectar
including Trichostema.

Perdita pallida Timberlake, 1954. Calif. Univ. Pubs. Ent. 9: 349. 2, ¢.

NOMEN NUDUM IN PERDITA
Perdita excisa Timberlake, 1928. Amer. Mus. Novitates 321: 10.

Family HALICTIDAE

This is among the largest and most widely distributed families of bees. All three of its sub-
families occur in the Nearctic Region, but only the Dufoureinae and the Halictinae are well
represented by numerous species.

Superfamily APOIDEA 1933

Revision: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 515-521, figs. (spp. of eastern
U.S.).

Biology: Sakagami and Michener, 1962. The Nest Architecture of the Sweat Bees. Kansas
Univ. Press, 135 pp., 181 figs. —Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc.,
Jour. 75: 145-146, table 1 (biological differences among subfamilies).

SUBFAMILY DUFOUREINAE

Although this subfamily is chiefly Holarctic in distribution, it is also represented in the
Ethiopian, Oriental and Neotropical regions by at least one or more species. It is absent in the
Australian Region. Many, if not most of the species, are apparently highly restricted in their in-
trafloral relationships and thus oligolecty is a relatively common phenomenon of this subfamily.

Taxonomy: Michener, 1965. Ent. Soc. Amer., Ann. 58: 321-326, 28 figs. (generic review).

Genus DUFOUREA Lepeletier

Taxonomy: Bohart, 1947. Ent. Soc. Amer., Ann. 40: 692-704. —Bohart, 1948. Ent. Soc. Amer.,
Ann. 41: 119-136. —Bohart, 1949. Ent. Soc. Amer., Ann. 42: 55-62.

Genus DUFOUREA Subgenus DUFOUREA Lepeletier

Dufourea Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 227.
Type-species: Dufourea minuta Lepeletier. Desig. by Richards, 1935.
convergens Bohart. Calif. (San Diego Co.).
Dufourea convergens Bohart, 1949. Ent. Soc. Amer., Ann. 42: 59. d.
leachi Timberlake. Calif. (Humboldt and Mendocino Cos.). Pollen: Unknown, but visits flowers
of Brodiaea.
Dufourea leachi Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 398. d.
viridis Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Eschscholzia
californica, Platystemon californicus.
Dufourea (Dufourea) viridis Timberlake, 1941. Ent. Soc. Amer., Ann. 34: 39. 6, °.

Genus DUFOUREA Subgenus HALICTOIDES Nylander

Halictoides Nylander, 1848. Notiser Sallskapet Faune Flora Fenn., v. 1, p. 195.
Type-species: Halictoides dentiventris Nylander. Desig. by Cockerell and Porter,
1899.
Halictoides subg. Epihalictoides Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7)
4: 420.
Type-species: Panurgus marginatus Cresson. Monotypic.
Halictoides subg. Parahalictoides Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7)
4: 420.
Type-species: Halictoides campanulae Cockerell. Orig. desig.
Conohalictoides Viereck, 1904. Ent. News 15: 245.
Type-species: Panurgus novaeangliae Robertson. Monotypic and orig. desig.
(=Conohalictoides lovelli Viereck).
Neohalictoides Viereck, 1904. Ent. News 15: 261.
Type-species: Panurgus maurus Cresson. Monotypic and orig. desig.
Cryptohalictoides Viereck, 1904. Ent. News 15: 261.
Type-species: Cryptohalictoides spiniferus Viereck. Monotypic and orig. desig.
Mimulapis Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 162.
Type-species: Mimulapis versatilis Bridwell. Monotypic and orig. desig.
Betheliella Cockerell, 1924. Ent. News 35: 169.
Type-species: Betheliella calochorti Cockerell. Monotypic.
Halictoides subg. Cephalictoides Cockerell, 1924. Psyche 31: 244.
Type-species: Halictoides paradoxus Morawitz. Monotypic and orig. desig.
afasciata Bohart. Oreg., Calif. Pollen: Unknown, but visits flowers of Trifolium.
Dufourea afasciata Bohart, 1948. Ent. Soc. Amer., Ann. 41: 180. d, ?.

1934 Hymenoptera in America North of Mexico

australis australis (Michener). South. Calif. Pollen: Unknown, but visits flowers of Chaenactis,
Encelia farinosa, Erigeron foliosus, Eriophyllum, Lasthenia gracilis. A third
subspecies occurs in Baja California.
Halictoides australis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 398. d.
australis dammersi Timberlake. South. Calif. (deserts).
Dufourea (Halictoides) dammersi Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 407. 3.
bernardina (Michener). Calif. Pollen: Unknown, but visits flowers of Agoseris aurantiaca,
Erodium, Gilia capitata, Nemophila integrifolia, Lasthenia chrysostoma, Phacelia
douglasit.
Halictoides bernardina Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 401. 6, °.
boregoensis (Michener). Ariz., Calif. Pollen: Collects pollen from flowers of Camissonia
including C. campestris, C. clavaeformis, C. pallida halli, C. refracta, C. scapoidea, but
also visits flowers of Abronia villosa, Cryptantha barbigera, C. dumetorum, Larrea
tridentata, Malacothrix californica, Malva pusilla, Nama demissum, Phacelia
fremontii.
Halictoides boregoensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 398. 6.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 19 (floral
relationships). —Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 76 (floral
relationships). —Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 41
(floral relationships).

brevicornis Timberlake. Calif. Pollen: Unknown, but visits flowers of Cryptantha intermedia,
Linanthus dianthiflorus, Sphaenosciadium capitellatum.
Dufourea (Halictoides) brevicornis Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 403. ¢, °.
calientensis Timberlake. Calif. Pollen: Unknown, but visits flowers of Coreopsis bigelovit,
Eschscholzia californica, Gilia tricolor, Nemophila, Oenothera, Plagiobothrys
nothofulvus, Prunus subcordata.
Dufourea (Halictoides) calientensis Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 405. 6, @.
californica (Michener). Calif. Pollen: Unknown, but visits flowers of Lasthenia.
Halictoides californicus Michener, 1935. Pan-Pacific Ent. 11: 181. d.
calochorti calochorti (Cockerell). Calif. Pollen: Possibly an oligolege of Calochortus, visits
flowers of C. caeruleus var. maweanus.
Betheliella calochorti Cockerell, 1924. Ent. News 35: 170. 6.
calochorti sculleni (Cockerell). Oreg. Pollen: Possibly an oligolege of Calochortus.
Betheliella calochorti sculleni Cockerell, 1937. Canad. Ent. 69: 33. ¢.
campanulae (Cockerell). Wash. Pollen: Unknown, but visits flowers of Calochortus,
Campanula.
Halictoides campanulae Cockerell, 1897. Canad. Ent. 29: 289. 3.
crassipes (Cockerell). Oreg.
Halictoides crassipes Cockerell, 1924. Psyche 31: 243. d.
cuprea Bohart. Calif. Pollen: Unknown, but visits flowers of Gilia, Nemophila, Phacelia.
Dufourea cuprea Bohart, 1948. Ent. Soc. Amer., Ann. 41: 119. 6, 9.
davidsoni (Cockerell). Calif. Pollen: Unknown, but visits flowers of Eriogonum nudum,
Gayophytum nuttallu, G. ramosissimum.
Halictoides davidsoni Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 140. o.

Taxonomy: Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 199. d, @.
dentipes Bohart. Calif. Parasite: Neopasites fulviventris (Cress.)? Pollen: Unknown, but
apparently oligolege of Calochortus albus, C. monophyllus; also visits Clarkia
rhomboidea, Hypochoeris glabra, Wyethia angustifolia.
Dufourea dentipes Bohart, 1948. Ent. Soc. Amer., Ann. 41: 132. d, 9.

Biology: Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 144 (parasite).
desertorum Timberlake. South. Calif. (deserts). Pollen: Unknown, but visits flowers of
Eschscholzia including E. minutiflora var. darwinensis.
Dufourea (Halictoides) desertorum Timberlake, 1939. Ent. Soe. Amer., Ann. 32: 408. 3, 2.
dilatipes Bohart. Alta., Mont.
Dufourea dilatipes Bohart, 1948. Ent. Soe. Amer., Ann. 41: 1385. ¢, °.

Superfamily APOIDEA 1935

echinocacti Timberlake. South. Calif. (deserts). Pollen: Apparently an oligolege of
Echinocactus including E. acanthodes, E. cylindraceus.
Dufourea (Halictoides) echinocacti Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 399. d, °.
fallugiae (Cockerell). N. Mex. Pollen: Possibly an oligolege of Fallugia.
Halictoides fallugiae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 359. °.
femorata Bohart. Calif. Pollen: Presumably oligolege of Gilia including G. capitata, G. tricolor,
but also visits flowers of Collinsia, Cryptantha, Layia glandulosa, Nemophila, Salvia.
Dufourea femorata Bohart, 1947. Ent. Soc. Amer., Ann. 40: 701. d, 2.
fimbriata fimbriata (Cresson). Colo., N. Mex., Ariz., Calif. (high mountains). Pollen:
Apparently an oligolege of Potentilla including P. glandulosa, but also visits flowers of
Penstemon.
Panurgus fimbriatus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 63. °.
Halictoides Harveyi Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 223. 2.
fimbriata sierrae (Michener). Calif. (Sierra Nevada and White Mts.). Pollen: Unknown, but
visits flowers of Ranunculus.
Halictoides harveyi sierrae Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 397. 2.
gilia Bohart. Calif. (16 mi. E. Mt. Hamilton). Pollen: Unknown, but visits flowers of Gilia.
Dufourea gilia Bohart, 1947. Ent. Soc. Amer., Ann. 40: 692. 3, °.
holocyanea (Cockerell). Oreg., Calif. Pollen: Unknown, but visits flowers of Ceanothus,
Symphoricarpos.
Halictoides holocyaneus Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 199. d.
impunctata Bohart. Calif. (“So. Cal.”).
Dufourea impunctata Bohart, 1949. Ent. Soc. Amer., Ann. 42: 58. d.
latifrons Timberlake. South. Calif. (deserts). Pollen: Collects pollen from flowers of
Camissonia including C. brevipes, C. clavaeformis.
Dufourea (Halictoides) latifrons Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 409. ¢.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 19 (floral
relationships).
linanthi Timberlake. South. Calif. Pollen: Apparently an oligolege of Linanthus including L.
aureus, L. dianthiflorus, L. lemmonii, but also visits flowers of Malacothrix glabrata.
Dufourea (Halictoides) linanthi Timberlake, 1939. Ent. Soe. Amer., Ann. 32: 402. 6, 2.
longiceps Bohart. Calif. (Panamint Mts.). Pollen: Unknown, but visits flowers of Phacelia.
Dufourea longiceps Bohart, 1948. Ent. Soc. Amer., Ann. 41: 128. d, @.
macswaini Bohart. Calif. Pollen: Unknown, but visits flowers of Clarkia cylindrica, C.
dudleyana, C. purpurea.
Dufourea macswaini Bohart, 1969. Pan-Pacifie Ent. 45: 57. d, °.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 49 (floral
relationships).
malacothricis Timberlake. N. Mex., Ariz. and Calif. (deserts). Pollen: Presumably an oligolege
of Malacothriz including M. californica, M. glabrata, but also visits flowers of
Anisocoma acaulis, Camissonia pallida hallii, Cryptantha barbigera, Eschscholzia

minutiflora var. darwinensis, Phacelia fremontii.
Dufourea (Halictoides) malacothricis Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 406. d,
OF

Biology: Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 141-142 (nest
architecture, provisioning, development).
marginata halictella Michener. Wyo., Utah, Colo., N. Mex. Pollen: Unknown, but visits

flowers of the Compositae including Helianthus.
Panurgus halictulus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 63. ¢. Preoce.
Dufourea (Halictoides) marginata halictella Michener, 1951. In Muesebeck, Krombein,

Townes, U. S. Dept. Agr., Agr. Monog. 2: 1132. N. name.
Taxonomy: Cockerell, 1922. Amer. Mus. Novitates 40: 3.

1936 Hymenoptera in America North of Mexico

marginata marginata (Cresson). Minn., Wis. and Ill. to Miss., west to Alta., Colo. and Utah.
Pollen: Collects pollen from flowers of Helianthus annuus, but also visits flowers of
Bidens aristosa, Erigeron macranthus.
Panurgus marginatus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 62. ?.
Panurgus autumnalis Robertson, 1895. Amer. Ent. Soc., Trans. 22: 121. 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 516-518, figs.

Biology: Stephen, Bohart and Torchio, 1969. The biology and external morphology of bees, p.
122 (floral relationships).
maura (Cresson). N. Dak., Wyo., Idaho, Nebr., Colo., N. Mex. Pollen: Unknown, but visits
flowers of Campanula. Predator: Philanthus pulcher Dalla Torre.
Panurgus maurus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 61. “2” = 6.
monardae (Viereck). Wis., Mich., Ill. and Tenn. Pollen: Oligolege of labiates, collects pollen
from flowers of Agastache foeniculum, Monarda including M. fistulosa.
Conohalictoides monardae Viereck, 1924. Ent. Soe. Wash., Proc. 26: 14. g, 9.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 518-519, figs.

Biology: Bouseman, 1976. Kansas Ent. Soc., Jour. 49: 531-532 (floral relationships).
mulleri (Cockerell). Wyo., Calif., Ariz. Parasite: Eurystylops sp., Neoposites cressoni Cwfd.
Pollen: Oligolege of Phacelia including P. affinis, P. distans, P popei var. arizonica, P.
ramosissima, P. tanacetifolia, but also visits flowers of Camissonia brevipes,
Cryptantha including C. intermedia, Nemophila.
Halictoides mulleri Cockerell, 1898. Amer. Ent. Soe., Trans. 25: 190. °.

Taxonomy: Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 24. 6. —Michener, 1937. Ann. and
Mag. Nat. Hist. (10) 19: 400. 2, 6.

Biology: Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 132-138, figs.
1-6 (nest architecture, provisioning and development, adult activity, pollen source,
parasites).

nemophilae (Michener). Calif. Pollen: Presumably an oligolege of Phacelia including P.
davidsonii, but also visits flowers of Montia perfoliata, Nemophila integrifolia.
Halictoides nemophilae Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 400. 6, 2.
neocalifornica Bohart. Calif.
Dufourea neocalifornica Bohart, 1947. Ent. Soc. Amer., Ann. 40: 697. ¢.
novaeangliae (Robertson). Mo. and Mich. east to N. J. and Maine. Pollen: Unknown, but visits
flowers of Fagopyrum and Pontederia.
Panurgus novae-angliae Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 339. 3.
Conohalictoides lovelli Viereck, 1904. Ent. News 15: 245. 2, ¢.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 519-520, figs. (redescription).
nudicornis Timberlake. South. Calif. (deserts). Pollen: Presumably an oligolege of Onagraceae
including Camissonia clavaeformis, C. c. aurantiaca, C. piersonii, Oenothera deltoides,
but also visits flowers of Eschscholzia, Phacelia crenulata.
Dufourea (Halictoides) nudicornis Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 412. 2, 6.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 19 (floral
relationships).
oenotherae Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Camissonia
bistorta.
Dufourea (Halictoides) oenotherae Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 413. 6.
oryx (Viereck). B. C., Colo., Ariz., N. Mex.
Halictoides (Parahalictoides) oryx Viereck, 1903. Amer. Ent. Soc., Trans. 29: 48. ¢.

Taxonomy: Cockerell, 1916. Ent. News 27: 62. °.

pectinipes Bohart. Calif. Pollen: Unknown, but visits flowers of Gilia, Mimulus including M.
layneae.
Dufourea pectinipes Bohart, 1948. Ent. Soc. Amer., Ann. 41: 126. 6, 2.

Superfamily APOIDEA 1937

pulchricornis (Cockerell). N. Mex., Ariz. Pollen: Oligolege of Lesquerella, including L. gordonii.
Halictoides pulchricornis Cockerell, 1916. Ent. News 27: 63. 3, 2.

Biology: Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 142-143 (nest
architecture, provisioning, development, adult activity).
rhamni (Michener). South. Calif. Pollen: Unknown, but visits flowers of Camissonia bistorta,
Cryptantha intermedia, Dendromecon rigida, Eschscholzia californica, Rhamnus,
Plagiobothrys.
Halictoides rhamni Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 394. 6, °@.
sandhouseae arida Bohart. South. Calif. Pollen: Unknown, but visits flowers of Phacelia.
Dufourea sandhouseae arida Bohart, 1949. Ent. Soe. Amer., Ann. 42: 61. 6, 9.
sandhouseae sandhouseae (Michener). Calif.; Mexico (Baja California). Pollen: Unknown, but
visits a wide variety of flowers including Arenaria douglasii, Agoseris heterophylla,
Calandrinia menziesii, Camissonia bistorta, C. cheiranthifolia, C. clavaeformis
aurantiaca, C. parishii, Chaenactis, Coreopsis californica, Cryptantha intermedia,
Convolvulus occidentalis, Dendromecon rigida, Encelia farinosa, Erodium botrys,
Eschscholzia californica, Gilia multicaulis, G. tricolor, Hesperochiron californicus,
Lasthenia chrysostoma, Layia elegans, L. platyglossa, Lessingia, Linanthus aureus,
Lomatium, Malacothrix californica, Nemophila menziesii, Phacelia, Prunus andersonii,
Salvia columbariae, Sisymbrium irio, Stenotopsis linearifolius.
Halictoides sandhouseae Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 328. d, 9.
saundersi (Cockerell). Calif. (Central Valley). Pollen: Unknown, but visits flowers of
Camissonia campestris.
Halictoides saundersi Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 189. 2, 6.
scabricornis Bohart. Calif., Oreg., Utah. Pollen: Unknown, but visits flowers of Gayophytum,
Symphoricarpos. Predator: Philanthus pacificus arizonae Dunning, P. pulcher Dalla
Torre.
Dufourea scabricornis Bohart, 1949. Ent. Soc. Amer., Ann. 42: 57. d, °.
scintilla (Cockerell). South. Calif. Pollen: Presumably an oligolege of Camissonia including C.
bistorta, C. campestris, C. cheiranthifolia, C. contorta, C. dentata, C. micrantha, C.
pallida hallii, C. sierrae, C. veitchiana, but also visits flowers of Clarkia, Cryptantha
intermedia, C. muricata, Coreopsis californica, Eschscholzia californica, Euphorbia
albomarginata, Phacelia douglasii, Plagiobothrys, Trifolium variegatum.
Diandrena scintilla Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 50. °.
Halictoides eschscholtziae Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 393. d, 2.

Biology: Linsley, MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 71: 41-42 (floral
relationships).
spilura (Cockerell). Calif. Pollen: Unknown, but visits flowers of Eriogonuwm nudum,
Gayophytum, Oenothera.
Halictoides spilurus Cockerell, 1925. Calif. Acad. Sci., Proce. (4) 14: 200. °.
spinifera (Viereck). Calif., Nev. Pollen: Unknown, but visits flowers of Arenaria kingii,
Sidalcea apicata, Trifoliwm monanthum.
Cryptohalictoides spiniferus Viereck, 1904. Ent. News 15: 261. d.
Taxonomy: Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 199. 6, 2. —Michener, 1937. Ann.
and Mag. Nat. Hist. (10) 19: 402. °.
subdavidsoni Bohart. Calif. Pollen: Unknown, but visits flowers of Gayophytum diffusum.
Dufourea subdavidsoni Bohart, 1949. Ent. Soe. Amer., Ann. 42: 55. 6, 2.
tarsata Bohart. Calif. Pollen: Unknown, but visits flowers of Cryptantha, Gilia, Linanthus
parviflorus, Lupinus, Nemophila.
Dufourea tarsata Bohart, 1947. Ent. Soc. Amer., Ann. 40: 698. 4, 2.

timberlakei Bohart. Calif. (Mt. Pinos).
Dufourea timberlakei Bohart, 1947. Ent. Soc. Amer., Ann. 40: 700. d.

tinsleyi (Cockerell). N. Mex.
Halictoides tinsleyi Cockerell, 1898. Canad. Ent. 30: 52. °.

1938 | Hymenoptera in America North of Mexico

trochantera Bohart. Wash., Oreg., Wyo., Utah, Calif. Parasite: Neopasites sp. Pollen: Oligolege
of Phacelia ineluding P. leucophylla, P. pringlei, but also visits flowers of Calochortus

venustus.
Dufourea trochantera Bohart, 1948. Ent. Soc. Amer., Ann. 41: 121. 6, °.

Biology: Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 138-141 (nest
architecture, provisioning, development, adult activity, pollen source, parasite).
truncata Timberlake. South. Calif. Pollen: Presumably an oligolege of Camissonia including C.
bistorta, C. dentata, C. veitchiana, but also visits flowers of Astragalus, Coreopsis
californica, Cryptantha intermedia.
Dufourea (Halictoides) truncata Timberlake, 1939. Ent. Soe. Amer., Ann. 32: 410. 6, 9.
tularensis Timberlake. Calif. (Three Rivers). Pollen: Unknown, but visits flowers of
Anisocoma acaulis, Eschscholzia californica, Layia glandulosa, Phacelia distans, P.
~ douglasii.
Dufourea (Halictoides) tularensis Timberlake, 1941. Ent. Soc. Amer., Ann. 34: 41. dg, 9.
tuolumne Bohart. Calif. Pollen: Unknown, but visits flowers of Cryptantha, Gilia, Linanthus
filipes, L. montanus, Lomatium.
Dufourea tuolumne Bohart, 1947. Ent. Soc. Amer., Ann. 40: 694. 6, °.
vanduzeei Bohart. Calif. (Monterey Co.).
Dufourea vanduzeei Bohart, 1947. Ent. Soc. Amer., Ann. 40: 696. 6.
vandykei Bohart. Calif. Pollen: Unknown, but visits flowers of Sphaeralcea ambigua.
Dufourea vandykei Bohart, 1948. Ent. Soc. Amer., Ann. 41: 124. 6, 9.
vernalis Timberlake. Calif. (deserts). Pollen: Evidently polylectic, usually visits flowers of
Eschscholzia including E. minutiflora, E. parishii, but also visits other flowers
including Argemone, Camissonia brevipes, C. clavaeformis, C. refracta, Malacothrix
glabrata, Mentzelia involucrata, Phacelia distans.
Dufourea (Halictoides) vernalis Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 400. d, °.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 19 (floral
relationships).
versatilis rubriventris Michener. Calif.; Mexico (Baja California). Pollen: Oligolege of Mimulus
including M. diffusus, M. fremontii, M. guttatus, M. layneae, M. montioides, M.
moschatus, M. primuloides, M. suksdorfi, but also visits flowers of Gilia, Nemophila.
Mimulapis versatilis rufiventris Timberlake, 1941. Ent. Soe. Amer., Ann. 34: 38. 2, 6.
Preoce.
Dufourea (Halictoides) rubriventris Michener, 1951. In Muesebeck, Krombein, Townes, U.
S. Dept. Agr., Agr. Monog. 2: 1133. N. name.
versatilis versatilis (Bridwell). South. Calif. Pollen: Apparently an oligolege of Mimulus, but
visits flowers of Gilia achillaefolia, G. splendens.
Mimulapis versatilis Bridwell, 1919. Hawaii. Ent. Soc., Proc. 4: 163. 6, °.
virgata (Cockerell). Calif. Pollen: Collects pollen from flowers of Camissonia boothii, but visits
a wide variety of flowers including Amsinckia, Brodiaea capitata, Camissonia
campestris, Convolvulus villosus, Coreopsis bigelovii, C. californica, Cryptantha
intermedia, C. muricata, Descurainia sophia, Encelia californica, E. farinosa,
Eriophyllum lanatum, Eschscholzia californica, Euphorbia albomarginata, Gilia
achillaefolia, G. multicaulis, G. tenuifolia, G. tricolor diffusa, Hesperochiron, Layia
elegans, L. glandulosa, Linanthus aureus, Lotus scoparius, Lupinus, Malacothriz,
Nemophila, Phacelia douglasii, Plagiobothrys nothofulvus, Platystemon, Salvia
columbariae, Stenotopsis linearifolius, Trifoliwm.
Halictoides virgatus Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 191. 2, 6.

Taxonomy: Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 140.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 19 (floral
relationships).

viridescens (Crawford). Calif.
Halictoides viridescens Crawford, 1916. Insecutor Inscitiae Menstruus 4: 139. 6.

Superfamily APOIDEA = 1939

Genus MICRALICTOIDES Timberlake

Dufourea subg. Micralictoides Timberlake, 1939. Ent. Soc. Amer., Ann. 32: 397.
Type-species: Halictoides ruficaudus Michener. Orig. desig.
Revision: Bohart, 1942. Pan-Pacifie Ent. 18: 119-123.

altadenae (Michener). South. Calif. Pollen: Unknown, but visits flowers of Chaenactis
glabriuscula, Eriophyllum confertiflorum.
Halictoides altadenae Michener, 1936. Ann. and Mag. Nat. Hist. (10) 19: 395. 2, 6.

mojavensis Bohart. South. Calif. Pollen: Unknown, but visits flowers of Chaenactis
brachypappa, Gilia multicaulis, Eschscholzia californica, Lasthenia gracilis, Layia
platyglossa, Malacothrix californica, M. glabrata, Phacelia fremontii, Salvia
columbariae.
Micralictoides mojavensis Bohart, 1942. Pan-Pacifie Ent. 18: 120. d, 2.

ruficaudus (Michener). Calif. Pollen: Apparently an oligolege of Eschscholzia californica, but
also visits flowers of Gilia multicaulis, Nemophila menziesii.
Halictoides ruficaudus Michener, 1936. Ann. and Mag. Nat. Hist. (10) 19: 397. 9.

Genus PROTODUFOUREA Timberlake

Protodufourea Timberlake, 1955. Pan-Pacific Ent. 31: 105.
Type-species: Protodufourea wasbaueri Timberlake. Orig. desig.
Revision: Timberlake, 1955. Pan-Pacific Ent. 31: 105-108.
parca Timberlake. Calif. (Riverside Co.). Pollen: Oligolege of Phacelia including P. distans, P.
hispida.
Protodufourea parca Timberlake, 1955. Pan-Pacifie Ent. 31: 106. d.

wasbaueri Timberlake. Calif. (San Benito Co.). Pollen: Oligolege of Emmenanthe penduliflora.
Protodufourea wasbaueri Timberlake, 1955. Pan-Pacifie Ent. 31: 107. d, °.

Genus CONANTHALICTUS Cockerell

Revision: Timberlake, 1961. Pan-Pacific Ent. 37: 145-160.
Biology: Rozen and McGinley, 1976. Amer. Mus. Novitates 2602: 1-6, 3 figs.

Genus CONANTHALICTUS Subgenus CONANTHALICTUS Cockerell

Halictus subg. Conanthalictus Cockerell, 1901. Ent. News 12: 209.
Type-species: Halictus (Conanthalictus) conanthi Cockerell. Monotypice.
conanthi (Cockerell). N. Mex. Pollen: Unknown, but visits flowers of Nama hispidum.
Halictus (Conanthalictus) conanthi Cockerell, 1901. Ent. News 12: 208. 9.

cotullensis Crawford. Tex. (Cotulla).
Conanthalictus cotullensis Crawford, 1907. N. Y. Ent. Soc., Jour. 16: 182. 2, ¢

Genus CONANTHALICTUS Subgenus PHACELIAPIS Michener

Conanthalictus subg. Phaceliapis Michener, 1942. N. Y. Ent. Soc., Jour. 50: 277.
Type-species: Conanthalictus bakeri Crawford. Orig. desig.
bakeri Crawford. South. Calif. Pollen: Unknown, but visits flowers of Phacelia distans, P.
tanacetifolia.
Conanthalictus Bakeri Crawford, 1907. Invertebrata Pacifica 1: 197. °.
caerulescens Timberlake. Calif. (Riverside Co.), Nev. Pollen: Unknown, but visits flowers of
Phacelia crenulata, P. distans.
Conanthalictus caerulescens Timberlake, 1961. Pan-Pacifie Ent. 37: 148. d, °.
cockerelli Timberlake. Calif. (San Bernardino Co.). Pollen: Unknown, but visits flowers of
Eriodictyon, Phacelia fremontii.
Conanthalictus cockerelli Timberlake, 1961. Pan-Pacifie Ent. 37: 155. d, 2.

1940 Hymenoptera in America North of Mexico

deserticola Timberlake. South. Calif. (Imperial and Riverside Cos.). Pollen: Unknown, but
visits flowers of Camissonia brevipes, Nama demissum.

Conanthalictus deserticola Timberlake, 1961. Pan-Pacifie Ent. 37: 152. ¢. Described under
heading Conanthalictus namatophilus Timberlake, new species which is a printer’s
error according to Timberlake in litt.; also see Pan-Pacific Ent. 40: 200, 1964.

macrops Cockerell. South. Calif. (San Gabriel and San Bernardino Mts.). Pollen: Unknown, but
visits flowers of Malacothrix, Phacelia davidsonii, P. douglasii, Yucca whipplei.

Conanthalictus macrops Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 51. 6.

mentzeliae Timberlake. South. Calif. (San Bernardino, Kern, and Inyo Cos.), Nev. Pollen:
Unknown, but visits flowers of Malacothrix, Mentzelia affinis, M. albicaulis, M.
micrantha, Phacelia.

Conanthalictus mentzeliae Timberlake, 1961. Pan-Pacifie Ent. 37: 156. d, 2.

minor Timberlake. South. Calif. (San Bernardino, Riverside and Inyo Cos.). Pollen: Unknown,
but visits flowers of Camissonia clavaeformis, Nama demissum, Phacelia.

Conanthalictus minor Timberlake, 1961. Pan-Pacifie Ent. 37: 151. 6, 2. Described under
heading Conanthalictus deserticola Timberlake, new species which is a printer’s error
according to Timberlake zn litt.; also see Pan-Pacific Ent. 40: 200, 1964.

namatophilus Timberlake. South. Calif. (Imperial, Riverside and San Bernardino Cos.). Pollen:
Unknown, but visits flowers of Camissonia clavaeformis, Lotus scoparius, Nama
demissum, Phacelia aff. fremontii.

Conanthalictus namatophilus Timberlake, 1961. Pan-Pacifie Ent. 37: 153. 6, 2. Described
under heading Conanthalictus minor Timberlake, new species which is a printer’s error
according to Timberlake in litt.; also see Pan-Pacific Ent. 40: 200, 1964.

nigricans Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Cryptantha
intermedia, Emmenanthe, Phacelia brachyloba, P. distans, P. douglasii, P. grandiflora,
P. minor.
Conanthalictus nigricans Timberlake, 1961. Pan-Pacific Ent. 37: 149. 6, 2.
seminiger Michener. South Calif. Pollen: Collects pollen from flowers of Emmenanthe
penduliflora, but also visits flowers of Rhamnus crocea.
Conanthalictus seminiger Michener, 1937. Ann. and Mag. Nat. Hist. 10 (19): 326. 9.
wilmattae Cockerell. South. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers of
Phacelia distans, Prosopis juliflora.
Conanthalictus wilmattae Cockerell, 1936. Amer. Mus. Novitates 831: 2. °.

Genus SPHECODOSOMA Crawford

Sphecodosoma Crawford, 1907. N. Y. Ent. Soc., Jour. 15: 182.
Type-species: Sphecodosoma pratti Crawford. Monotypic and orig. desig.
Taxonomy: Timberlake, 1961. Pan-Pacific Ent. 37: 158-160 (as subgenus).

dicksoni (Timberlake). South. Calif., Tex. (Davis Mts); Mexico (Durango). Pollen: Unknown,
but visits flowers of Coldenia, Nama demissum, N. hispidum.
Conanthalictus dicksoni Timberlake, 1961. Pan-Pacific Ent. 37: 158. °.

pratti Crawford. Tex. Pollen: Unknown, but visits flowers of Lesquerella.
Sphecodosoma pratti Crawford, 1907. N. Y. Ent. Soc., Jour. 15: 183. 2, ¢.

Genus MICHENERULA Bohart

Michenerula Bohart, 1965. Ent. Soc. Amer., Ann. 58: 320.
Type-species: Michenerula beameri Bohart. Monotypic and orig. desig.

beameri Bohart. Tex. (Marathon). Pollen: Unknown, but visits flowers of Nama hispidum.
Michenerula beameri Bohart, 1965. Ent. Soe. Amer., Ann. 58: 320. d, 2.

Genus XERALICTUS Cockerell

Xeralictus Cockerell, 1927. Pan-Pacifie Ent. 4: 41.
Type-species: Xeralictus timberlakei Cockerell. Monotypic.

Taxonomy: Eickwort, 1969. Ent. Soc. Amer., Ann. 62: 654 (subfamilial assignment).

Superfamily APOIDEA 194]

timberlakei Cockerell. Calif. (Colorado Desert). Pollen: Oligolege of Mentzelia involucrata, but
also visits other flowers for nectar including Mohavea confertiflora, Phacelia crenulata,
P. distans.
Xeralictus timberlakei Cockerell, 1927. Pan-Pacifie Ent. 4: 42. 3.

SUBFAMILY NOMIINAE

Members of this subfamily are especially numerous in the Old World tropics including those of
the Australian Region. They are relatively poorly represented in the Holarctic Region. Only the
widespread genus Noma is present in the Nearctic Region and apparently all of the species are
polyleges, obtaining their pollen and nectar from the flowers of a wide variety of both native
and introduced plants. However, some of the species (e.g., Nomia nevadensis Cresson and N. tri-
angulifera Vachal) appear to collect pollen only from the flowers of the Compositae. At least one
of the species, the alkali bee (N. melanderi Cockerell), is an exceptionally valuable pollinator of
alfalfa (Medicago sativa) and has been managed successfully in artificial nesting sites so as to
enhance the pollination of this important agricultural crop.

Genus NOMIA Latreille

Genus NOMIA Subgenus NOMIA Latreille

Nomia Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, p. 182.
Type-species: Andrena curvipes Fabricius. Monotypice.
Nitocris Rafinesque, 1815. Analyse de la nature, p. 123. Proposed unnecessarily for Nomia
Latreille.

This subgenus is found only in the Old World.
Genus NOMIA Subgenus CURVINOMIA Michener

Nomia subg. Paranomia Friese, 1897. Festschr. fuenfzig. bestehene Ver. Schles.
Insektenk. Breslau, p. 48. Preocc.
Type-species: Nomia chalybeata Smith. Desig. by Cockerell, 1910.
Nomia subg. Paranomina Michener, 1944. Amer. Mus. Nat. Hist., Bul. 82: 251. N. name.
Preoce.
Nomia subg. Curvinomia Michener, 1944. Amer. Mus. Nat. Hist., Bul. 82: 251.
Type-species: Nomia californiensis Michener. Monotypic and orig. desig. (=Nomia
tetrazonata Cockerell).

Revision: Ribble, 1965. Kansas Univ. Sci. Bul. 45: 277-309, figs. (N. Amer. spp.).

Taxonomy: Ribble, 1968. Wyo. Univ. Agr. Expt. Sta., Sci. Monog. 11: 15-18 (tax. notes, geogr.
and floral records).
angustitibialis Ribble. Southeastern Ariz. to Guerrero, Mex. Pollen: Unknown, but visits
flowers of Baccharis glutinosa, Condalia lycioides, Larrea tridentata, Melilotus alba.
Nomia (Curvinomia) angustitibialis Ribble, 1965. Kansas Univ. Sci. Bul. 45: 292. d, °.
fedorensis Cockerell. Southwestern Kans., Okla., Tex., south to Veracruz, Mexico. Pollen:
Unknown, but visits flowers of Cassia, Dalea grisea, Gaillardia, Helianthus, Mimosa,
Rhynchosia americana, Tephrosia virginiana.
Nomia fedorensis Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 6: 277. 9, d.

Biology: Cockerell, 1934. Amer. Mus. Novitates 697: 4 (nest).
foxii Dalla Torre. Colo. (Denver), S. Dak. (Big Stone City). Senior synonym of either Nomia
universitatis Cockerell or N. tetrazonata Cockerell, see Hurd and Linsley, 1974. Ent.
Soc. Wash., Proc. 76: 198.
Nomia punctata Fox, 1893. Ent. News, 4: 135. 2. Preoce.
Nomia foxii Dalla Torre, 1896. Cat. Hym., v. 10, p. 167. N. name.

Taxonomy: Hurd and Linsley, 1974. Ent. Soc. Wash., Proc. 76: 198 (tax. status).

maneei Cockerell. N. J., N. C. to Fla. west to Okla. and Ala. Parasite: Psewdomethoca
vanduzeei Bradley. Pollen: Unknown, but visits flowers of Asclepias tuberosa,

1942 Hymenoptera in America North of Mexico

Desmodium ciliare, Eriogonum baileyi, Galactia, Lespedeza capitata, L. repens,
Phaseolus polystachios, Strophostyles.
Nomia maneei Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 6: 276. 2.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 508, figs. (redescription).

mesillensis Cockerell. South. Colo., Utah, Ariz., N. Mex., Tex., south to Jalisco, Mexico. Pollen:
Polylege of a wide variety of flowers including Aloysia wrightii, Asclepias linaria, A.
subverticillata, Baccharis glutinosa, Cevallia sinuata, Cirsium, Cleome jonesi, C.
serrulata, Crotalaria pumila, Croton, Crusea subulata, Eriogonum abertianum, E.
annuum, Eysenhardtia polystacha, Gossypium, Guardiola tulocarpa, Gutierrezia
sarothrae, Haplopappus, Helianthus annuus, Koeberlinia spinosa, Larrea tridentata,
Lepidium thurberi, Lotus wrightii, Medicago sativa, Melilotus alba, M. officinalis,
Mentzelia pumila, Nolina, Petalostemon candidum, Psoralea tenuifolia, Solanum
elaeagnifolium, S. rostratum, Verbesina encelioides. Predator: Blepharepium secabile
(Walk.), Mallophora bromleyi Curran.

Nomia mesillensis Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 334. °.

Taxonomy: Hurd and Linsley, 1974. Ent. Soc. Wash., Proc. 76: 198-199 (tax. status).

tetrazonata tetrazonata Cockerell. South. Calif. and Nev. to N. Mex. south into Mexico (Baja

California and Sonora). Pollen: Polylege of a wide variety of flowers including Acacia,
Allionia incarnata, Antigonon, Asclepias, Astragalus preussii, Baccharis glutinosa, B.
sarothroides, Boerhaavia, Cercidium floridum, Chrysothamnus, Cissus trifoliata,
Cleome, Cleomella longipes, Colubrina, Condalia lycioides, Croton californicus,
Cuscuta, Dalea, Encelia farinosa, Eriogonum fasciculatum, E. heermannii, E.
inflatum, E. nodosum, Euphorbia, Glycine max, Gossypium, Gutierrezia sarothrae,
Heliotropium curassavicum, Heterotheca subaxillaris, Hymenothrix wislizenii, Hyptis
emoryi, Kallstroemia grandiflora, Larrea tridentata, Lepidium thurberi, Lotus
scoparius, Medicago sativa, Melaleuca, Melilotus alba, Nolina parryi, Pectis papposa,
Penstemon palmeri, Philbertia heterophylla, Prosopis juliflora, Sapindus saponaria,
Solanum elaeagnifolium, S. rostratum, Stanleya pinnata, Tamarix gallica, Verbesina
encelioides, Wislizenia refracta, Zea mays.

Nomia tetrazonata Cockerell, 1910. U. S. Natl. Mus., Proc. 38: 297. 6.

Nomia moctezumae Crawford, 1911. U.S. Natl. Mus., Proc. 39: 618. °.

Nomia californiensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 318. 9.

tetrazonata uvaldensis Cockerell. N. Mex., Tex.; Mexico. Pollen: Unknown, but visits flowers

of Acacia greggii, Baccharis glutinosa, Cercidium texanum, Melilotus alba,
Parkinsonia aculeata, Prosopis juliflora, Solanum elaeagnifolium, Teucrium cubense.

Nomia uvaldensis Cockerell, 1930. In Cockerell and Blair, Amer. Mus. Novitates 433: 13.

oF
Nomia zabriskii Cockerell and Blair, 1930. Amer. Mus. Novitates 433: 18. 6.
Nomia parksi Cockerell, 1934. Amer. Mus. Novitates 697: 7. 2.
universitatis Cockerell. N. Dak. to Tex. between Mississippi River and Rocky Mts. Pollen:
Unknown, but visits flowers of Amorpha canescens, Astragalus gracilis, Dalea,
Medicago sativa, Melilotus alba, M. officinalis, Petalostemon purpureum, Psoralea
tenuiflora, Rudbeckia, Symphoricarpos occidentalis. Predator: Diogmites

angustipennis Loew.
Nomia universitatis Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 334. d.

Genus NOMIA Subgenus ACUNOMIA Cockerell

Nomia subg. Acunomia Cockerell, 1930. In Cockerell and Blair, Amer. Mus. Novitates
433: 11.
Type-species: Nomia nortoni Cresson. Orig. desig.

Revision: Ribble, 1965. Kansas Univ. Sci. Bul. 45: 282-283, 309-359, figs. (N. Amer. spp.,
includes summary of biological literature).

Taxonomy: Ribble, 1968. Wyo. Univ. Agr. Expt. Sta., Sci. Monog. 11: 1-15 (alkali bee and
related species, including summary of biological literature).

Superfamily APOIDEA 1943

howardi Crawford. South. Utah, south to South. Calif. and southwestern Ariz.; Mexico (Baja

California). Pollen: Polylege of a wide variety of flowers including Acacia greggii, Aster
spinosus, Carthamnus tinctorius, Cucumis melo, Gossypium, Heliotropium
curassavicum, Isocoma veneta, Lippia lanceolata, Medicago sativa, Melilotus alba,
Phyla nodiflora, Pluchea sericea, Prosopis pubescens, Salix exigua, Tamarix gallica, T.
petandra. Predator: Imparipes americanus (Banks).

Nomia howardi Crawford, 1911. U. S. Natl. Mus., Proce. 39: 617. @.

Nomia howardi vanduzeei Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 540. °.

Nomia californica peninsularis Cockerell, 1930. In Cockerell and Blair, Amer. Mus.
Novitates 433: 12.

Biology: Linsley, 1946. Jour. Econ. Ent. 6: 18-29 (pollination).
melanderi Cockerell. Centr. Wash. to South. Calif., east to Wyo. Introduced into S. Dak.

(Brookings and Heela) and B. C. (Ashcroft and Kamloops). Parasite: Dasymutilla vesta
(Cress.)?, Euphytomima nomiivora James, Heterostylum robustum O. S., Nemognatha
sp., Nomada suavis Cress., Physocephala texana (Will.), Physoconops fronto (Will.),
Pseudomethoca propinqua (Cress.)?, Zodion obliquefasciatum (Macq.), Zonitis
atripennis flavida LeC. Pollen: Polylege of a wide variety of plants especially the
Leguminosae including alfalfa, but also visits the flowers of Achillea, Allium, Apium
graveolens, Asclepias speciosa, A. subverticillata, Aster, Astragalus, Baccharis emoryi,
Berula erecta, Beta vulgaris, Brassica geniculata, B. incana, Centromadia pungens,
Chrysothamnus nauseosus, Cirsium, Citrullus vulgaris, Cleome lutea, C. serrulata,
Convolvulus arvense, Coreopsis, Crepis glauca, C. runcinata, Cressa cretica, Croton
californicus, Cuscuta salina, Daucus carota, Dipsacus, Eremocarpus setigerus,
Erigeron, Eriogonum, E'schscholzia californica, Gossypium, Grindelia, Haplopappus
acradenius, Helianthus, Heliotropium curassavicum, Hemizonia pungens, Lippia
lanceolata, Lotus americanus, L. purshianus, Melilotus alba, M. indica, Mentha spicata,
Phyla lanceolata, Pluchea camphorata, P. sericea, Polygonum lapathifolium, Ribes,
Salsola kali, Sicyos, Sisymbrium altissimum, Solanum elaeagnifolium, S. tuberosum,
Solidago occidentalis, Tamarix gallica, Tribulus terrestris, Trifolium hybridum, T.
pratense, T. repens, Wislizenia refracta, Zea mays. Predator: Badgers, blackbirds,
Cicindela haemorrhagica LeC., C. pusillus Say, Efferia straminea (Will.), Formica
fusca Linn., Imparipes americanus (Banks), magpies, meadowlarks, Microtus sp., Mus
musculus Linn., Pogonomyrmesx occidentalis (Cress.), robins, skunks, English sparrows,
starlings, weasels.

Nomia melandri Cockerell, 1906. Canad. Ent. 38: 279. 2°. Lapsus calami.

Nomia melanderi Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 334. Emend.

Nomia californica Cockerell, 1910. U. S. Natl. Mus., Proc. 38: 296. 9.

Nomia acus Cockerell, 1910. U. S. Natl. Mus., Proc. 38: 296. ¢.

Nomia melanderi paysoni Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 621. d.

Taxonomy: Michener, 1953. Kansas Univ. Sci. Bul. 35: 1031, figs. (larva).

Biology: Stephen and Evans, 1960. Oreg. Agr. Expt. Sta. Tech. Bul. 52: 1-39, 5 figs, 2 tables
(life history, parasites, associates). —Frick, 1962. Ent. Soc. Amer., Ann. 55: 5-15 (ecology,
parasite). —Hackwell and Stephen, 1966. Pan-Pacific Ent. 42: 196-200 (eclosion and
duration of larval development). —Ribble, 1968. Wyo. Univ. Agr. Expt. Sta., Sci. Monog.
11: 2-11 (summary of literature). —Cross and Bohart, 1969. Kansas Ent. Soc., Jour. 42:
195-219, 6 figs., 5 tables (phoretic behavior of associated mites). —Telford, Johansen and
Eves, 1972. Mededel. Fakult. Landbouwwetensch. Gent 37: 776-783 (management practices
and insecticide poisoning). —Parker and Potter, 1974. Environ. Ent. 3: 739-743, 5 figs.
(introduction and establishment). —Batra, 1976. Kans. Ent. Soc., Jour. 49: 18-22, 2 tables
(comparative efficiency in alfalfa pollination). —Mayer and Johansen, 1976. Ent. Soc.
Amer., Bul. 22: 423-425 (bibliography).

nortoni cressoni Westwood. Ariz. (Santa Cruz Co.), south to Puebla, Mexico. Pollen: Unknown,
but visits flowers of Asclepias, Dalea, Eysenhardtia polystacha, Stachys bigelovii.
Nomia Cressoni Westwood, 1875. Ent. Soc. London, Trans., p. 218. ¢.
nortoni nortoni Cresson. Penn., south to Fla., west to Idaho, Colo. and N. Mex.; Mexico
(Coahuila). Pollen: Polylege of a wide variety of flowers including Amorpha fruticosa,

1944. Hymenoptera in America North of Mexico

Aster, Borreria, Brazoria truncata, Callirhoe leiocarpa, Cassia cinerea, C. fasciculata,
Cirsium altissimum, C. vulgare, Clethra, Coreopsis cardaminefolia, Dalea aurea, D.
multiflora, Desmodium paniculatum, Engelmannia pinnatifida, Eryngium
yuccifolium, Euipomoea, Euphorbia marginata, Gaillardia, Gossypium, Grindelia
squarrosa, Helenium autumnale, Helianthus maximiliani, Ligustrum, Lyciwm
halimifolium, Marrubium vulgare, Medicago sativa, Melilotus alba, Mentha longifolia,
Monarda citriodora, M. punctata, Oenothera biennis, Petalostemon microphyllum, P.
purpureum, Phacelia, Polygonum, Pontederia cordata, Prosopis, Pycnanthemum
flexuosum, P. virginianum, Ratibida columnifera, R. pinnata, Rhus copalina, Solanum
rostratum, S. torryei, Solidago rigida, Verbena officinalis, Yucca glauca.

Nomia nortoni Cresson, 1868. Amer. Ent. Soc., Trans. 1: 385. 2, 6.

Nomia nortoni var. plebeia Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 6: 276. °.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 510, figs. (redescription).

Biology: Cockerell, 1934. Amer. Mus. Novitates 697: 7. —Ribble, 1965. Kansas Univ. Sci. Bul.
45: 316-318 (nesting habits, parasites, predators).

Genus NOMIA Subgenus EPINOMIA Ashmead

Epinomia Ashmead, 1899. ‘Amer. Ent. Soe., Trans. 26: 88.
Type-species: Nomia triangulifera Vachal. Monotypic and orig. desig. (=Nomia
persimilis Cockerell).

Revision: Cross, 1958. Kansas Univ. Sci. Bul. 38: 1261-1301, 4 figs. (included spp., contains
biological information).
micheneri Cross. South. Ariz. and Tex., south to Jalisco and Guanajuato, Mexico. Pollen:
Unknown, but males visit flowers of Eysenhardtia polystachya, Guardiola tulocarpa,
Helianthus.
Nomia (Epinomia) micheneri Cross, 1958. Kansas Univ. Sci. Bul. 38: 1273. d, °.
nevadensis angelesia Cockerell. Oreg., Calif., Nev.; Mexico (extreme northwest Baja
California). Parasite: Rhipiphorus epinomiae Linsley and MacSwain. Pollen: Polylege of
a wide variety of flowers including Aster spinosa, Brassica incana, Calendula,
Centromadia pungens, Chrysothamnus, Coreopsis lanceolata, Croton californicus,
Erigeron canadense, E. linifolius, Eriogonum fasciculatum, E. gracile, Grindelia
camporum, Gnaphalium beneolens, Gutierrezia californica, G. sarothrae, Haplopappus
palmeri, H. vernonioides, Helianthus gracilentus, Heterotheca grandiflora,
Lepidospartum squamatum, Lippia filiformis, Lotus americanus, Marrubium vulgare,
Medicago sativa, Melilotus alba, Pectis papposa, Pluchea camphorata, Polygonum
lapathifolium, Senecio douglasii, Solidago occidentalis, Stephanomeria exigua, S.
virgata, Tamarix gallica, Trichostema, Wislizenia refracta.
Nomia arizonensis angelesia Cockerell, 1910. U. S. Natl. Mus., Proe. 38: 293. 6.

Taxonomy: Michener, 1953. Kansas Univ. Sci. Bul. 35: 1029, figs. (larva).

nevadensis arizonensis Cockerell. Utah, Ariz., south. Calif. (Colorado Desert), N. Mex., south
to Jalisco, Mexico. Pollen: Polylege of a wide variety of flowers including Asclepias,
Baccharis glutinosa, Chrysothamnus, Cleome serrulata, Eriogonum, Euphorbia,
Eysenhardtia polystacha, Grindelia squarrosa, Guardiola tulocarpa, Gutierrezia
microcephala, G. sarothrae, Haplopappus acradenius, H. spinulosus, Helianthus,
Hymenothrix wislizeni, Hymenoxys, Lygodesmia juncea, Pectis papposa, Psilostrophe
cooperi, Solanum elaeagnifolium, Sphaeralcea, Verbesina encelioides, Wislizenia
refracta. Predator: Philanthus ventilabris Fabr.

Nomia arizonensis Cockerell, 1899. Entomologist, 32: 128. d.

Biology: Aleock and Gamboa, 1975. Ariz. Acad. Sci. 10: 163 (predator).
nevadensis bakeri Cockerell. Fla. west to Wyo., Colo., N. Mex.; Mexico (Tamaulipas). Parasite:
Rhipiphorus nomiae Rivnay. Pollen: Polylege of a wide variety of flowers including
Aster, Bidens involucrata, Boltonia asteroides, Chrysopis microcephala, Coreopsis,
Dalea lanata, Grindelia, Haplopappus, Helenium, Heterotheca, Isopappus, Melilotus
alba, Prionopsis, Prosopis, Solidago, Thelesperma gracile, Verbesina encelioides.
Nomia bakeri Cockerell, 1898. Entomologist, 31: 32. ¢.

Superfamily APOIDEA 1945

Nomia pattoni Cockerell, 1910. U. S. Natl. Mus., Proc. 38: 292. 2, d.
Nomia bakeri var. rufibasis Cockerell, 1930. In Cockerell and Blair, Amer. Mus. Novitates
433: 14.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 512-518, figs. (redescription).

Biology: Hicks, 1926. Colo. Univ., Studies 15: 222. —Cockerell, 1934. Amer. Mus. Novitates
697: 4. —Kerfoot, 1964. Kansas Ent. Soc., Jour. 37: 152-157, 1 fig.
nevadensis nevadensis Cresson. Nev. (Nye and White Pine Cos.).
Nomia nevadensis Cresson, 1874. Amer. Ent. Soc., Trans. 5: 101. 2, d.
triangulifera Vachal. N. Dak., Minn. and III. west to Utah and N. Mex. Parasite: Rhipiphorus
solidaginis (Pierce), Triepeolus mesillae Cockerell. Pollen: Unknown, but visits flowers
of Bidens involucrata, Cleome serrulata, Gilia, Grindelia squarrosa, Helianthus
annuus, H. lenticularis, H. maximiliani, H. petiolaris, Medicago sativa, Polygonum,
Rudbeckia trilobata, Silphium perfoliatum, Solidago, Vernonia.
Nomia triangulifera Vachal, 1897. Mise. Ent. 5: 9. d.
Nomia persimilis Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 50, 72. °, d.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 513-515, figs. (redescription).

Biology: Pierce, 1904. Nebr. Univ., Studies, 4: 181 (habits, parasite). —Rau, 1929. Psyche 36:
243. —Cross and Bohart, 1960. Kansas Univ. Sci. Bul. 41: 761-792.

Genus NOMIA Subgenus DIEUNOMIA Cockerell

Eunomia Cresson, 1875. In Wheeler, Rpt. Geog. Geol. Expl. Survey west of 100th
Meridian, v. 5, p. 722. Preocc.
Type-species: Eunomia marginipennis Cresson. Desig. by Cockerell, 1910.
(=Nomia heteropoda kirbii Smith).
Dieunomia Cockerell, 1899. Entomologist 32: 14. N. name.
Monia(!) Howard, 1901. Insect Book, pl. IV, figs. 34, 38.

Revision: Blair, 1935. N. Y. Ent. Soc., Jour. 43: 201-214, pl. XVI.
apacha Cresson. Nebr., Colo., N. Mex., Tex.
Nomia? apacha Cresson, 1868. Amer. Ent. Soc., Trans. 1: 386. 9.
bolliana bolliana Cockerell. N. Mex., Tex.
Nomia bolliana Cockerell, 1910. Ann. and Mag. Nat. Hist. 8 (5): 259. °.

Taxonomy: Cockerell, 1910. U. S. Natl. Mus., Proc. 38: 295. °, d.
bolliana helenii Cockerell. Tex.
Nomia bolliana helenii Cockerell, 1936. Amer. Mus. Novitates 831: 2. 2.
heteropoda heteropoda Say. Man., Minn., Wis., Ill., Ind., Ohio south to Md., Ga., Fla. and Miss.
Pollen: Unknown, but visits flowers of Bidens bipinnata, Eupatorium, Helenium,
Helianthus.
Nomia? heteropoda Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 349. d.
Andrena valida Say, 1837. Boston Jour. Nat. Hist. 1: 393. °.
Nomia heteropoda validescens Blair, 1935. N. Y. Ent. Soce., Jour. 43: 206. d.
Nomia heteropoda subvalida Blair, 1935. N. Y. Ent. Soe., Jour. 43: 206. °.

Taxonomy: Cockerell, 1934. Amer. Mus. Novitates 697: 4. —Mitchell, 1960. N. C. Agr. Expt.
Sta. Tech. Bul. 141: 507-508, figs. (redescription).
heteropoda kirbii Smith. Ga. and Fla. west to South. Calif., north to Colo., Nebr., and IIl.,
south to Tex. and Mexico; apparently incorrectly recordea from Brazil. Pollen:
Unknown, but visits flowers of Helianthus.
Nomia Kirbii Smith, 1865. Ent. Soc. London, Trans. (3) 2: 398. é.
Eunomia marginipennis Cresson, 1875. In Wheeler, Rpt. Geog. Geol. Expl. Survey west
of 100th Meridian, v. 5, p. 722. 2, d.
Nomia heteropoda var. semivalida Cockerell, 1934. Amer. Mus. Novitates 697: 5. d.
Nomia heteropoda var. atripennis Cockerell, 1934. Amer. Mus. Novitates 697: Cpe Neye
Nomia heteropoda semirubra Cockerell, 1934. Amer. Mus. Novitates 697: 6. 2, 6.
Nomia heteropoda var. subvalidior Blair, 1935. N. Y. Ent. Soc., Jour. 43: 208. °.

1946 Hymenoptera in America North of Mexico

Taxonomy: Westwood, 1875. Ent. Soc. London, Trans. p./217. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul. 141: 508 (tax. characters).

mesillae (Cockerell). Colo., N. Mex., Tex.
Dieunomia mesillae Cockerell, 1899. Entomologist 32: 266. d.

Taxonomy: Cockerell, 1930. Amer. Mus. Novitates 433: 10. d, @.

xerophila (Cockerell). Colo., Utah, N. Mex., Ariz.
Dieunomia xerophila Cockerell, 1899. Entomologist 32: 265. ¢, 2.

Genus NOMIA Subgenus UNASSIGNED

tacita Cameron. N. Mex. Probably not Nearctic.
Nomia tacita Cameron, 1902. Amer. Ent. Soc., Trans. 28: 376. 6.

NOMEN NUDUM IN NoMIA LATREILLE
Nomia birkmanii Birkman, 1899. Ent. News 23: 244.

SUBFAMILY HALICTINAE

This very large, diverse and nearly cosmopolitan subfamily of bees exhibits both solitary
behavior and nearly every degree of sociality ranging from subsocial through communal, semiso-
cial to eusocial behavior. Most of the species nest in the ground, but some nest in pre-existing
holes in wood and sometimes in rotten or decaying wood.

Although the species of this subfamily are known to visit the flowers of a great variety of
plants for pollen and nectar, some groups of species are oligolectic (e.g., Hemihalictus and
Sphecodogastra). Since much of the floral information recorded in the literature is in need of
critical evaluation, no attempt has been made to list the floral records for the majority of the
species in this subfamily. Partial summaries of the floral records or comments about them may
be found in a number of studies including Mitchell (1960. N. C. Agr. Expt. Sta. Tech. Bul. 141:
332-506), Moldenke and Neff (1974. Origin and Structure of Ecosystems (IBP) Tech. Rpt. 74-75:
9-135), Ordway (1966. Kansas Univ. Sci. Bul. 46: 509-624), Roberts (1969. Kansas Univ. Sci. Bul.
48: 689-719), and Robertson (1928. Flowers and insects, Carlinville, II]., 221 pp.).

Taxonomy: Hickwort and Fischer, 1963. Ent. Soc. Amer., Ann. 56: 348-354 (hind tibial spurs).
—Eickwort, 1969. Ent. Soc. Amer., Ann. 62: 652-660 (classification and nest architecture).

Biology: Michener, 1958. Ecology 39: 207-217 (distribution of nests in earth banks).
—Michener and Lange, 1958. Science 127: 1046-1047 (primitive social behavior).
—Hamilton, 1964. Jour. Theoret. Biol. 7: 17-52 (genetical evolution). —Michener, 1964. Ins.
Sociaux 11: 317-342 (reproductive efficiency). —Lin, 1964. Ins. Sociaux 11: 187-192
(parasitism as a factor in the evolution of social behavior). —Batra, 1966. Indian Jour. Ent.
28: 375-393. —Knerer and Atwood, 1966. Science 152: 1262-1263 (polymorphism). —Knerer
and Atwood, 1966. Canad. Ent. 98: 13837-1339 (nest architecture and taxonomy). —Knerer
and Atwood, 1967 (1966). Ent. Soc. Ontario, Proc. 97: 103-110 (parasitization). —Knerer
and Plateaux-Quenu, 1966. Acad. Sci. Paris, Compt. Rend. 263: 1622-1625 (cell coverings).
—Knerer and Plateaux-Quenu, 1966. Acad. Sci. Paris, Compt. Rend. 263: 1759-1761
(polymorphism). —Knerer and Plateaux-Quenu, 1966. Acad. Sci. Paris, Compt. Rend. 263:
2014-2017 (polygyny). —Michener, 1966. Animal Behav. 14: 126-129 (worker interaction).
—Knerer and Plateaux-Quenu, 1967. Acad. Sci. Paris, Compt. Rend. 264: 1-3 (life history).
—Knerer and Plateaux-Quenu, 1967. Acad. Sci. Paris, Compt. Rend. 264: 651-653
(production of lining). —Knerer and Plateaux-Quenu, 1967. Acad. Sci. Paris, Compt. Rend.
264: 1096-1099 (male production). —Plateaux-Quenu, 1967. Soc. Ent. France, Ann. (n. s.) 3:
859-866 (social evolution). —Batra, 1968. Kansas Ent. Soc., Jour. 41: 120-133 (behavior in
nests). —Bonelli, 1969. Soc. Ent. Italiana, Bol. 48: 68-78 (social evolution). —Knerer, 1969.
Internat]. Union Study Social Insects VI Congr. Bern, Proc. pp. 101-107 (social structure).
—Knerer, 1969. Science 164: 429-430 (brood care). —Knerer, 1969. Ent. News 80: 141-147
(nest architecture and defense). —Knerer, 1969. Canad. Jour. Zool. 47: 925-930 (synergistic
evolution). —Houston, 1970. Australian Jour. Zool. 18: 345-351 (male soldier caste). —May,
1970. Science 170: 651-652 (brood care). —-Batra, 1970. Science 170: 652 (brood care).
—Plateaux-Quenu, 1972. La Biologie des abeilles primitives. Masson et Cie, Paris, 200 pp.

Superfamily APOIDEA 1947

—Bell and Hawkins, 1974. Jour. Compar. Physiol. 93: 183-193 (intraspecific agonistic
interactions). —Barrows, 1975. Ins. Sociaux 22: 307-332, 12 figs., 2 tables (mating
behavior). —Barrows, 1976. Kans. Ent. Soe., Jour. 49: 105-119, 2 figs., 3 tables (mating
behavior). —Waddington, 1976. Psyche 83: 112-118 (foraging patterns at flowers of
Convolvulus arvensis).

Morphology: Eickwort and Fischer, 1963. Ent. Soc. Amer., Ann. 56: 348-354, 18 figs. (tibial
spurs). —Lello, 1971. Kansas Ent. Soc., Jour. 44: 14-20 (sting apparatus).

TRIBE AUGOCHLORINI

Taxonomy: Eickwort, 1969. Ent. Soe. Amer., Ann. 62: 652-653 (classification and nest
architecture). —Eickwort, 1969. Kansas Univ. Sci. Bul. 48: 325-524, 418 figs., 3 tables
(generic classification).

Genus AUGOCHLOROPSIS Cockerell

Genus AUGOCHLOROPSIS Subgenus AUGOCHLOROPSIS Cockerell

Augochlora subg. Augochloropsis Cockerell, 1897. Canad. Ent. 29: 4.
Type-species: Augochlora (Augochloropsis) subignita Cockerell. Orig. desig.
(=Augochlora ignita Smith).

This subgenus is found only south of the United States.
Genus AUGOCHLOROPSIS Subgenus PARAUGOCHLOROPSIS Schrottky

Augochloropsis subg. Paraugochloropsis Schrottky, 1906. Ztschr. System. Hym. Dipt. 6:
312.
Type-species: Augochloropsis (Paraugochloropsis) lycorias Schrottky. Monotypic.
Augochlora subg. Tetrachlora Schrottky, 1909. Deut. Ent. Ztschr., p. 481.
Type-species: Halictus multiplex Vachal. Monotypic.
Paraugochlora Schrottky, 1910. Deut. Ent. Ztschr., p. 540.
Type-species: Augochlora spinolae Cockerell. Orig. desig.
Rivalisia Strand, 1921. Arch. Naturges. Abt. A, 87: 270.
Type-species: Rivalisia metallica Strand. Monotypic.
Augochlora subg. Glyptobasis Moure, 1941. Arq. Zool. Estado Sao Paulo 2: 48. Preoce.
Type-species: Augochlora (Glyptobasis) chloera Moure. Orig. desig.
Glytobasia Moure, 1941. Arq. do Mus. Paranaense 1: 98. N. name.
Augochloropsis subg. Glyptochlora Moure, 1958. N. Y. Ent. Soe., Jour. 66: 188. This
subgenus has questionably been placed as a synonym of Paraugochloropsis.
Type-species: Megalopta ornata Smith. Orig. desig.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 463-469 (spp. of United
States). —Eickwort, 1969. Kansas Univ. Sci. Bul. 48: 414-416 (generic synonymy and tax.
characters).

anonyma (Cockerell). Coastal Plains, N. C. to Fla. Pollen: Polylege, visiting flowers of
Baccharis, Bidens, Cirsium, Erigeron, Helenium, Hypericum, Ilex, Linaria, Melilotus,
Metopium, Ocimum, Polygonum, Pycnanthemum, Rhus, Rubus, Trifolium.
Augochlora anonyma Cockerell, 1922. U. S. Natl. Mus., Proc. 60: 15. d.
metallica fulgida (Smith). Mich. to New England, south to La. and Fla. Pollen: Polylege,
visiting flowers of Aronia, Aster, Azalea, Barbarea, Chrysanthemum, Cornus,
Eryngium, Fragaria, Geranium, Hypericum, Ilex, Linaria, Melilotus, Rhus, Rubus,
Solidago, Taxaracum.
Augochlora fulgida Smith, 1853. Cat. Hym. Brit. Mus. v. 1, p. 79. °.
Augochlora lucidula Smith, 1853. Cat. Hym. Brit. Mus. v. 1, p. 81. °.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 465 (tax. characters and

status).
metallica metallica (Fabricius). New England and Ont., south to Fla. and Ariz.; Mexico to
Panama and Colombia (Gorgona Island). Pollen: Polylege, visiting flowers of Apocynum,

1948 Hymenoptera in America North of Mexico

Arabis, Aralia, Aronia, Asclepias, Aster, Bidens, Blephilia, Brassica, Cacalia, Cassia,
Caulophyllum, Ceanothus, Celastrus, Cercis, Chrysanthemum, Cicuta, Circaea,
Claytonia, Coreopsis, Cornus, Crataegus, Cornuta grandifloria, Cuphea balsamona,
Diospyros, Dodecatheon, Erigenia, Eryngium, Eupatorium, Fragaria, Geranium,
Gerardia, Gnaphalium, Gonolobus, Haplopappus, Hydrangea, Hydrophyllum,
Hypericum, Ilex, Lepachys, Lippia, Lycopus, Malva, Melilotus, Nepeta, Osmorrhiza,
Parthenium, Petalostemon, Polygonum, Polytaenia, Prunus, Pycnanthemum,
Rhamnus, Rhus, Ribes, Rosa, Rubus, Sagittaria, Salix, Sanicula, Scrophularia,
Smilacina, Solidago, Specularia, Smyphoricarpos, Taenidia, Thaspium, Tilia,
Trifolium, Verbascum, Verbena, Veronica, Zanthoxyllum, Zizia. Predator: Philanthus
gibbosus (Fabr.), P. solivagus Say.

Andrena metallica Fabricius, 1793. Ent. syst. 2: 309. 9.

Augochlora cuprea Smith, 1853. Cat. Hym. Brit. Mus., v. 1,.p. 79. 2.

Augochlora viridula Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 81. ¢.

Augochlora fervida Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 81. d.

Augochlora cleomis Titus, 1901. Canad. Ent. 33: 135. 2, 6.

Halictus (Augochlora) chorisis Vachal, 1903. Mise. Ent. 11: 186. 2.

Halictus (Augochlora) viridissimus Viereck, 1910. In Smith, N. J. State Mus., Ann. Rpt.
1909: 688. N. name; proposed to replace viridula Smith, preoccupied in Halictus.

Augochlora fulvofimbriata Friese, 1916. Stettin. Ent. Ztg. 77: 315. @.

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 88-90 (redescription, as
fulvofimbriata). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 465-466, fig. 110
(tax. characters, synonymy). —Moure, 1960. Studia Ent. 3: 102-103 (notes on type,
synonymy).

sumptuosa (Smith). Maine to Fla., west to S. Dak., Colo., Ariz., and Tex. Pollen: Polylege,

visiting flowers of Asclepias, Berlandiera, Bidens, Clethra, Crataegus, Eriogonum,
Eryngium, Helianthus, Hypericum, Melilotus, Silphium, Stachys, Vaccinium.

Augochlora sumptuosa Smith, 1853. Cat. Hym. Brit. Mus., vol. 1, p. 82. 2.

Augochlora humeralis Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 365. 2, d.

Agapostemon caeruleus Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 7. °.

Halictus pattoni Vachal, 1903. Mise. Ent. 11: 132. N. name. Preocc.

Augochlora sumptuosa bolliana Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 31. 9.

Augochlora lacustris Cockerell, 1922. U. S. Natl. Mus., Proc. 60 (18): 14. °.

Augochlora floridica Cockerell, 1922. U. S. Natl. Mus., Proc. 60 (18): 14. ¢.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 466-467, 469, figs. 110, 111
(redescription, synonymy).

Biology: Smith, 1898. Ent. News 9: 157, 320. —Smith, 1901. N. Y. Ent. Soc., Jour. 9: 52.
—Michener and Lange, 1959. Amer. Mus. Novitates 1924: 36.

Genus AUGOCHLORA Smith
Genus AUGOCHLORA Subgenus AUGOCHLORA Smith

Augochlora Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 73.
Type-species: Halictus purus Say. Desig. by Cockerell, 1923.

Oxystoglossa Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 83.
Type-species: Oxystoglossa decorata Smith. Monotypic.

Odontochlora Schrottky, 1909. La Plata Mus., Rev. 16: 14.
Type-species: Augochlora muelleri Cockerell. Orig. desig.

The species of this subgenus nest in dead wood either in pre-existing burrows of other insects
or in soft rotting wood.

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 59 (generic status). — Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 456-458 (spp. of eastern U. S.). —Eickwort,
1969. Kansas Univ. Sci. Bul. 48: 420-422 (tax. characters).

Biology: Eickwort and Eickwort, 1973. Kansas Ent. Soc., Jour. 46: 17-22, figs. (nests).

Superfamily APOIDEA 1949

azteca (Vachal). South. Tex.; Mexico.
Halictus (Oxystoglossa) azteca Vachal, 1911. Mise. Ent. 19: 45, 110. 9.
nigrocyanea Cockerell. South. Tex.; Mexico to Panama.
Augochlora nigrocyanea Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 144. 9.
pura mosieri Cockerell. South. Fla. Pollen: Polylege, visiting flowers of Bidens, Cattleya,
Flaveria, Hamelia, Melaleuca, Psychotrea, Salix, Sida.
Augochlora mosieri Cockerell, 1922. U. S. Natl. Mus., Proc. 60: 15. 9.
pura pura (Say). Que. and Maine to Minn., south to Tex. and Fla. Parasite: Rhipiphorus
schwarzi LeC., R. stylopoides Newm.? Pollen: Polylege, visits a wide variety of flowers
including Acer, Agastache, Althaea, Amelanchier, Arabis, Aruncus, Asclepias, Aster,
Barbarea, Bidens, Blephilia, Cacalia, Caulophyllum, Cercis, Chaerophyllum, Cicuta,
Coreopsis, Cornus, Cryptotaenia, Diospyros, Dirca, Evonymus, Fragaria, Geranium,
Gerardia, Haplopappus, Heliopsis, Helianthus, Houstonia, Hydrangea, Ilex, Lippia,
Ludwigia, Malus, Malva, Melilotus, Osmorrhiza, Padus, Polemonium, Polygonum,
Prunus, Ranunculus, Rhamnus, Rhus, Rudbeckia, Rubus, Salix, Smilacina, Solidago,
Smyphoricarpos, Taenidia, Teucrium, Thaspium, Tradescantia, Vaccinium, Verbena,
Verbesina, Vernonia, Vitis, Zizia. Predator: Philanthus gibbosus (Fabr.), P. lepidus
Cress., P. solivagus Say.
Halictus purus Say, 1837. Boston Jour. Nat. Hist. 1: 395. 2, 6.
Augochlora festiva Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 80. °.
Augochlora Robertsoni Cockerell, 1897. Canad. Ent. 29: 69. °.
Augochlora Banksiella Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 533. 9.
Halictus astios Vachal, 1911. Mise. Ent. 19: 44. 9.
Halictus astios var.(?) fuscatipes Vachal, 1911. Mise. Ent. 19: 45. 9.
Halictus asaphes Vachal, 1911. Mise. Ent. 19: 49. °.
Augochlora palmarum Cockerell, 1922. U.S. Natl. Mus., Proc. 60 (18): 15. 9, d.

Taxonomy: Michener, 1953. Kansas Univ. Sci. Bul. 35: 1028, figs. 57-59, 64 (larva).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7): 32. —Blackman and Stage, 1924. N. Y.
State Col. Forestry, Syracuse Univ. Tech. Pub. 17: 197. —Rau, 1934. Acad. Sci. St. Louis,
Trans. 28: 223. —Stockhammer, 1961. North Central Branch, Ent. Soc. Amer., Proc. 16:
17-18. —Stockhammer, 1966. Kansas Ent. Soc., Jour. 39: 157-192 (nesting habits, life cycle).
—May and Stockhammer, 1968. Kansas Ent. Soc., Jour. 41: 339-341, 1 fig. (induced nesting
in cellulose powder). —May, 1972. Kansas Ent. Soc., Jour. 45: 439-449, 4 figs. (water uptake
during larval development). —Barrows, 1973. Kansas Ent. Soc., Jour. 46: 496-499 (induced
soil nesting). —May, 1973. Kansas Ent. Soc., Jour. 46: 301-310, 6 tables (brood cell
recognition). —Barrows, 1974. Fla. Entomologist 57: 189-193 (aggregation behavior).
—May, 1974. Kansas Ent. Soc., Jour. 47: 504-516, 5 figs. (nature and origin of brood cell
linings).

Genus AUGOCHLORA Subgenus OXYSTOGLOSSELLA Eickwort

Augochlora subg. Oxystoglossella Eickwort, 1969. Kansas Univ. Sci. Bul. 48: 422.
Type-species: Augochlora (Oxystoglossella) cordiaefloris Cockerell. Orig. desig.

All the members of this subgenus which have been studied have been found to construct their
nests in the ground.
Taxonomy: Eickwort, 1969. Kansas Univ. Sci. Bul. 48: 422-423.
Biology: Eickwort and Eickwort, 1972. Kansas Ent. Soc., Jour. 45: 18-45 (life history, larva,
pupa).
aztecula Cockerell. South. Tex.; Mexico.
Augochlora aztecula Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 534. 9.
Genus AUGOCHLORA Subgenus UNKNOWN

obliqua Provancher. B. C. (Vancouver).
Augochlora obliqua Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 318. °.

1950 Hymenoptera in America North of Mexico

Genus AUGOCHLORELLA Sandhouse

Augochlorella Sandhouse, 1937. Wash. Acad. Sci., Jour. 27: 66.
Type-species: Augochlora gratiosa Smith. Orig. desig.
Oxystoglossidia Moure, 1943. Rev. Ent. 14: 473.
Type-species: Oxystoglossidia uraniella Moure. Orig. desig.

Revision: Ordway, 1966. Kansas Univ. Sci. Bul. 509-624, 92 figs., 3 tables (spp. north of
Mexico). —Eickwort, 1969. Kansas Univ. Sci. Bul. 48: 424-426 (generic classification).

Biology: Ordway, 1961. North Central Branch, Ent. Soc. Amer., Proe. 16: 17. —Ordway, 1964.
Kansas Ent. Soc., Jour. 37: 139-152. (parasite). —Eickwort, 1966. Kansas Ent. Soce., Jour.
39: 410-429. (phoretic mite).
aurata (Smith). N. C., Ga., Fla., Ala., Tex.; Mexico. Pollen: Polylege, visiting flowers of Aster,
Cirsium, Citrus, Crataegus, Lythrum, Opuntia, Polygonum, Rubus, Viburnum.
Augochlora aurata Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 82. 2.
Augochlora austrina Robertson, 1893. Amer. Ent. Soc., Trans. 20: 147. 2.

bracteata Ordway. Northeast. Tex., south into eastern Mexico (Hidalgo).
Augochlorella bracteata Ordway, 1966. Kansas Univ. Sci. Bul. 46: 570. 2, 6.
gratiosa (Smith). N. J. and D. C., south to Fla., east to Tex. Pollen: Polylege, visiting flowers
of Aster, Ampelopsis, Asclepias, Berteroa, Bidens, Callicarpa, Cassia, Chrysobalanus,
Chrysopsis, Cirsium, Citrus, Clethra, Crataegus, Crotonopsis, Cunila, Erigeron,
Eryngium, Galactia, Gerardia, Gossypium, Helianthus, Hypericum, Ilex, Ixora,
Jacquemontia, Lepidium, Malva, Melilotus, Oenothera, Opuntia, Piriqueta, Polygala,
Polygonum, Pterocaulon, Pycnothymus, Rhus, Rubus, Sabal, Senecio, Solidago,
Taraxacum, Teucrium, Vaccinium, Verbena, Warea.
Augochlora gratiosa Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 80. °.
neglectula neglectula (Cockerell). Ariz., N. Mex., Tex., south to Panama. Pollen: Polylege,
visiting flowers of Acacia, Aesculus, Baccharis, Ceanothus, Chilopsis, Chrysanthemum,
Chrysopsis, Dalea, Descurainia, Echinocactus, Erigeron, Eschscholzia, Fendlera,
Gaillardia, Gossypium, Gutierrezia, Helenium, Heterotheca, Lepidium, Manzanita,
Melilotus, Opuntia, Penstemon, Prunus, Pyrus, Rosa, Senecio, Sida, Sisymbrium,
Sphaeralcea, Ungnadia. The subspecies Augochlorella neglectula maritima Ordway
occurs in the sand dunes along the west coast of Mexico.
Augochlora neglectula Cockerell, 1897. N. Mex. Agr. Expt. Sta. Bul. 24: 43. 2, 6.
Augochlora dimissa Cockerell, 1923. U. S. Natl. Mus., Proc. 63: 5. @.
persimilis (Viereck). Pa., west to Minn., south to Ga., Ala., La., and Tex. Parasite:
Myrmilloides grandiceps (Blake), Psewdomethoca frigida (Sm.), Sphecodes pimpinellae
Robt. Pollen: Polylege, visits a great variety of flowers including Achillea, Agastache,
Ailanthus, Alisma, Althaea, Ammannia, Amorpha, Antennaria, Anthemis, Aphanes,
Apocynum, Arabis, Asclepias, Asparagus, Aster, Barbarea, Bidens, Blephilia, Borago,
Brassica, Callirhoe, Camassia, Campanula, Capsella, Cardamine, Cassia, Ceanothus,
Celastrus, Cerastium, Chrysanthemum, Chrysopsis, Cichorium, Cirsium, Citrullus,
Claytonia, Convolvulus, Coreopsis, Cornus, Cotoneaster, Crataegus, Cucurbita, Daucus,
Descurainia, Diospyros, Echinacea, Erigeron, Erysimum, Eupatorium, Euphorbia,
Fragaria, Geranium, Geum, Gnaphalium, Gutierrezia, Hedeoma, Helenium, Helianthus,
Heliopsis, Heterotheca, Heuchera, Houstonia, Hypoxis, Ipomoea, Justicia, Kolkwitzia,
Krigia, Lepidium, Lespedeza, Lippia, Lobelia, Lotus, Ludwigia, Lycopus, Malva,
Medicago, Melilotus, Monarda, Nepeta, Nothoscordum, Oenothera, Oxalis, Paeonia,
Parosela, Parthenium, Passiflora, Pastinaca, Petalostemon, Phacelia, Plantago,
Polemonium, Polygonum, Polytaenia, Potentilla, Prunus, Psoralea, Pycnanthemum,
Ranunculus, Raphanus, Rhus, Rorippa, Rosa, Rubus, Rudbeckia, Sabatia, Sagittaria,
Salix, Salvia, Senecio, Sida, Silphium, Sisyrinchium, Smilacina, Smilax, Solidago,
Specularia, Stellaria, Symphoricarpos, Taenidia, Taraxacum, Thaspium, Tradescantia,
Trifolium, Valerianella, Verbena, Verbesina, Vernonia, Veronica, Viburnum,
Zigadenus, Zizia. Predator: Philanthus bilunatus Cress., P. gibbosus (Fabr.), P.
lepidus Cress., P. politus politus Say, P. solivagus Say.
Augochlora similis Robertson, 1893. Amer. Ent. Soc.. Trans. 20: 146. 2. ¢. Preoce.

Superfamily APOIDEA 1951

Halictus (Oxystoglossa) persimilis Viereck, 1910. In Smith, N. J. State Mus., Ann. Rpt.
1909: 688. N. name.
Halictus xystris Vachal, 1911. Mise. Ent. 19: 50. °.

Biology: Pearson, 1933. Ecol. Monog. 3: 386, 396 (as Oxystoglossa similis). —Ordway, 1965.
Insectes Sociaux 12: 291-308 (caste differentiation). —Eickwort, 1966. Kansas Ent. Soc.,
Jour. 39: 410-429 (phoretic mite). —Ordway, 1966. Kansas Ent. Soc., Jour. 39: 270-313
(bionomics).

pomoniella (Cockerell). Calif., Nev., Utah, Ariz. N. Mex. (Willow Creek); Mexico (Baja

California, south along west coast to Chiapas and Yucatan), Guatemala and Costa Rica.
Ecology: Nests in abandoned cells of Sceliphron. Pollen: Polylege, visits a great variety
of flowers including Acacia, Achillea, Agoseris, Arctostaphylos, Arenaria, Argemone,
Asclepias, Aster, Astraglus, Baccharis, Baileya, Bebbia, Brassica, Brodiaea,
Calandrinia, Calochortus, Carnegiea, Centaurea, Cercidium, Chaenactis, Chrysopsis,
Chrysothamnus, Cirsium, Cissus, Clarkia, Cleome, Convolvulus, Coreopsis,
Corethrogyne, Croton, Cryptantha, Dalea, Datura, Diplacus, Encelia, Enceliopsis,
Eremocarpus, Eriastrum, Eriodictyon, Eriogonum, Eriophyllum, Erodium, Eryngium,
Eschscholzia, Eucnide, Fendlerella, Gazania, Gilia, Gnaphalium, Gossypium,
Grossularia, Gutierrezia, Haplopappus, Helianthus, Heliotropium, Hemizonia,
Heterotheca, Hymenothrix, Isomeris, Kallstroemia, Lantana, Layia, Lessingia,
Lomatium, Lotus, Malacothamnus, Marrubium, Melilotus, Mentha, Mirabilis,
Monardella, Oenothera, Opuntia, Osmoronia, Penstemon, Perideridia, Petalonyx,
Peucephyllum, Phacelia, Platystemon, Ranunculus, Raphanus, Rhus, Salix, Salvia,
Sanicula, Scrophularia, Senecio, Sisymbrium, Sisyrinchium, Solanum, Solidago,
Sphaeralcea, Stanleya, Stephanomeria, Tamarix, Trichostema, Wyethia Zauschneria.

Augochlora pomoniella Cockerell, 1915. Pomona Jour. Ent. Zool. 7: 232. °.

Augochlora utahensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 314. 2.

striata (Provancher). Southern Canada to south. Fla., west to Rocky Mountains. Parasite:
Myrmilloides grandiceps (Blake), Pseudomethoca frigida (Sm.); Sphecodes pimpinellae
Robt. Pollen: Polylege, visits a great variety of flowers including Achillea, Aesculus,
Agastache, Agoseris, Althaea, Amelanchier, Amorpha, Anemone, Anemonella,
Antennaria, Apocynum, Aquilegia, Arabis, Aralia, Argemone, Aruncus, Asclepias,
Aster, Astragalus, Barbarea, Berteroa, Bidens, Brassica, Caltirhoe, Calopogon,
Camassia, Camelina, Campanula, Capsella, Cardamine, Carduus, Cassia,
Caulophyllum, Ceanothus, Celastrus, Centaurea, Cephalanthus Cercis, Chaerophyllum,
Chrysanthemum, Chrysopsis, Cichorium, Cicuta, Circaea, Cirsium, Citrullus,
Claytonia, Clethra, Convolvulus, Coreopsis, Cornus, Crataegus, Cryptantha,
Cryptotaenia, Cubelium, Cucumis, Cucurbita, Cunila, Daucus, Dentaria, Diervilla,
Dodacatheon, Echinacea, Echium, Ellisia, Erigenia, Erigeron, Erysimum, Evonymus,
Eupatorium, Euphorbia, Fragaria, Gaillardia, Geranium, Gerardia, Glechoma,
Gnaphalium, Gossypium, Grindelia, Gutierrezia, Haplopappus, H edeoma, Heiracium,
Helenium, Helianthus, Heliopsis, Heracleum, Heterotheca, Heuchera, Houstonia,
Hybanthus, Hydrangea, Hydrocotyle, Hydrolea, Hydrophyllum, Hypericum, Impatiens,
Inula, Ipomoea, Iris, Isopyrum, Kolkwitzia, Krigia, Lactuca, Lathyrus, Lepidium,
Lespedeza, Lesquerella, Linum, Lippia, Lobelia, Lomatium, Lonicera, Lotus,
Lycopersicum, Lycopus, Lythrum, Malus, Malva, Medicago, Melilotus, Mentha,
Mertensia, Mikania, Monarda, Myosoton, Nigella, Nothoscordum, Oenothera,
Onopordum, Opuntia, Osmorrhiza, Oxalis, Paeonia, Parthenium, Parthenocissus,
Paspalum, Pastinaca, Penstemon, Perideridia, Petalostemon, Phryma, Physalis,
Polemonium, Polygonatum, Polygonum, Polymnia, Polytaenia, Pontederia, Potentilln,
Prenanthes, Prunella, Prunus, Psoralea, Ptelea, Pteridium, Pycnanthemum,
Pyrrhopappus, Pyrus, Ranunculus, Ratibida, Rhamnus, Rhus, Ribes, Rorippa, Rosa,
Rubus, Rudbeckia, Sagittaria, Salix, Salvia, Sanicula, Sapindus, Satureja,
Scrophularia, Scutellaria, Sedum, Senecio, Sericocarpus, Sida, Silphium, Sisymbrium,
Sisyrinchium, Smilacina, Smilax, Solanum, Solidago, Sonchus, Specularia,
Sphaeralcea, Spiraea, Stellaria, Stokesia, Strophostyles, Symphoricarpos, Syringa,
Taenidia, Tanacetum, Taraxacum, Teucrium, Thaspium, Tradescantia, Tragopogon,
Trifolium, Trillium, Triosteum, Vaccinium, Verbascum, Verbena, Vernonia, Viburnum,

1952 Hymenoptera in America North of Mexico

Vicia, Viola, Vitis, Waldsteinia, Xanthoxylum, Zizia. Predator: Philanthus bilunatus
Cress., P. gibbosus (Fabr.), P. lepidus Cress., P. politus politus Say, P. solivagus Say.
Augochlora striata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. Deer

Augochlora matilida Robertson, 1893. Amer. Ent. Soc., Trans. 20: Aen
Augochlora confusa Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 324. ?, d.
Augochlora Coloradensis Titus, 1901. Canad. Ent. 33: 133. 2, d.

Augochlora pseudopurella Strand, 1914. Arch. Naturgesch. 80 (Abt. A, Heft 1): 163.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 32 (as confusus). —Hicks, 1931. Canad.
Ent. 63: 176 (as coloradensis). —Evans and Lin, 1959. Wasmann Jour. Biol. 17: 120, 128,
127, 131 (as awrata and striata). —Michener and Wille, 1961. Kansas Univ. Sci. Bul. 42:
1130. —Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 174. —Ordway, 1964. Kansas
Ent. Soc., Jour. 37: 139-152. (parasite). —Evans, 1964. Psyche 71: 142, 147. —Michener,
1964. Amer. Zool. 4: 233. (nests). —Ordway, 1965. Ins. Sociaux 12: 291-308 (caste
differentiation). —Ordway, 1966. Kansas Ent. Soc., Jour. 39: 270-313 (bionomics).
—Eickwort, 1966. Kansas Ent. Soc., Jour. 39: 410-429 (phoretic mite).

Genus PSEUDAUGOCHLOROPSIS Schrottky

Augochloropsis subg. Pseudaugochloropsis Schrottky, 1906. Ztschr. System. Hym. Dipt. 6:
313.
Type-species: Halictus nigromarginatus Spinola. Desig. by Schrottky, 1909.
Caenaugochlora subg. Pseudaugochlora Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104:
77.
Type-species: Halictus nigromarginatus Spinola. Orig. desig.

Taxonomy: Eickwort, 1969. Kansas Univ. Sci. Bul. 48: 429-432 (generic classification).

Biology: Michener and Kerfoot, 1967. Kansas Ent. Soc., Jour. 40: 214-232 (nests, social
behavior).
graminea (Fabricius). Tex. to Argentina; West Indies. Pollen: Possibly restricted to the pollen

of Solanum (including S. wendlandii) and Cassia.

Megilla graminea Fabricius, 1804. Syst. Piez., p. 334.

Halictus nigromarginatus Spinola, 1841. Soc. Ent. France, Ann. 10: 137. °.

Augochlora camure Holmberg, 1884. Soc. Cient. Argentina, An. 18: 213. 2.

2Augochlora chapadae Cockerell, 1900. Acad. Nat. Sci. Phila., Proc., p. 361, 375.

Taxonomy: Moure, 1960. Studia Ent. 3: 105-106 (synonymy, notes on type). —Eickwort, 1967.
Kansas Ent. Soc., Jour. 40: 232-237 (tax. characters and status).

Biology: Michener and Kerfoot, 1967. Kansas Ent. Soc., Jour. 40: 214-232 (nests, sex ratio,
social behavior).

Genus TEMNOSOMA Smith

Temnosoma Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 238.
Type-species: Temnosoma metallicum Smith. Monotypic.

Micraugochlora Schrottky, 1909. Rev. Mus. La Plata, 16: 188.
Type-species: Micraugochlora sphaerocephala Schrottky. Monotypic.

Temnosoma subg. Temnosomula Ogloblin, 1953. Soc. Ent. Argentina, Bol. 2: 2.
Type-species: Temnosoma (Temnosomula) platensis Ogloblin. Monotypic and orig.

desig.
Taxonomy: Hickwort, 1969. Kansas Univ. Sci. Bul. 48: 446-448 (generic classification).

smaragdinum Smith. Ariz. (Tucson); Mexico, Centr. Amer.
Temnosoma smaragdinum Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 29. d.

TRIBE HALICTINI

Taxonomy: Eickwort, 1969. Ent. Soc. Amer., Ann. 62: 654-656 (classification and nest
architecture).

Superfamily APOIDEA 1953

Genus AGAPOSTEMON Guerin-Meneville

Andrena subg. Agapostemon Guerin-Meneville, 1844. Iconogr. Regne Anim., Ins., v. 3, p.
448.
Type-species: Apis femoralis Guerin-Meneville. Monotypic. (=Apis viridula
Fabricius).

The bees of this genus are known to visit a great variety of flowers for nectar and pollen.
Since the data pertaining to the floral visitations of these bees are conveniently available (see
discussion of the subfamily Halictinae), this information is not repeated here or presented in the
species treated below.

Revision: Roberts, 1972. Kansas Univ. Sci. Bul. 49: 437-590, 228 figs. (included spp.).

Taxonomy: Roberts, 1973. Oreg. State Univ. Agr. Expt. Sta. Tech. Bul. 125: 1-23 (northwest.
Amer. spp.).

Biology: Roberts, 1969. Kansas Univ. Sci. Bul. 48: 689-719, 21 figs. (natural history).
SPECIES GROUP SPLENDENS

angelicus Cockerell. N. Dak., Colo. and Iowa, south to Tex., N: Mex., Ariz. and south. Calif.;
Mexico (Chihuahua, Durango and Sonora). Predator: Philanthus albopilosus Cress.
Agapostemon angelicus Cockerell, 1924. Calif. Acad. Sei., Proce. (4) 12: 537. 2.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 159, fig. 5 (sleep). —Evans, 1975. Ent. Soc.
Amer., Ann. 68: 891 (predator).

texanus Cresson. Transcont. south. Canada and U. S., south to centr. Costa Rica. Predator:

Philanthus crabroniformis Smith.

Agapostemon texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 255. 2.

Agapostemon texanus subtilior Cockerell, 1898. Ent. News 9: 27. 2 ?

Agapostemon borealis Crawford, 1901. Nebr. Acad. Sci., Proc. 7: 160. 2.

Agapostemon californicus Crawford, 1901. Nebr. Acad. Sci., Proc. 7: 164. 2.

Halictus (Agapostemon) brachycerus Vachal, 1903. Mise. Ent. 11: 101. d.

Agapostemon texanus iowensis Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 363. 2.

Agapostemon proscriptus Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 10: 24. °.

Agapostemon joseanus Friese, 1916. Stettin. Ent. Ztg. 77: 310. 2.

Agapostemon sulfuripes Friese, 1916. Stettin. Ent. Ztg. 77: 310. d.

Agapostemon cyanozonus Cockerell, 1924. Calif. Acad. Sci., Proe. (4) 12: 539. d.

Agapostemon proscriptellus Cockerell, 1924. Calif. Acad. Sei., Proc. (4) 12: 538. 2.

Agapostemon texanus vandykei Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 191. °.

Agapostemon californicus psammobius Cockerell, 1937. Pan-Pacifie Ent. 13: 150. d, 2.

Agapostemon angelicus idahoensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 314.
Os

Agapostemon californicus clementinus Cockerell, 1939. Calif. Acad. Sci., Proc. (4) 28: 431.
oo)

splendens (Lepeletier). Ont. and Maine to Sask., south to Fla., Ala., Miss., La., Tex., N. Mex.,

Ariz.; Mexico (Gulf coast to Veracruz).

Halictus splendens Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 283. 2.

Agapostemon aeruginosus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 86. ?.

Biology: Stevens, 1921. Canad. Ent. 53: 68.
SPECIES GROUP SERICEUS
cockerelli Crawford. Wyo. Colo., Tex., N. Mex., Ariz.; Mexico.
Agapostemon Cockerelli Crawford, 1901. Nebr. Acad. Sci., Proc. 7: 161. °.
femoratus Crawford. B. C., Alta., N. Dak., south to N. Mex., Ariz., Nev., Calif., Mexico (Baja

California).
Agapostemon femoratus Crawford, 1901. Nebr. Acad. Sci., Proc. 7: 162. d, 2.
sericeus (Forster). Ont., Maine to Fla., west to N. Dak., south to Tex. Predator: Philanthus
sanbornii Cress.
Apis sericea Forster, 1771. Novae Species Insectorum, Centuria vol. I, p. 91. d.
Halictus radiatus Say, 1837. Boston Jour. Nat. Hist. 1: 394. 2.

1954 Hymenoptera in America North of Mexico

Agapostemon pulchra Smith, 1853. Cat. Hym. Brit. Mus. v. 1, p. 87. °.
Agapostemon sulcatulus Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 25. 6.

Taxonomy: van der Vecht, 1959. Ent. Ber. 19: 69 (tax. status). — Roberts, 1972. Kansas Univ.
Sei. Bul. 49: 554 (tax. status). —Day and Fitton, 1977. Biol. Jour. Linn. Soc. 9: 39. 3 (tax.
status of type, synonymy).

Biology: Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 219.
SPECIES GROUP VIRESCENS

coloradinus Crawford. S. Dak., Nebr., Kans., Okla., Tex., N. Mex., Colo., Wyo., Utah.
Agapostemon coloradensis Crawford, 1901. Nebr. Acad. Sci., Proc. 7: 163. 2. Preoce.
through secondary homonymy prior to 1961.
Halictus Coloradinus Vachal, 1903. Mise. Ent. 11: 90. N. name.
tyleri Cockerell. Ariz., N. Mex., Tex.; Mexico.

Agapostemon tyleri Cockerell, 1917. Ann. and Mag. Nat. Hist. (8) 20: 241. 2, 6.
Agapostemon martini Cockerell, 1927. Pan-Pacific Ent. 3: 153. 9 (¢ misdet.).
virescens (Fabricius). Ont. and Maine to Fla., west to B. C., Wash. and Oreg. Predator:

Philanthus gibbosus (Fabr.), P. solivagus Say.
Andrena virescens Fabricius, 1775. Systema Ent., p. 378. 2.
Andrena nigricornis Fabricius, 1793. Ent. System., v. 2, p. 313. 6.
Halictus dimidiatus Lepeletier, 1841. Hist. Nat. Ins., Hym. v. 2, p. 283. 2.
Halictus tricolor Lepeletier, 1841. Hist. Nat. Ins., Hym. v. 2, p. 389. ¢.
Agapostemon bicolor Robertson, 1893. Amer. Ent. Soc., Trans. 20: 148. 2, 6.

Taxonomy: Moure, 1960. Studia Ent. 3: 103-104 (synonymy, notes on type).
SPECIES GROUP MELLIVENTRIS

melliventris Cresson. Mont., Idaho, S. Dak., south to Tex., N. Mex., Ariz., and Calif.; Mexico.
Ecology: Nests in abandoned burrows and holes including vacated Sceliphron nests.
Predator: Philanthus arizonicus Bohart.
Agapostemon melliventris Cresson, 1874. Amer. Ent. Soc., Trans. 5: 101. 2.
Agapostemon fasciatus Crawford, 1901. Nebr. Acad. Sci., Proc. 7: 163. 2, 3.
Halictus (Agapostemon) plurifasciatus Vachal, 1903. Mise. Ent. 11: 93, 101. 2, 6.
Agapostemon digueti Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 539. 2, 6.
mexicanus Roberts. Mexico (Baja California and Sonora). May occur in U. S. (vicinity of San
Diego, Calif. or Yuma, Ariz.).
Agapostemon mexicanus Roberts, 1972. Kansas Univ. Sci. Bul. 49: 503. 6, ?.
peninsularis Roberts. Calif. (La Jolla and San Diego); Mexico (Baja California).
Agapostemon peninsularis Roberts, 1972. Kansas Univ. Sci. Bul. 49: 515. 6, 2.

SPECIES GRouP EREBUS

leunculus Vachal. Mexico to Colombia and Ecuador. May occur in U. S. since the species has
been collected 10 miles southwest of Pharr, Texas in Mexico.
Agapostemon leunculus Vachal, 1903. Mise. Ent. 11: 92. 2.
Agapostemon vulpicolor Crawford, 1906. Amer. Ent. Soc., Trans. 32: 162. 9.
Agapostemon nasutus var. ater Friese, 1916. Stettin. Ent. Ztg. 77: 310. °.

SPECIES GROUP NASUTUS

nasutus Smith. Tex. (Brownsville), south to South America (Colombia, Ecuador, Peru and
Venezuela) including Trinidad. Parasite: Zodion americanum Wied.
Agapostemon nasutus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 87. 3.
Agapostemon peruvianus Cameron, 1903. Amer. Ent. Soc., Trans. 29: 237. 6.
Agapostemon nasutus gualanicus Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 9: 556. d.
Agapostemon purpureopictus Cockerell, 1924. Calif. Acad. Sci., Proce. (4) 12: 538. °.
Agapostemon melanurus Cockerell, 1949. U. S. Natl. Mus., Proc. 98: 438. °.

Biology: Eickwort and Eickwort, 1969. Kansas Ent. Soc., Jour. 42: 421-452 (life history).

Superfamily APOIDEA 1955

Genus HALICTUS Latreille

Revision: Sandhouse, 1941. Ent. Amer. (n. s.) 21: 23-37. —Mitchell, 1960. N. C. Agr. Expt.
Sta. Tech. Bul. 141: 332-338, figs. 82-83 (eastern U. S. spp.).

Biology: Wille and Michener, 1971. Biol. Trop., Rev. 18: 17-31 (Neotropical spp.).

Taxonomy: Roberts, 1973. Oreg. State Univ. Agr. Expt. Sta. Tech. Bul. 126: 1-23 (northwest.
Amer. spp.).

Genus HALICTUS Subgenus HALICTUS Latreille

Halictus Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, p. 182.
Type-species: Apis quadricincta Fabricius. Desig. by Richards, 1935.
Odontalictus Robertson, 1918. Ent. News 29: 91.
Type-species: Halictus ligatuws Say. Monotypic and orig. desig.
Monilapis Cockerell, 1931. Ann. and Mag. Nat. Hist. (10) 7: 529.
Type-species: Hylaeus tomentosus Eversmann. Monotypic and orig. desig.
farinosus Smith. Mont., Nebr., and N. Mex., west to B. C. and Calif. Parasite: Sphecodes
arvensiformis Ckll. Predator: Philanthus crabroniformis Sm.
Halictus farinosus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 69. 2.
Halictus montanus Crawford, 1902. Canad. Ent. 34: 234. 2, 6.
Paranomia venablesii Ashmead, 1903. Canad. Ent. 35: 243. °.
Halictus denticulus Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 469. 2, d.
Halictus procerus Vachal, 1904. Soc. Sei. Hist. Arch. Correze, Bul. 26: 469. d.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 350. °.
ligatus Say. North America south of about 50 degrees latitude, southward to the West Indies

and Colombia. Parasite: Nomada sp., Thecophora modesta (Will.). Predator: Philanthus
albopilosus Cress., P. bilunatus Cress., P. gibbosus (Fabr.), P. lepidus Cress., P.
pacificus arizonae Dunn., P. solivagus Say, P. ventilabris Fabr.

Halictus ligatus Say, 1837. Boston Jour. Nat. Hist. 1: 396. 2, d.

Halictus poeyi Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 271. d.

Halictus capitosus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 67. °.

Halictus armaticeps Cresson, 1872. Amer. Ent. Soc., Trans. 4: 250. °.

Halictus teranus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 251. 2, 6.

Halictus ornatipes Cresson, 1872. Amer. Ent. Soc., Trans. 4: 252. d.

Halictus townsendi Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 293. °.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 31. —Hicks, 1926. Colo. Univ., Studies 15:
217. —Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 160.
parallelus Say. Ont. and N. J. to Fla., west to Mont., N. Mex., and Tex.
Halictus parallelus Say, 1837. Boston Jour. Nat. Hist. 1: 397. 2.
Halictus occidentalis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 250. °, ¢.

Biology: Packard, 1868. Amer. Nat. 1: 364. —Hungerford and Williams, 1912. Ent. News 23:
241.
rubicundus (Christ). Holarctic, in North America ranging from Alaska, B. C., N. W. T., and N.
S., south to Calif., Ariz., Tex., and Fla. Parasite: Nomada imbricata Sm., Sphecodes
dichrous Sm., Zodion cinerium (Fabr.). Predator: Philanthus albopilosus Cress., P.
crabroniformis Sm., P. solivagus Say.
Apis rubicunda Christ, 1791. Naturges. Klassif. Nomencl. Ins. Bienen, Wespen,
Ameisengeschlect...Hym., p. 190. °.
Halictus lerouxi Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 272. 2.
Halictus lerouxi var. ruborum Cockerell, 1898. Canad. Ent. 30: 52. °.
Halictus (Halictus) lerouxii Lovell, 1908. Psyche 15: 34. Emend.
Halictus lupinelli Cockerell, 1936. Pan-Pacific Ent. 12: 158. °.

Taxonomy: Cockerell, 1937. Canad. Ent. 69: 88.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 217. —Hicks, 1934. Colo. Univ., Studies 21: 265.
—Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 162.

1956 Hymenoptera in America North of Mexico
Genus HALICTUS Subgenus SELADONIA Robertson

Seladonia Robertson, 1918. Ent. News 29: 91.
Type-species: Apis seladonius Fabricius. Orig. desig.
confusus arapahonum Cockerell. N. Dak. to N. Mex., west to B. C., Oreg., Calif. and Utah.
Halictus arapahonum Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 316. 9.
Halictus provancheri nearcticus Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 661.

Preoce.
Halictus (Chloralictus) olivarius Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (2532): 10. °.

Taxonomy: Michener, 1951. Jn Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr.
Monog. 2: 1105 (synonymy).
confusus confusus Smith. Alaska, to N. S. south to Fla. west to N. Dak. and Tex. Predator:

Philanthus albopilosus Cress., P. bilunatus Cress., P. crabroniformis Sm., P. gibbosus
(Fabr.), P. pacificus arizonae Dunn., P. politus politus Say, P. pulcher Dalla Torre, P.
solivagus Say.

Halictus confusus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 70. 2, ¢.

Halictus constrictus Provancher, 1882. Nat. Canad. 13: 202. ?. Preoce.

Halictus provancheri Dalla Torre, 1896. Cat. Hym., v. 10, p. 77. N. name.

Halictus nearcticus Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 470. 2, 6.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 350. 2. —Cockerell, 1911. Ann. and
Mag. Nat. Hist. (8) 8: 660. —Michener, 1951. Jn Muesebeck, Krombein and Townes, U. S.
Dept. Agr., Agr. Monog. 2: 1105 (synonymy).

Biology: Hicks, 1936. Canad. Ent. 68: 47. —Knerer and Atwood, 1962. Ent. Soc. Ontario,
Proc. 92: 161.

harmonius Sandhouse. South. Calif.
Halictus (Halictus) harmonius Sandhouse, 1941. Ent. Amer. (n. s.) 21: 36. 2.
tripartitus Cockerell. Idaho, Colo., and Tex. to Wash., Calif., Mexico (Baja California).
Predator: Philanthus crabroniformis Sm., P. gibbosus (Fabr.), P. pacificus arizonae
Dunn., P. pulcher Dalla Torre.
Halictus tripartitws Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 63. 2.
Halictus meliloti Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 67. °.
Halictus catalinensis Cockerell, 1903. South. Calif. Acad. Sci., Bul. 2: 84. 2.

Taxonomy: Michener, 1953. Kansas Univ. Sci. Bul. 35: 1027, figs. 60, 61 (larva).
virgatellus Cockerell. N. W. T., B. C., Alta., Oreg., Colo., N. Mex.
Halictus virgatellus Cockerell, 1901. Psyche 9: 284. 2.
Halictus sansoni Crawford, 1911. U.S. Natl. Mus., Proc. 41: 267. 2.
Halictus fraserae Cockerell, 1916. Entomologist 49: 100. 9°.
Halictus typographicus Cockerell, 1918. Entomologist 51: 261. d.
Halictus (Seladonia) ororyctes Cockerell, 1933. Ent. Soc. Amer., Ann. 26: 40. 9.

Taxonomy: Cockerell, 1919. Ent. News 30: 288. 6.
NoMINA NuDA IN HALICTUS LATREILLE

Halictus californica Bridwell, 1899. Kans. Acad. Sci., Trans. 16: 210.

Halictus emarginata Smith, 1910. N. J. State Mus., Ann. Rpt. 1909, p. 688.

Halictus fendleri Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 252.
Halictus medionitens Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 252.
Halictus missouriensis Bridwell, 1899. Kans. Acad. Sci., Trans. 16: 210.

Halictus pulzenus Clements and Long, 1923. Carnegie Inst. Wash., Pub. 336: 252.
Halictus rhoptoides Bray, 1917. Pomona Jour. Ent. Zool. 9: 99.

Genus LASIOGLOSSUM Curtis

Lasioglossum Curtis, 1833. Brit. Ent. v. 10, p. 448.
Type-species: Melitta xanthopus Kirby. Monotypic and orig. desig. (=Lasioglossum
tricingulum Curtis).
Curtisapis Robertson, 1918. Ent. News 29: 91.

Superfamily APOIDEA 1957

Type-species: Halictus coriaceus Smith. Orig. desig.
Pachyhalictus Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 589.
Type-species: Halictus merescens Cockerell. Orig. desig.

Revision: Sandhouse, 1933. Ent. Soc. Wash., Proc. 35: 80-83 (eastern U. S. spp.). —Mitchell,
1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 338-347, figs. 81, 84-87 (eastern U. S. spp.).

athabascense (Sandhouse). N. S. to B. C., south to Wis., Mich., Ohio and in the eastern U. S. to
N.C

Halictus athabascense Sandhouse, 1933. Ent. Soe. Wash., Proc. 35: 78. 6, °.

bardum (Cresson). Tex. to Calif.
Halictus bardus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 251. 2.

Taxonomy: Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 163.
citerior (Vachal). No locality recorded.
Halictus citerior Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 473. °.
coriaceum (Smith). N. S. to Ga., west to Alta. and N. Mex. ranging southward in Mississippi
Valley to south. Ill. Predator: Philanthus gibbosus (Fabr.), P. solivagus Say.
Halictus coriaceus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 70. °.
Halictus subquadratus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 72. d.
Halictus 6-cinctus Provancher, 1882. Nat. Canad. 13: 200. ¢.
Halictus sexcinctus Dalla Torre, 1896. Cat. Hym., v. 10, p. 59. Emend.
cyaneiceps (Cockerell). N. Mex., Ariz.
Halictus cyaneiceps Cockerell, 1916. Canad. Ent. 48: 254. 2, 6.
forbesii (Robertson). N. B. west to Wash., south to Tex. and Ga. Predator: Philanthus
sanborni Cress.
Halictus Forbesii Robertson, 1890. Amer. Ent. Soc., Trans. 17: 315. @, 6.
Halictus forbesi Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 163. Emend.

Taxonomy: Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 294.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 163.
fuscipenne (Smith). N.S. to Ontario, south to Tex. and Fla.
Halictus fuscipennis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 67. 2, d.
Halictus capitulatus Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 472. °.
heterorhinum (Cockerell). Colo., Ariz.
Halictus heterorhinus Cockerell, 1930. Amer. Mus. Novitates 397: 6. 2.
leucozonium (Schrank). Holarctic, widespread in Europe and Canada; Maine, N. Y. Predator:
Philanthus albopilosus Cress., P. bilunatus Cress., P. solivagus Say.
Apis leucozonia Schrank, 1781. Enum. Ins. Austr., p. 406.
Halictus similis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 69. °.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 352.

Biology: Atwood, 1933. Canad. Jour. Res. 9: 449. —Knerer and Atwood, 1962. Ent. Soc.
Ontario, Proc. 92: 163.
manitouellum (Cockerell). Colo.
Halictus manitouellus Cockerell, 1908. Ent. Soc. Wash., Proc. 9: 119. 2.
mellipes (Crawford). Calif.; Mexico (Baja California).
Halictus mellipes Crawford, 1907. Invertebrata Pacifica 19: 190. °.
olympiae (Cockerell). B. C., Wash., Oreg., Calif.,? Colo.
Halictus olympiae Cockerell, 1898. Canad. Ent. 30: 51. 2.
Halictus olympiae var. subangustus Cockerell, 1898. Canad. Ent. 30: 51. 2.
Halictus olympiae var. subangustatus Crawford, 1906. Canad. Ent. 38: 301. Error for
subangustus.

Taxonomy: Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 476. d ?
pacificum (Cockerell). B. C., Wash., Calif.
Halictus pacificus Cockerell, 1898. Acad. Nat. Sci. Phila., Proc. 50: 50. 2, d.

pavonotum (Cockerell). Calif.
Halictus pavonotus Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 188. 2, d.

1958 Hymenoptera in America North of Mexico

ripariellum (Cockerell). Colo.
Halictus ripariellus Cockerell, 1918. Entomologist 51: 261. 9.
sisymbrii (Cockerell). Wyo. to N. Mex., west to B. C. and Calif.; Mexico (Baja California).
Predator: Philanthus ventilabris Fabr.
Halictus sisymbrii Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 63. 9.

Taxonomy: Cockerell, 1897. Amer. Ent. Soc., Trans. 25: 165. 2, d.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 159. —Aleock and Gamboa, 1975. Ariz.
Acad. Sci. 10: 163 (predator).
titusi (Crawford). Oreg., Calif.
Halictus Titusi Crawford, 1902. Canad. Ent. 34: 235. 9°.
trizonatum (Cresson). Alta. to N. Mex., west to B. C. and Calif.
Halictus trizonatus Cresson, 1874. Amer. Ent. Soc., Trans. 5: 101. 2.
Halictus egregius Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 476. 2.
Halictus colatus Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 476. ¢.
zonulum (Smith). Holarctic; N. S. to Minn., south to N. Y., Mich., Wis. Predator: Phalanthus
gibbosus (Fabr.).
Halictus zonulus Smith, 1848. Zoologist 6: 2171. 2, 6.
Halictus craterus Lovell, 1908. Psyche 15: 35. @, ¢.

Biology: Brittain, 1933. Canad. Dept. Agr., Bul. (n. s.) 162: 94. —Knerer and Atwood, 1962.
Ent. Soc. Ontario, Proc. 92: 163.

Genus EVYLAEUS Robertson

Evylaeus Robertson, 1902. Canad. Ent. 34: 244.
Type-species: Halictus arcuatus Robertson. Orig. desig. (=Halictus cinctipes
Provancher).

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 348-363, figs. 89-91 (eastern
U.S. spp.).

Biology: Knerer and Plateaux, 1967. Acad. Sci. Paris, Compt. Rend. 265: 455-458 (nest
architecture).
aberrans (Crawford). N. Dak., Nebr., Colo., N. Mex., Alta., Oreg., Calif., Utah. Parasite:
Sphecodes sp. Pollen: Collects pollen principally from Oenothera, but also visits these
and a wide variety of other plants for nectar or occasionally pollen. Predator:
Philanthus gibbosus (Fabr.).
Halictus aberrans Crawford, 1903. Canad. Ent. 35: 336. °.
Halictus galpinsiae Cockerell, 1903. Canad. Ent. 35: 342. 9.
Halictus gelidus Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 481. d.

Biology: Bohart and Youssef, 1977 (1976). Wasmann Jour. Biol. 34: 185-234, 18 figs., 4 tables
(nest architecture, life history, behavior, parasite, as galpinsiae).
absurdiceps (Timberlake). Calif. (Central Valley).
Halictus (Evylaeus) absurdiceps Timberlake, 1962. Ent. Soc. Wash., Proc. 62: 105. ¢.
allonotus (Cockerell). Calif.
Halictus allonotus Cockerell, 1936. Pan-Pacific Ent. 12: 156. 2.
amicus (Cockerell). N. Mex., Ariz., Calif.
Halictus amicus Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 164. °.
angustior (Cockerell). N. Mex., Ariz.
Halictus angustior Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 165, 167. 9, 6.
argemonis (Cockerell). Calif.; Mexico.
Halictus arcuatus var. argemonis Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 146. °.
Halictus latifrons Crawford, 1907. Invertebrata Pacifica 1: 192. °.

Taxonomy: Michener, 1951. Jn Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr.
Monog. 2: 1107 (synonymy).
arizonensis (Crawford). Ariz., Calif.; Mexico (Baja California).
Halictus arizonensis Crawford, 1907. N. Y. Ent. Soc., Jour. 15: 189. 2.

Superfamily APOIDEA 1959

aspilurus (Cockerell). Calif.
Halictus aspilurus Cockerell, 1925. Calif. Acad. Sci., Proce. (4) 14: 190. 9.

avalonensis (Cockerell). Calif.
Halictus avalonensis Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 150. 9.

Taxonomy: Cockerell, 1939. Calif. Acad. Sci., Proc. (4) 23: 435. d.
birkmanni (Crawford). Tex.
Halictus Birkmanni Crawford, 1906. Canad. Ent. 38: 5. 9.
bradleyi Mitchell. N. J. (Pitman). This may be the male of Evylaeus pectinatus (Robertson).
Evylaeus bradleyi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 356, figs. 88, 91. d.
cinctipes (Provancher). N. S. to Fla., west to N. Dak. and Colo.
Halictus cinctipes Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 316. d.
Halictus arcuatus Robertson, 1893. Amer. Ent. Soc., Trans. 20: 145. °.
Halictus crassus Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 477. °.
Halictus adelipus Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 479. ¢.
Halictus arcuatus var. parisus Lovell, 1908. Psyche 15: 36. °.
Halictus arcuatus gulosus Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 661. 2.
Halictus gulosus punctiferus Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 286. 2.

Taxonomy: Knerer and Atwood, 1964. Canad. Ent. 96: 960-962, fig. 1.

Biology: Brittain, 1933. Canad. Dept. Agr., Bul. (n. s.) 162: 94. —Atwood, 1933. Canad. Jour.
Res. 9: 448. —Knerer and Atwood, 1964. Canad. Ent. 96: 960-962, fig. 1. —Knerer and
Plateaux-Quenu, 1966. Acad. Sci. Paris, Compt. Rend. 263: 1622-1625 (nests).

comagensis Knerer and Atwood. Ont., Alaska.
Evylaeus comagensis Knerer and Atwood, 1964. Canad. Ent. 96: 959, fig. 1. 2, 6.
cooleyi (Crawford). Alta., B. C., Mont., Colo., N. Mex., Tex., Oreg., Calif. Predator: Philanthus
crabroniformis Sm.
Halictus cooleyi Crawferd, 1906. In Viereck, Canad. Ent. 38: 301. 2, d.
cordleyi (Crawford). B. C., Oreg., Calif.
Halictus Cordleyi Crawford, 1906. In Viereck, Canad. Ent. 38: 302. 2, d.
dasiphorae (Cockerell). N. Mex., Calif.
Halictus dasiphorae Cockerell, 1901. Psyche 9: 285. °.
diatretus (Vachal). B. C., Wash., Nev., Colo.,? Oreg.
Halictus diatretus Vachal, 1904. Soe. Sei. Hist. Arch. Correze, Bul. 26: 481. 2, d.
divergenoides Mitchell. Mich., Ind. Predator: Philanthus lepidus Cress.
Evylaeus divergenoides Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 351, fig. 91. 6.
divergens (Lovell). N. S. to Minn., south to Ga.
Halictus divergens Lovell, 1905. Canad. Ent. 37: 299. °.

Taxonomy: Knerer and Atwood, 1964. Canad. Ent. 96: 958, fig. 1. d.
fartus (Vachal). Wash.
Halictus fartus Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 483. °.
fedorensis (Crawford). Tex.
Halictus Fedorensis Crawford, 1906. Canad. Ent. 38: 4. °.
foxii (Robertson). N. S. to Man. and Minn., south to Ga.
Halictus gracilis Robertson, 1890. Amer. Ent. Soc., Trans. 17: 316. 2°, d. Preoce.
Halictus foxii Robertson, 1895. Amer. Ent. Soc., Trans. 22: 117. N. name.
Halictus gracillimus Dalla Torre, 1896. Cat. Hym., v. 10, p. 68. N. name.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 164.

giffardi (Michener). Calif.
Halictus giffardi Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 315. °.
glabriventris (Crawford). Oreg., Calif., Colo.
Halictus Vachali Crawford, 1906. In Viereck, Canad. Ent. 38: 300. 2. Preoce.
Halictus glabriventris Crawford, 1907. Canad. Ent. 39: 21. N. name.

Biology: Hicks, 1936. Canad. Ent. 68: 47.

1960 Hymenoptera in America North of Mexico

granosus (Vachal). Colo.

Halictus granosus Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 477. °, d.
gularis (Vachal). Calif.

Halictus gularis Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 481. 2.
hammondi (Cockerell). Calif.

Halictus hammondi Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 149. °.
humboldtensis (Michener). Calif.

Halictus (Evylaeus) humboldtensis Michener, 1936. Pan-Pacific Ent. 12: 166. °.
inconditus (Cockerell). Alaska, Wash., Colo.

Halictus inconditus Cockerell, 1916. Entomologist 49: 101. °.
kincaidii (Cockerell). Wash., Oreg., Calif.

Halictus kincaidii Cockerell, 1898. Canad. Ent. 30: 51. °.

Taxonomy: Cockerell, 1903. Entomologist 36: 208. —Michener, 1937. Ann. and Mag. Nat.
Hist. (10) 19: 318. 6, 2. —Michener, 1953. Kansas Univ. Sci. Bul. 35: 1025, figs. 48-538
(larva).

lusorius (Cresson). Tex., N. Mex. |
Halictus lusorius Cresson, 1872. Amer. Ent. Soc., Trans. 4: 252. 9.

Taxonomy: Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 166. —Linsley and MacSwain, 1962.
Pan-Pacifie Ent. 38: 45.
macoupinensis (Robertson). Minn. to N. B., south to Ga. and Ala.
Halictus 4-maculatus Robertson, 1890. Amer. Ent. Soc., Trans. 17: 316. 2, d. Preoce.
Halictus quadrimaculatus Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 65. Emend.
Halictus macoupinensis Robertson, 1895. Amer. Ent. Soc., Trans. 22: 117. N. name.
mendocinensis (Michener). Calif.; Mexico (Baja California).
Halictus (Evylaeus) mendocinensis Michener, 1936. Pan-Pacific Ent. 12: 167. °.
miguelensis (Cockerell). Calif.; Mexico (Baja California).
Halictus miguelensis Cockerell, 1937. Pan-Pacifie Ent. 18: 156. ¢.
Halictus cooleyi obscurior Cockerell, 1939. Calif. Acad. Sci., Proce. (4) 28: 435. 2, d. Preoce.
nelumbonis (Robertson). Maine to Minn., south to Fla. and Tex.
Halictus nelumbonis Robertson, 1890. Amer. Ent. Soc., Trans. 17: 316. 2, d.
nigrescens (Crawford). Calif.; Mexico (Baja California).
Halictus nigrescens Crawford, 1907. Invertebrata Pacifica 1: 191. °.

Taxonomy: Cockerell, 1936. Pan-Pacific Ent. 12: 159. °.

nigricollis (Vachal). Colo.
Halictus nigricollis Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 480. 3.
nigridens (Vachal). Calif.
Halictus nigridens Vachal, 1904. Soe. Sci. Hist. Arch. Correze, Bul. 26: 480. d.
niger (Viereck). Labrador, N.S., N. Y., Alta., B. C., south in mountains to N. Mex.; Eurasia.
Predator: Philanthus crabroniformis Sm., P. pacificus arizonae Dunn., P. pulcher Dalla
Torre.
Halictus niger Viereck, 1903. Amer. Ent. Soc., Trans. 29: 56. 2, d.
Halictus fratellus Perez, 1903. Soc. Linn. de Bordeaux, Actes 58: cexiv. 2.
Halictus frey-gessneri Alfken, 1905. Nat. Ver. Bremen, Abh. 18: 73. 2, d.

Taxonomy: Eidman, 1935. Arb. ueber Morph. u. Taxonom. Ent. 2: 104.

occultus (Vachal). Wash.
Halictus occultus Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 478. °.
oenotherae (Stevens). N. S. south to Ga., west to N. Dak. and Kans. Pollen: Oligolege of
Oenothera. Evylaeus pineolensis Mitchell is possibly the male of this species.
Halictus (Evylaeus) oenotherae Stevens, 1920. Ent. News 31: 37. @.
Halictus ralenci Crawford, 1932. Ent. Soc. Wash., Proc. 34: 70. 2.

Taxonomy: Linsley and MacSwain, 1962. Pan-Pacific Ent. 38: 45. —Knerer and Atwood, 1964.
Canad. Ent. 96: 958-959 (geogr. and floral records).

Superfamily APOIDEA 1961

Biology: Knerer and MacKay, 1969. Canad. Jour. Zool. 47: 289-294 (nest architecture, life
history).
orthocarpi (Cockerell). Calif.
Halictus orthocarpi Cockerell, 1936. Pan-Pacifie Ent. 12: 159. 9
Taxonomy: Michener, 1936. Pan-Pacific Ent. 12: 171. 6.

ovaliceps (Cockerell). Colo., N. Mex., B. C., Calif.
Halictus ovaliceps Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 45. 2. Republished
by Cockerell, 1898. N. Mex. Univ., Bul. 1: 45. female.

Taxonomy: Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 317. d.

Biology: Hicks, 1936. Canad. Ent. 68: 47.
pecosensis (Crawford). N. Mex.
Halictus Pecosensis Crawford, 1906. Canad. Ent. 38: 6. 2.
pectinatus (Robertson). Conn., N. J., Md., Ill., Mo.
Halictus pectinatus Robertson, 1890. Amer. Ent. Soc., Trans. 17: 315. °.
pectoralis (Smith). N.S. to Fla., west to Wis., Nebr., Tex.
Halictus pectoralis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 68. °.

Taxonomy: Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 64. —Robertson, 1898. Acad. Sci.
St. Louis, Trans. 8: 44. 6. —Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 350. 9.

pectoraloides (Cockerell). Man., B. C., Colo., N. Mex., Ariz., Calif.; north. Mexico.
Halictus pectoraloides Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 64. °.
Halictus pectoraloides var. beatulus Cockerell, 1918. Entomologist 51: 262. 2.
Halictus (Evylaeus) grindeliae Cockerell, 1934. Ent. News 45: 29. °.

Taxonomy: Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 166. 2, d.
peralpinus (Cockerell). Colo.
Halictus peralpinus Cockerell, 1919. Ent. News 30: 289. °.
peraltus (Cockerell). Wash., Wyo., Colo., N. Mex., Calif. Predator: Philanthus crabroniformis

Sm.
Halictus peraltus Cockerell, 1901. Psyche 9: 164. d.

pineolensis Mitchell. N. C. (Pineola). Possibly male of Evylaeus oenotherae (Stevens).
Evylaeus pineolensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 358. ¢.
pullilabris (Vachal). Calif.
Halictus pullilabris Vachal, 1904. Soc. Sci. Hist. Arch. Correze, Bul. 26: 479. 3.
Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ., Pubs. Ent. 70: 37-40 (flower
relationships).
pulveris (Cockerell). Colo.
Halictus pulveris Cockerell, 1930. Amer. Mus. Novitates 397: 7. 2, d.
quebecensis (Crawford). Minn. to Newfoundland, south to Ga.
Halictus quebecensis Crawford, 1907. N. Y. Ent. Soc., Jour. 15: 189. °.
regis (Cockerell). Colo.
Halictus regis Cockerell, 1916. Entomologist 49: 102. 2.
robertsoni (Crawford). Tex.
Halictus Robertsoni Crawford, 1906. Canad. Ent. 38: 4. °.
robustus (Crawford). Calif.
Halictus robustus Crawford, 1907. Invertebrata Pacifica 1: 191. °.
ruficornis (Crawford). Nev., Calif., Ariz.
Halictus ruficornis Crawford, 1907. Invertebrata Pacifica 1: 192. d.
rufitarsis (Zetterstedt). Holarctic; Alaska, N. S., N. B., Ont., Mich.
Halictus rufitarsis Zetterstedt, 1838. Ins. Lappon., v. 1, p. 462. 2, ¢.
Hylaeus minutissimus Eversmann, 1852. Soc. Nat. Moscou, Bul. 25: 42. 6.
Halictus atricornis Smith, 1870. Ent. Ann., p. 26. 2, d.

Taxonomy: Eidmann, 1935. Arb. ueber Morph. u. Taxonom. Ent. 2: 104.

1962 Hymenoptera in America North of Mexico

sanfrancisconis (Strand). Ariz.
Halictus sanfrancisconis Strand, 1917. Arch. f. Naturgesch. 83 (Abt. A, Heft 11): 57. ¢.

sequoiae (Michener). Calif.
Halictus (Evylaeus) sequoiae Michener, 1936. Pan-Pacifie Ent. 12: 165. 2.
sopinci (Crawford). N. J., N. C., Ga.
Halictus sopinci Crawford, 1932. Ent. Soe. Wash., Proce. 34: 69. 9.
subobscurus (Cockerell). N. Mex.
Halictus subobscurus Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 65. @.
Halictus Cockerelli Crawford, 1902. Canad. Ent. 34: 236. 2.
supranitens (Cockerell). Colo.
Halictus supranitens Cockerell, 1919. Ent. News 30: 289. ?.
swenki (Crawford). N. Dak., Nebr.
Halictus swenki Crawford, 1906. Ent. News 17: 275. 9.
synthyridis (Cockerell). Alta., Colo. Predator: Philanthus crabroniformis Sm., P. pacificus
arizonae Dunn.
Halictus synthyridis Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 427. °, 6.

tracyi (Cockerell). Calif.
Halictus tracyi Cockerell, 1936. Pan-Pacifie Ent. 12: 161. ?.

Taxonomy: Michener, 1936. Pan-Pacifie Ent. 12: 170. 2, d.
truncatus (Robertson). Que. and Maine to Ga., west to Kans., Nebr., and Wash.
Halictus truncatus Robertson, 1901. Canad. Ent. 38: 230. 2, d.
Halictus fulgidus Crawford, 1902. Canad. Ent. 34: 235. 9.

Taxonomy: Knerer and Atwood, 1964. Canad. Ent. 96: 962, fig. 1.

vaporellus (Cockerell). Colo.
Halictus vaporellus Cockerell, 1910. Canad. Ent. 42: 366. °.

Genus HEMIHALICTUS Cockerell

Hemihalictus Cockerell, 1897. Canad. Ent. 29: 288.
Type-species: Panurgus lustrans Cockerell. Monotypic.
lustrans (Cockerell). Va. to Fla., Ind. and Mich. to Miss. and Tex. Pollen: Oligolege of
Pyrrhopappus carolinianus, but occassionally visits other flowers including Cichorium
intybus and Cucurbita presumably for nectar; males also obtain nearly their entire
nectar supply from the flowers of Pyrrhopappus carolinianus.
Panurgus lustrans Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 147. @.

Taxonomy: Michener, 1947. N. Y. Ent. Soc., Jour. 55: 50. d. —Mitchell, 1960. N. C. Agr.
Expt. Sta. Tech. Bul., 141: 363-364, fig. 81.

Biology: Michener, 1947. N. Y. Ent. Soc., Jour. 55: 50. —Daly, 1961. Kans. Ent. Soc., Jour. 34:
134-140, 13 figs. (biology and taxonomy).

Genus SPHECODOGASTRA Ashmead

Sphecodogastra Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 92.
Type-species: Parasphecodes texana Cresson. Monotypic and orig. desig.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 364-366. —Linsley and
MacSwain, 1962. Pan-Pacific Ent. 38: 45.

Biology: Stevens, 1920. Ent. News 31: 39-44.
noctivaga (Linsley and MacSwain). Utah, N. Mex., west Tex. Pollen: Collects pollen from
flowers of Oenothera hartwegii, but visits flowers of other plants for nectar.
Lasioglossum (Sphecodogastra) noctivaga Linsley and MacSwain, 1962. Pan-Pacific Ent.
38: 46. 2.
texana (Cresson). N. Dak. and Mich., south to Tex., N. Mex. and Mexico, west to Colo., east to
Ind. Pollen: Oligolege of Oenothera including O. caespitosa, O. rhombipetala, O.
runcinata, but visits the flowers of other plants for nectar.
Parasphecodes texanus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 249. 2.

Superfamily APOIDEA 1963

Biology: Graenicher, 1911. Pub. Mus. Milwaukee, Bul. 1: 222-225. —Stevens, 1920. Ent. News
31: 39. —Hicks, 1936. Canad. Ent. 68: 51. —Chandler, 1962. Ind. Acad. Sci., Proc. for 1961,
71: 124-129, 2 figs., 2 tables. —Kerfoot, 1967. Kansas Ent. Soc., Jour. 40: 84-93, 9 figs., 2
tables (nest architecture, behavior). —Kerfoot, 1967. Anim. Behavior 15: 479-486 (lunar
periodicity).

Genus DIALICTUS Robertson

Dialictus Robertson, 1902. Canad. Ent. 34: 48.
Type-species: Halictus anomalus Robertson. Monotypic and orig. desig.
Chloralictus Robertson, 1902. Canad. Ent. 34: 248.
Type-species: Halictus cressoni Robertson. Orig. desig.
Halictus subg. Gastrohalictus Ducke, 1902. Ztschr. System. Hym. Dipt. 2: 102.
Type-species: Halictus osmioides Ducke. Monotypic.
Halictomorpha Schrottky, 1911. Rev. Mus. Paulista 8: 81.
Type-species: Halictomorpha phaedra Schrottky. Monotypic and orig. desig.

Revision: Sandhouse, 1923. Canad. Ent. 55: 193. —Sandhouse, 1924. U. S. Natl. Mus., Proc. 65
(19): 1-10. —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 368-445, figs. 95-102,
tables 11-12 (eastern U. S. spp.).

abanci (Crawford). Minn. to N. B., south to Ga.
Halictus abanci Crawford, 1932. Ent. Soe. Wash., Proc. 34: 71. 2.
abietus (Michener). Colo.
Halictus (Chloralictus) abietum Michener, 1938. Ann. and Mag. Nat. Hist. (10) 18: 281. °,
3.
absimilis (Sandhouse). Colo.
Halictus absimilis Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 21. °.
abundus (Sandhouse). N. Mex.
Halictus abundus Sandhouse, 1924. U. S. Natl. Mus., Proce. 65 (19): 32. ¢.
academicus (Sandhouse). Iowa, Colo.
Halictus academicus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 12. @.
Halictus vintonensis Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 22. 9.

Taxonomy: Michener, 1951. In Muesebeck, Krombein and Townes, U. S. Dept. Agr., Agr.

Monog. 2: 1111 (synonymy).
accentus (Sandhouse). Colo.

Halictus accentus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 38. ¢.
achilleae Mitchell. Mass. to N. C., Mich.

Dialictus achilleae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 377. 9.
actinosus (Sandhouse). Calif.

Halictus actinosus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 13. 2.
actuarius (Sandhouse). Colo.

Halictus actuarius Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 25. 6.
admirandus (Sandhouse). Minn. to N.S., south to La. and Fla.

Halictus admirandus Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 14. @.
advertus Mitchell. Mass. (Reading).

Dialictus advertus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 433, fig. 102. ¢.
alachuensis Mitchell. Ga., Fla.

Dialictus alachuensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 378. 2.
albipennis (Robertson). N. S. to Oreg., south to N. C., Ill., and Colo. Predator: Philanthus

politus politus Say.

Halictus albipennis Robertson, 1890. Amer. Ent. Soc., Trans. 17: 317. 2, d.

Halictus nubilis Lovell, 1905. Canad. Ent. 37: 40. 9.
albitarsis (Cresson). Tex.,? Ont.

Halictus albitarsis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 254. 6.

Taxonomy: Warncke, 1973. Soc. Royale Sci. Liege 42 (7-8): 294 (proposed replacement name).

1964 Hymenoptera in America North of Mexico

albohirtus (Crawford). Nev.
Halictus albohirtus Crawford, 1907. Invertebrata Pacifica 1: 193. 2.

albuquerquensis (Michener). N. Mex.
Halictus (Chloralictus) albuquerquensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19:
Gm oe
alius (Sandhouse). N. Mex., Ariz.
Halictus alius Sandhouse, 1924. U. S. Natl. Mus., Proce. 65 (19): 16. 2.
alternatus Mitchell. Ont. and Mass. to Va., Ohio, Mich.
Dialictus alternatus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 433. 6.
anomalus (Robertson). Alta. and Colo. east to N. S. south to Ala. and Ga.
Halictus anomalus Robertson, 1892. Amer. Nat. 26: 272. 9°.
apertus (Sandhouse). Minn. to Maine, south to La. and Ga. Predator: Philanthus lepidus Cress.
Halictus apertus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 35. ¢.
apocyni Mitchell. W. Va., Ohio, Ind., Tenn.
Dialictus apocyni Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 381. 2.
apopkensis (Robertson). N. C. to Fla., Miss.
Halictus apopkensis Robertson, 1892. Amer. Nat. 26: 272. °.
aquilae (Cockerell). N. Mex.; north. Mexico.
Halictus aquilae Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 450. °.

Taxonomy: Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 415. 6.
arcanus (Sandhouse). N. Mex.
Halictus arcanus Sandhouse, 1924. U. S. Natl. Mus., Proce. 65 (19): 19. @.
astutus (Sandhouse). N. Mex.
Halictus astutus Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 31. 6.
atlanticus Mitchell. Ont., Mass., N. Y., W. Va., N. C.
Dialictus atlanticus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 383. fig. 99. 9°.

Taxonomy: Knerer and Atwood, 1966. Canad. Ent. 98: 881. ¢.
atriventris (Crawford). Alta., B. C.
Halictus atriventris Crawford, 1906. In Viereck, Canad. Ent. 38: 303. 2, 6.
basilicus (Sandhouse). Minn. to Conn., N. H., Ont.
Halictus basilicus Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 36. d.

Taxonomy: Knerer and Atwood, 1966. Canad. Ent. 98: 881. °.
bernardinensis (Michener). Calif.
Halictus (Chloralictus) bernardinensis Michener, 1936. Ann. and Mag. Nat. Hist. (10) 18:
286. °.
brassicae Mitchell. N. C., Fla.
Dialictus brassicae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 384. °.
brevibasis (Cockerell). Sask.
Halictus (Chloralictus) brevibasis Cockerell, 1938. Amer. Mus. Novitates 983: 3. ¢.
bruneri (Crawford). Ont. and N. Y. to Fla., west to Nebr.
Halictus Bruneri Crawford, 1902. Canad. Ent. 34: 237. @.
Halictus brimleyi Crawford, 1932. Ent. Soc. Wash., Proc. 34: 71. °.
brunneiventris (Crawford). Nev., Calif.
Halictus brunneiventris Crawford, 1907. Invertebrata Pacifica 1: 194. 9.
cabrilli (Cockerell). Calif. (San Miguel Island).
Halictus (Chloralictus) cabrilli Cockerell, 1937. Pan-Pacifie Ent. 13: 155. 6.
caducus (Sandhouse). N. Mex.
Halictus caducus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 17. °.
californiae (Ellis). Calif.
Halictus perpunctatus var. californiae Ellis, 1924. U.S. Natl. Mus., Proe. 65 (19): 6. 2.
Lasioglossum (Chloralictus) calforniae(!) Michener, 1951. In Muesebeck, Krombein and
Townes, U.S. Dept. Agr., Agr. Monog. 2: 1112. Lapsuws calami.

Taxonomy: Timberlake, 1940. South. Calif. Acad. Sci., Bul. 39: 190. 9, d.

Superfamily APOIDEA 1965

callidus (Sandhouse). Il. and Mich. to New England, south to N. C.
Halictus callidus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 34. 6.
cattellae (Ellis). Ohio, Que., New England, south to Ga.
Halictus cattellae Ellis, 1913. Ent. News 24: 209. 2, 6.
ceanothi Mitchell. N. C., Mich., Mo.
Dialictus ceanothi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 386, fig. 99. 2.
clarissimus (Ellis). N. Mex., Ariz.
Halictus clarissimus Ellis, 1914. N. Y. Ent. Soe., Jour. 22: 222. 9.
clematisellus (Cockerell). N. Mex., Ariz., Calif., Utah. Predator: Philanthus multimaculatus
Cam.
Halictus clematisellus Cockerell, 1904. Canad. Ent. 36: 13. °.

Taxonomy: Cockerell, 1904. Entomologist 39: 6. d, °.
coactus (Cresson). Tex.
Halictus coactus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 254. 9.
coeruleus (Robertson). Minn. to Mass., south to Ga. Ecology: Nests in decomposed wood, but

has been induced to nest in the soil.
Halictus coeruleus Robertson, 1893. Amer. Ent. Soc., Trans. 20: 146. 9, d.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 167. —Stockhammer, 1967.
Kansas Ent. Soc., Jour. 40: 177-189 (life history). —Barrows, 1973. Kansas Ent. Soc., Jour.
46: 496-499 (induced soil nesting experimentation).
comulus (Michener). N. Mex.
Halictus comis Sandhouse, 1924. U. S. Natl. Mus., Proce. 65 (19): 18. 2. Preoce. by Vachal,
1911.
Lasioglossum (Chloralictus) comulum Michener, 1951. In Muesebeck, Krombein, Townes,
U.S. Dept. Agr., Agr. Monog. 2: 1113. N. name.
connexus (Cresson). Wis., Tex.
Halictus connexus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 253. 9.
coreopsis (Robertson). Ill. and Mich. to Mass., south to Fla.
Halictus longiceps Robertson, 1892. Amer. Nat. 26: 272. 2. Preoce. by Saunders, 1879.
Chloralictus cereopsis(!) Robertson, 1902. Canad. Ent. 34: 249. 2.
Chloralictus coreopsis Robertson, 1902. Canad. Ent. 34: 250. ¢.
Lasioglossum (Chloralictus) robertsonellum Michener, 1951. In Muesebeck, Krombein,
Townes, U.S. Dept. Agr., Agr. Monog. 2: 1117. N. name.

Biology: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 388-389, fig. 102 (synonymy,
redescription).
crassiceps (Ellis). N. Mex.
Halictus crassiceps Ellis, 1914. Ent. News 25: 108. °.
creberrimus (Smith). N. C. to Fla., La.
Halictus creberrimus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 72. 9.
Halictus ashmeadii Robertson, 1892. Amer. Nat. 26: 271. °.
Halictus ashmeadi Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 294. Emend.
cressonii (Robertson). N. S. to N. C., west to Alta., Wash., and Colo. Ecology: Nests in rotten
wood above ground. Predator: Philanthus lepidus Cress., P. politus politus Say.
Halictus Cressonii Robertson, 1890. Amer. Ent. Soc., Trans. 17: 317. 2, d.
Halictus (Chloralictus) cressoni Viereck, 1916. Conn. State Geol. and Nat. Hist. Survey,
Bul. 22: 707. Emend.
cyaneonotus (Crawford). Nev.
Halictus cyaneonotus Crawford, 1907. Invertebrata Pacifica 1: 193. °.
cyanurus (Cockerell). Calif.
Halictus cyanurus Cockerell, 1936. Pan-Pacifie Ent. 12: 157. °.
daggetti (Cockerell). Mexico (Baja California), near the border.
Halictus daggetti Cockerell, 1916. Canad. Ent. 48: 57. d.
delectatus Mitchell. N. B. to N. C., W. Va., Ohio, Ont.
Dialictus delectatus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 435. ¢.

1966 Hymenoptera in America North of Mexico

disbanci Knerer and Atwood. Ont. and Que.
Dialictus disbanci Knerer and Atwood, 1966. Canad. Ent. 98: 882. 2, ¢.
disparilis (Cresson). Kans., Tex., ?N. J., ?Alta., 7Nev.
Halictus disparilis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 253. 2.
diversopunctatus (Ellis). Idaho, Wash., Calif., Ariz.
Halictus diversopunctatus Ellis, 1914. Ent. News 25: 154. @.
dreisbachi Mitchell. N. Y., Md., Mich., Minn.
Dialictus dreisbachi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 391, fig. 101. 2,
3.
dubitatus Mitchell. N. Y. (Cortland Co.).
Dialictus dubitalus(!) Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 436. ¢. Spelled
dubitatus in key (p. 376) and in index (p. 533).
eophilius (Ellis). N. Mex., Ariz.
Halictus eophilius Ellis, 1914. Ent. News 25: 153. 9.
evestigatus (Sandhouse). Colo.
Halictus evestigatus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 27. 6.
fattigi Mitchell. N. C., Ga.
Dialictus fattigi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 392. °.
flaveriae Mitchell. Fla.
Dialictus flaveriae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 393, figs. 101, 102.
OMG
foveolatus (Robertson). IIl., Mich.
Chloralictus foveolatus Robertson, 1902. Canad. Ent. 34: 250. ¢.
genuinus (Sandhouse). Ont. to N. C.
Halictus genuinus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 36. ¢.
grinnelli (Cockerell). Calif.; Mexico (Baja California).
Halictus grinnelli Cockerell, 1916. Canad. Ent. 48: 56. 2.
halophitus (Graenicher). N. C. to Fla., La.
Halictus halophitus Graenicher, 1927. Psyche 34: 206. 2, ¢.
Halictus halophilus Graenicher, 1930. Ent. Soc. Amer., Ann. 23: 156. Emend.
hartii (Robertson). Nebr., Minn. and IIl. to La. and N. C.
Halictus hartii Robertson, 1892. Amer. Nat. 26: 268. °.
Halictus rugosus Crawford, 1902. Canad. Ent. 34: 237. @, 6.
hemimelas (Cockerell). Colo., N. Mex.
Halictus hemimelas Cockerell, 1901. Psyche, 9: 285. 2.
heterognathus Mitchell. N. B. to Minn., south to N. C. and Tenn.
Dialictus heterognathus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 397, figs. 95,
99. 2.
Dialictus banksi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 434, fig. 102. 3.

Taxonomy: Knerer and Atwood, 1963. Ent. Soc. Wash., Proc. 65: 167. 6. —Knerer and
Atwood, 1966. Canad. Ent. 98: 882 (synonymy).
highlandicus Mitchell. N. C., Ont.
Dialictus highlandicus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 398. 9.
hudsoniellus (Cockerell). Colo., Ariz.
Halictus hudsoniellus Cockerell, 1919. Ent. News 30: 290. °.
hunteri (Crawford). Tex. to Calif.
Halictus hunteri Crawford, 1932. Ent. Soc. Wash., Proc. 34: 72. °.
hyalinus (Crawford). Wash. to Utah, Ariz., Nev. and Calif.
Halictus hyalinus Crawford, 1907. Invertebrata Pacifica 1: 194. 9.
illinoensis (Robertson). Nebr., and Minn. to N. S., south to Tex. and Ga.
Halictus illinoensis Robertson, 1892. Amer. Nat. 26: 271. ?.
Halictus illinoisensis Smith, 1910. N. J. State Mus., Ann. Rpt. 1909, p. 688. Emend.
Halictus (Chloralictus) politissimus Cockerell, 1917. In W. P. Cockerell, N. Y. Ent. Soc.,
Jour. 25: 189. 2. é

Superfamily APOIDEA 1967

imitatus (Smith). New England and Que. west to Minn., south to Fla. and west to Calif.

(Riverside Co.). Parasite: Paralictus cephalotes (Dalla Torre), Psewdomethoca frigida
frigida (Sm.). Predator: Philanthus albopilosus Cress., P. bilunatus Cress., P. gibbosus
(Fabr.), P. lepidus Cress., P. politus politus Say, P. ventilabris Fabr.

Halictus imitatus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 71. 6.

Halictus inconspicuus Smith, 1853. Cat. Hym. Brit. Mus., Veuls pa(oamer

Halictus stultus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 254. 9.

Chloralictus sparsus Robertson, 1902. Canad. Ent. DANZA DONG

Halictus hortensis Lovell, 1905. Canad. Ent. 37: 39. 2.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 350 (as imitatus). —Cockerell, 1905.
Amer. Ent. Soce., Trans. 31: 349 (as inconspicuus). —Michener, 1953. Kansas Univ. Sci.
Bul. 35:-1027 (larva, as sparsus). —Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92:
168.

Biology: Michener and Wille, 1961. Kansas Univ. Sci. Bul. 42: 1123-1202, 44 figs., 11 tables (as
inconspicuus).

Morphology: Knerer and Atwood, 1964. Ent. Soc. Wash., Proc. 66: 111-112 (metanotal
anomaly).
impavidus (Sandhouse). Calif., Nev., Ariz.
Halictus impavidus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 18. 9.
impurus (Cresson). Tex.
Halictus impurus Cresson, 1872. Amer. Ent. Soc., Trans. 4: 252. 9.
incompletus (Crawford). N. Mex., Nev., Calif. Predator: Philanthus crabroniformis Sm., P.
gibbosus (Fabr.), P. pacificus arizonae Dunn.
Halictus incompletus Crawford, 1907. Invertebrata Pacifica 1: 195. °.

Taxonomy: Michener, 1936. Ann. and Mag. Nat. Hist. (10) 19: 284. ¢.

insolitus (Sandhouse). Iowa.

Halictus insolitus Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 41. 6.
insulsus (Sandhouse). N. Mex.

Halictus insulsus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 29. 3.
intrepidus Mitchell. Ga. (Stone Mt.).

Dialictus intrepidus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 487. ¢.
jameseae (Cockerell). Colo.

Halictus (Chloralictus) jameseae Cockerell, 1933. Ent. Soc. Amer., Ann. 26: 41. d.
junaluskensis Mitchell. N. C., N. Y., Mich., Minn.

Dialictus junaluskensis Mitchell, 1960 N. C. Agr. Expt. Sta. Tech. Bul. 141: 487. d.
kunzei (Cockerell). Ariz., Calif.

Halictus Kunzei Cockerell, 1898. Canad. Ent. 30: 238. °.
lactineus (Sandhouse). Colo.

Halictus lactineus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 34. 6.
laevissimus (Smith). N. S. to Alta., south to Ga. and Calif. Predator: Philanthus

crabroniformis (Sm.), P. gibbosus (Fabr.), P. lepidus Cress., P. pacificus arizonae Dunn.,
P. pulcher Dalla Torre.

Halictus laevissimus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 72. 2.

Halictus levissimus Dalla Torre, 1896. Cat. Hym., v. 10, p. 68. Emend.

Halictus smilacinae Robertson, 1899. Acad. Sci. St. Louis, Trans. 7: 322. 9.

Halictus euryceps Ellis, 1914. Ent. News 25: 98. 2.

Taxonomy: Knerer and Atwood, 1962. Canad. Ent. 94: 1228. d.
Biology: Brittain, 1933. Canada Dept. Agr. Bul. (n. s.) 162: 94 (as smilacinae). —Knerer and
Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 168.
latus (Sandhouse). Colo.
Halictus latus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 20. °.
lazulis (Ellis). Colo.
Halictus lazulis Ellis, 1913. Ent. News 24: 207. °.

1968 Hymenoptera in America North of Mexico

lectus Mitchell. Ga., D. C., N. J., Ind.
Dialictus lectus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 438. ¢.
leviensis Mitchell. N. C. to Fla.
Dialictus leviensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 403. °.
lineatulus (Crawford). Minn. to Newfoundland, south to Tex. and Ga. Predator: Philanthus
albopilosus Cress., P. gibbosus (Fabr.)., P. politus politus Say, P. solivagus Say.
Halictus lineatulus Crawford, 1906. Canad. Ent. 38: 5. 2.
Halictus subconnexus Ellis, 1915. Ent. News 26: 291. 2.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 169.

Morphology: Knerer and Atwood, 1964. Ent. Soc. Wash., Proc. 66: 111-112 (metanotal

anomaly).
lionotus Sandhouse. Colo.

Dialictus lionotus Sandhouse, 1923. Canad. Ent. 55: 194. 6.
longicornis (Crawford). Calif.

Halictus longicornis Crawford, 1907. Invertebrata Pacifica 1: 195. d.
mactus (Sandhouse). Colo.

Halictus mactus Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 37. 3.
malinus (Sandhouse). Va.

Halictus malinus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 40. d.
marinensis (Michener). Calif.

Halictus (Chloralictus) marinensis Michener, 1936. Pan-Pacifie Ent. 12: 167. °.
marinus (Crawford). Mass. to Fla.

Halictus marinus Crawford, 1904. Ent. News 15: 99. °.

Taxonomy: Graenicher, 1927. Psyche 34: 204. 6. —Graenicher, 1930. Ent. Soc. Amer., Ann.
23: 156.
megastictus (Cockerell). Calif. (San Miguel Island).
Halictus (Chloralictus) megastictus Cockerell, 1937. Pan-Pacifie Ent. 13: 152. 2, 6.
meritus (Sandhouse). Colo.
Halictus meritus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 19. 2.
merosus (Sandhouse). N. Mex.
Halictus merosus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 40. d.
mesillensis (Cockerell). Tex. to Nev. and Calif.; Mexico (Hidalgo).
Halictus nymphalis race mesillensis Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 47.
2. Republished by Cockerell, 1898. N. Mex. Univ., Bul. 1: 47. female.
microlepoides (Ellis). N. Mex., Ariz., Calif.; north. Mexico. Predator: Philanthus gibbosus
(Fabr.), P. multimaculatus Cam., P. ventilabris Fabr.
Halictus microlepoides Ellis, 1914. Ent. News 25: 152. 9.

Biology: Aleock and Gamboa, 1975. Ariz. Acad. Sci. 10: 163 (predator).
miniatulus Mitchell. Fla.
Dialictus miniatulus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 405. °.
mollis (Sandhouse). Colo.
Halictus mollis Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 39. d.
nevadensis (Crawford). Ariz., Nev., Calif.; Mexico (Baja California).
Halictus nevadensis Crawford, 1907. Invertebrata Pacifica 1: 195. °.
Halictus (Chloralictus) pasadenae Michener, 1936. Ann. and Mag. Nat. Hist. (10) 18: 285.
2. Oe
Taxonomy: Timberlake Jn Michener, 1951. In Muesebeck, Krombein and Townes, U. S. Dept.
Agr., Agr. Monog. 2: 1115 (synonymy).
nigroviridis (Graenicher). B. C. to N. B., south to Ga.
Halictus nigro-viridis Graenicher, 1910. Pub. Mus. City Milwaukee, Bul. 1: 2338. ?.
novascotiae Mitchell. N. S. to New England.
Dialictus novascotiae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 407. °.

Taxonomy: Knerer and Atwood, 1964. Ent. News 75: 5, fig. 1. 6.

Superfamily APOIDEA 1969

nymphaearum (Robertson). Minn. and Kans. to N.S., south to Ga.
Halictus palustris Robertson, 1890. Amer. Ent. Soc., Trans. 17: 317. 2, d. Preoce.
Halictus nymphaearum Robertson, 1895. Amer. Ent. Soc., Trans. 22: 117. N. name.
Halictus paludicola Dalla Torre, 1896. Cat. Hym., v. 10, p. 75. N. name.
Halictus oceanicus Cockerell, 1916. Brooklyn Ent. Soe., Bul. 11: 11. 2.

nymphalis (Smith). Minn. and Mass. south to Tex. and Fla.
Halictus nymphalis Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 68. °.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 352.
oblongus (Lovell). Colo., Minn. and N. S., south to N. Mex., La. and Ga. Ecology: Nests in
rotten wood above ground.
Halictus oblongus Lovell, 1905. Canad. Ent. 37: 40. 2.
Halictus planatus Lovell, 1905. Canad. Ent. 37: 300. °.

Taxonomy: Lovell, 1908. Psyche 15: 39. d.
obnubilus (Sandhouse). Colo.
Halictus (Chloralictus) obnubilus Sandhouse, 1924. U. S. Natl. Mus., Proe. 65 (19): 28. d.
obscurus (Robertson). Minn. and III. to Conn. and Ont., south to Ga. Predator: Philanthus
gibbosus (Fabr.).
Halictus obscurus Robertson, 1892. Amer. Nat. 26: 270. °.
occidentalis Crawford. Colo., N. Mex.
Dialictus occidentalis Crawford, 1902. Canad. Ent. 34: 318. 2.
Halictus galei Cockerell, 1919. Canad. Ent. 51: 272. °.
oleosus (Cockerell). Colo., N. Mex.
Halictus oleosus Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 47. 9. Republished by
Cockerell, 1898. N. Mex. Univ., Bul. 1: 47, female.
orbitatus Mitchell. Minn. to Conn. and Ont., south to N. C. and W. Va.
Dialictus orbitatus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 440, fig. 101. d.
ornduffi Hurd. Calif. Pollen: Apparently a narrowly polylectic visitor to the flowers of
Jepsonia heterandra, but also visits flowers of Compositae apparently for nectar.
Dialictus ornduffi Hurd, 1970. Pan-Pacifie Ent. 46: 210. 2, d.

Biology: Ornduff, 1971. Evolution 25: 300-311, 1 fig., 7 tables (role in pollination of Jepsonia
heterandra).
otsegoensis Mitchell. Mich. (Otsego Co.).
Dialictus otsegoensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 440. d.
pacatus (Sandhouse). Colo.
Halictus pacatus Sandhouse, 1924. U. S. Natl. Mus., Proce. 65 (19): 16. °.
pallidellus (Ellis). N. Mex., Utah, Calif.
Halictus pallidellus Ellis, 1914. Ent. News 25: 151. °.
paradmirandus Knerer and Atwood. Ont.
Dialictus paradmirandus Knerer and Atwood, 1966. Canad. Ent. 98: 886. ?, d.

paululus (Sandhouse). Colo.
Halictus paululus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 2350r

pavoninus (Ellis). Colo.
Halictus pavoninus Ellis, 1913. Ent. News 24: 206. °.
pensitus (Sandhouse). Colo.
Halictus pensitus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 38. d.
perdifficilis (Cockerell). Colo., N. Mex.; north. Mexico.
Halictus perdifficilis Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 68. 2.
Halictus perfficilis(!) Cockerell, 1897. N. Mex. Agr. Expt. Sta., Bul. 24: 19.

Taxonomy: Cockerell, 1901. Ann. and Mag. Nat. Hist. (7) 7: 126.

perexiguus (Sandhouse). N. Mex.
Halictus perexiguus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 26. d.

perichlarus (Cockerell). Calif.
Halictus (Chloralictus) perichlarus Cockerell, 1937. Pan-Pacifie Ent. 13: 153. 2, 6.
Halictus perichlorus Cockerell, 1939. Calif. Acad. Sci., Proe. (4) 23: 428. Emend.

1970 Hymenoptera in America North of Mexico

perparvus (Ellis). Ariz., Calif; Mexico. Predator: Philanthus multimaculatus Cam.

Halictus perparvus Ellis, 1914. Ent. News 25: 102. 9.
perpunctatulus Knerer and Atwood. Boreal Canada.

Dialictus perpunctatulus Knerer and Atwood, 1966. Canad. Ent. 98: 884. 2, d.
perpunctatus (Ellis). N. B. to Ga., west to N. Mex., Colo. and Calif.

Halictus perpunctatus Ellis, 1913. Ent. News 24: 210. @.

Halictus brycinci Crawford, 1932. Ent. Soe. Wash., Proc. 34: 70. °.
perspicuus Knerer and Atwood. Ont., south to N. Y., west to Minn.

Dialictus perspicuus Knerer and Atwood, 1966. Canad. Ent. 98: 883. 2, 6.
petrellus (Cockerell). Calif.; Mexico (Baja California).

Halictus petrellus Cockerell, 1903. South. Calif. Acad. Sci., Bul. 2: 84. 9.

Halictus coronadensis Cockerell, 1916. Canad. Ent. 48: 56. 6.
phaceliarum (Cockerell). Colo.

Halictus phaceliarum Cockerell, 1919. N. Y. Ent. Soe., Jour. 27: 299. 9°.
philanthanus Mitchell. N. C., N. Y., Mich., Ont. Predator: Philanthus sp.

Dialictus philanthanus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 441. ¢.
pictus (Crawford). Nebr., Minn., Wis., Mich., Ill., Ont.

Halictus pictus Crawford, 1902. Canad. Ent. 34: 236. 9.

Halictus graenicheri Ellis, 1914. N. Y. Ent. Soe., Jour. 22: 221. @.

Dialictus muskegonensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 439. d.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 170.

pikei (Sandhouse). Colo.
Halictus pikei Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 28. ¢.
pilosellus (Cockerell). Calif.
Halictus pilosellus Cockerell, 1936. Pan-Pacific Ent. 12: 160. °.
pilosicaudus (Cockerell). Calif. (San Miguel Isl.).
Halictus (Chloralictus) pilosicaudus Cockerell, 1937. Pan-Pacifie Ent. 13: 155. 2.
pilosus floridanus (Robertson). N. C. to Fla.
Halictus floridanus Robertson, 1892. Amer. Nat. 26: 269. 2.
pilosus pilosus (Smith). Colo. and Minn. to N. S., south. to Ga. Parasite: Sphecodes atlantis
Mitchell. Predator: Philanthus albopilosus, P. gibbosus (Fabr.), P. politus politus Say.
Halictus pilosus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 71. ?.
Halictus pilosus var. leucocomus Lovell, 1908. Psyche 15: 37. °.
Halictus floridanus caesareus Cockerell, 1916. Brooklyn Ent. Soc., Bul. 11: 11. 2.

Taxonomy: Robertson, 1895. Amer. Ent. Soc., Trans. 22: 117. 6. —Cockerell, 1905. Amer.
Ent. Soc., Trans. 31: 351. —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 152: 547.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 170.

placidensis Mitchell. Fla.
Dialictus placidensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 441. 6.
praepes (Sandhouse). Colo.
Halictus praepes Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 30. 3.
pruinosiformis (Crawford). Alta., S. D., Iowa, Nebr., N. Mex., Tex., Ariz., Nev., Calif. Predator:
Philanthus multimaculatus Cam.
Halictus pruinosiformis Crawford, 1906. Canad. Ent. 38: 284. 2, d.
Halictus glaucovirens Cockerell, 1919. Ent. News 30: 290. °.
Halictus exalbidus Sandhouse, 1924. U.S. Natl. Mus., Proce. 65 (19): 24. 6.
pruinosus (Robertson). New England states, south to N. J., west to Alta. and Ariz. Parasite:
Rhipiphorus fasciatus Say. Predator: Philanthus ventilabris Fabr.
Halictus pruinosus Robertson, 1892. Amer. Nat. 26: 269. °, d.

Biology: Melander and Brues, 1903. Biol. Bul. 5: 1.

pseudotegularis (Cockerell). N. Mex.; Mexico
Halictus pseudotegularis Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 294. °.

pudicus (Sandhouse). Colo.
Halictus pudicus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 33. d.

Superfamily APOIDEA 1971

punctatoventris (Crawford). Calif.; Mexico (Baja California).

Halictus punctatoventris Crawford, 1907. Invertebrata Pacifica 1: 196. 9, d.
punctiferellus (Cockerell). Calif. (San Miguel Island).

Halictus (Chloralictus) punctiferellus Cockerell, 1937. Pan-Pacifie Ent. 13: 154. 2.
raleighensis (Crawford). N. C. to Fla.

Halictus raleighensis Crawford, 1932. Ent. Soc. Wash., Proce. 34: 73. 9.
reticulatus (Robertson). Ill. and Mich. to Maine, south to Fla.

Halictus fulvipes Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 67. 2. Preoce.

Halictus reticulatus Robertson, 1892. Amer. Nat. 26: 268. °.

Halictus rhododactylus Dalla Torre, 1896. Cat. Hym., v. 10, p. 80. 9. N. name.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 351. —Mitchell, 1958. In Krombein,
U.S. Dept. Agr., Agr. Monog. 2, sup. 1: 230. (synonymy).
rhodognathus (Cockerell). Tex.
Halictus (Chloralictus) rhodognathus Cockerell, 1917. In W. P. Cockerell, N. Y. Ent. Soc.,
Jour. 25: 190. 2.
richardsoni (Cockerell). Alta.
Halictus (Chloralictus) richardsoni Cockerell, 1937. Canad. Ent. 69: 113. °.
rohweri (Ellis). Minn. to Maine and Ont., south to Ga. Parasite: Psewdomethoca frigida frigida
(Sm.). Predator: Philanthus gibbosus (F.).
Halictus subconnexus rohweri Ellis, 1915. Ent. News 26: 292. °.

Taxonomy: Knerer and Atwood, 1962. Canad. Ent. 94: 12380. d.

Biology: Knerer and Atwood, 1962. Ent. Soc. Ontario, Proc. 92: 171. —Breed, 1975. Kansas
Ent. Soc., Jour. 48: 64-80, 3 figs., 7 tables (life cycle, behavior).
rufulipes (Cockerell). Sask.
Halictus (Chloralictus) rufulipes Cockerell, 1938. Amer. Mus. Novitates 983: 3. 2.
ruidosensis (Cockerell). Colo., N. Mex., Ariz. Predator: Philanthus pacificus arizonae Dunn.,
P. pulcher Dalla Torre.
Halictus ruidosensis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 142. 9, d.
sagax (Sandhouse). Colo.
Halictus sagax Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 25. d.
sandhouseae (Michener). Colo., Minn., Ont.
Halictus occultus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 27. 3. Preoce. by

Vachal, 1904.
Lasioglossum (Chloralictus) sandhouseae Michener, 1951. In Muesebeck, Krombein,

Townes, U. S. Dept. Agr., Agr. Monog. 2: 1117. N. name.

Taxonomy: Knerer and Atwood, 1964. Ent. News 75: 7. (female).

scrophulariae (Cockerell). Colo.
Halictus (Chloralictus) scrophulariae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 428.
OF
sedi (Sandhouse). Calif., Oreg., Colo.
Halictus sedi Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 12. °.

Taxonomy: Timberlake Jn Michener, 1958. In Krombein, U.S. Dept. Agr., Agr. Monog. 2,
Sup. 1: 230 (tax. status).
semibrunneus (Cockerell). N. Mex.
Halictus semibrunneus Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 67. 9.
semicaeruleus (Cockerell). N. Mex.
Halictus semicaeruleus Cockerell, 1895. Ann. and Mag. Nat. Hist. (6) 16: 66. 9.
solidaginis Mitchell. Mich. to Maine, south to Ind., W. Va., and N. Y.
Dialictus solidaginis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 448, fig. 102. d.
stictaspis Sandhouse. N. Mex.
Dialictus stictaspis Sandhouse, 1923. Canad. Ent. 55: 195. d.

stuartensis Mitchell. Fla. (Stuart).
Dialictus stuartensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 419. °.

1972 Hymenoptera in America North of Mexico

subversans Mitchell. Mich., N. Y.
Dialictus subversans Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 419. 2.
subviridatus (Cockerell). Sask.
Halictus (Chloralictus) subviridatus Cockerell, 1938. Amer. Mus. Novitates 983: 2. 2, d.
succinipennis (Ellis). Colo.
Halictus succinipennis Ellis, 1913. Ent. News 24: 205. °.
supraclypeatus Mitchell. Va., Mich., Tenn., Mo.
Dialictus supraclypeatus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 420, fig. 99.
OF
surianae Mitchell. Fla.
Dialictus surianae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 420. 9.
tahitensis Mitchell. Fla. (Tahiti Beach).
Dialictus tahitensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 421. 9.
tamiamensis Mitchell. Fla.
Dialictus tamiamensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 421, figs. 98,
101, 102. 2, 6.
tarponensis Mitchell. Fla., Ga.
Dialictus tarponensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 423. 9.
tegulariformis (Crawford). Idaho, Utah, Nev., N. Mex., Ariz., Calif.; Mexico (Baja California).
Predator: Philanthus pacificus arizonae Dunn.
Halictus tegulariformis Crawford, 1907. Invertebrata Pacifica 1: 194. 2, ¢.
Halictus helianthi Cockerell, 1916. Canad. Ent. 48: 77. °.
Halictus gaudialis Sandhouse, 1924. U. S. Natl. Mus., Proe. 65 (19): 28. 3.

Taxonomy: Michener, 1936. Ann. and Mag. Nat. Hist. (10) 18: 283. ¢.

Biology: Timberlake Jn Michener, 1951. Jn Muesebeck, Krombein, Townes, U.S. Dept. Agr.,
Agr. Monog. 2: 1118 (synonymy).
tegularis (Robertson). Minn. to N. H., south to Tex. and Fla.; recorded probably erroneously
from Mont., Colo., Calif., and Mexico. Predator: Philanthus lepidus Cress.
Halictus tegularis Robertson, 1890. Amer. Ent. Soc., Trans. 17: 318. 2, 6.
Halictus ellisiae Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 11. 9.
Halictus lepidvi Graenicher, 1927. Psyche 34: 204. 2, ¢.
tenax (Sandhouse). Colo.
Halictus tenax Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 15. 9.
tenuis (Ellis). Colo.
Halictus tenuis Ellis, 1913. Ent. News 24: 208. 9.
testaceus (Robertson). N. C., Wis., IIl., N. Dak.
Halictus testaceus Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 323. 2.
theodori Crawford. N. Mex.
Dialictus Theodori Crawford, 1902. Canad. Ent. 34: 318. °.
tranquillus (Sandhouse). N. Mex.
Halictus tranquillus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 32. ¢.
unicus (Sandhouse). Minn., Mich., N. H., Ont., Va., N. C.
Halictus wnicus Sandhouse, 1924. U. S. Natl. Mus., Proc. 65 (19): 21. @.

vanduzeei (Sandhouse and Cockerell). South. Calif. (Inyo County); Mexico (Baja California).
Halictus vanduzeei Sandhouse and Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 13: 334. °.

veganus (Cockerell). N. Mex., Calif.
Halictus veganus Cockerell, 1901. Psyche 9: 286. 2.
versans (Lovell). Minn. to Newfoundland, south to Ga.; reportedly introduced in vicinity of San
Francisco. Predator: Philanthus lepidus Cress., P. solivagus Say.
Halictus versans Lovell, 1905. Canad. Ent. 37: 39. 2 (d misdet.).
Halictus consonus Sandhouse, 1924. U.S. Natl. Mus., Proc. 65 (19): 30. 6.
versatus (Robertson). Minn. to Ont., south to Ga. and La. Parasite: Psewdomethoca frigida
frigida (Sm.)? Predator: Philanthus ventilabris Fabr.
Chloralictus versatus Robertson, 1902. Canad. Ent. 34: 249. 2, 6.

Superfamily APOIDEA 1973

Biology: Michener, 1966. Kansas Ent. Soc., Jour. 39: 193-217 (bionomics).

vierecki (Crawford). Minn. to New England, south to La. and Ga.
Halictus vierecki Crawford, 1904. Ent. News 15: 97. 9.

Taxonomy: Graenicher, 1910. Canad. Ent. 42: 158. 6.
viridatulus (Cockerell). Colo., Calif.
Halictus viridatulus Cockerell, 1919. Ent. News 30: 290. 9.
viridatus (Lovell). Minn. to N. S., south to La. and Ga.
Halictus viridatus Lovell, 1905. Canad. Ent. 37: 300. @.
Dialictus lepidus Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 438. 6.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 547.

Biology: Atwood, 1933. Canad. Jour. Res. 9: 453.
weemsi Mitchell. N. C., Ga.
Dialictus weemsi Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 431, fig. 99. °.
wheeleri Mitchell. Mass. (Forest Hills).
Dialictus wheeleri Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 445. d.
zephyrus (Smith). Maine to Ga., west to Minn., Colo. and Tex., Utah, Oreg. Parasite: Pa ralictus
cephalotes (Dalla Torre), Pseudomethoca frigida (Sm.). Predator: Philanthus gibbosus
(F.).
Halictus zephyrus Smith, 1853. Cat. Hym. Brit. Mus., v. 2, p. 68. 6.

Taxonomy: Robertson, 1895. Amer. Ent. Soc., Trans. 22: 117. 2. —Cockerell, 1905. Amer.
Ent. Soc., Trans. 31: 352. 6.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7): 1. —Rau, 1926. Acad. Sci. St. Louis,
Trans. 25: 157. —LaBerge and Isakson, 1963. Ent. News 74: 113-116. —Lin, 1964. Ins.
Sociaux 11: 187-192 (reaction to mutillid parasitism). —Batra, 1964. Ins. Sociaux 11:
159-186, 20 figs. (behavior). —Batra, 1965. Kansas Ent. Soc., Jour. 38: 367-389 (associated
organisms). —Batra, 1966. Kansas Univ. Sci. Bul 46: 359-423 (life cycle, behavior).
—Michener and Brothers, 1971. Kansas Ent. Soc., Jour. 44: 236-239. —Michener, Brothers
and Kamm, 1971. Kansas Ent. Soe., Jour. 44: 276-279 (queen-worker relationships).
—Michener, Brothers and Kamm, 1971. Natl. Acad. Sci. U. S. A., Proc. 68: 1241-1245
(interactions within colonies). —Bell, 1973. Ins. Sociaux 20: 189-196. —Bell, 1973. Ins.
Sociaux 20: 253-260 (factors controlling initiation of vitellogenesis). —Barrows and Snyder,
1973. Ent. News 84: 314-316. —Kamm, 1974. Kansas Ent. Soc., Jour. 47: 8-18 (effects of
temperature, day length and number of adults on size of cells and offspring). —Brothers
and Michener, 1974. Jour. Compar. Physiol. 90: 129-168 (ethometry of division of labor).
—Brothers, 1974. Acad. Natl. Sci. U. S. A., Proc. 71: 671-674 (origin of altruism). —Bell,
Breed, Richards and Michener, 1974. Jour. Compar. Physiol. 93: 173-181 (nest defense).
—Bell and Hawkins, 1974. Jour. Compar. Physiol. 13: 183-193 (nest defense). —Bell, 1974.
Jour. Compar. Physiol. 13: 195-202 (intraspecific defense of nests). —Barrows, 1975.
Behavioral Biol. 15: 57-64 (female odor production, male response). —Kumar, 1975. Kansas
Ent. Soc., Jour. 48: 374-380, 3 tables (cell size). —Barrows, Bell, and Michener, 1975. Natl.
Acad. Sci. U.S. A., Proc. 72: 2824-2828 (individual odor differences and their social
functions). —Breed and Gamboa, 1977. Science 195: 694-696, 2 figs. (behavioral control of
workers by queens).

zophops (Ellis). Colo.
Halictus zophops Ellis, 1914. Ent. News 25: 97. °.

Genus PARALICTUS Robertson

Paralictus Robertson, 1901. Canad. Ent. 33: 299.
Type-species: Halictus cephalotes Dalla Torre. Orig. desig. (=Halictus cephalicus
Robertson).

Members of this genus are believed to be inquilines in the nests of the genus Dialictus.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 445-450, figs. 103, 104, table
13 (included species).

1974 Hymenoptera in America North of Mexico

asteris Mitchell. N. Y. to N. C., west to Mich., Ind. and Tenn. Ecology: Visits flowers of Aster
and Robinia.
Paralictus asteris Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 446. 2, d.
cephalotes (Dalla Torre). Minn. to Conn., south to N. C. Ecology: Visits flowers of Gonolobus
laevis, Leucanthemum, Salix nigra. Host: Dialictus imitatus (Sm.).
Halictus cephalicus Robertson, 1892. Amer. Nat. 26: 270. 2, 3. Preoce.
Halictus cephalotes Dalla Torre, 1896. Cat. Hym., v. 10, p. 57. N. name.
michiganensis Mitchell. Mich. (Wayne Co.).
Paralictus michiganensis Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 148. 9.
platyparius (Robertson). Minn., Mich., and Md., south to N. C. Ecology: Visits flowers of Aster,
Cucurbita, Erigeron, Gnaphalium, Gonolobus, Koellia, Rhus, Salix, Solidago,
Taraxacum, Zanthoxylum, Zizia.
Halictus platyparius Robertson, 1895. Amer. Ent. Soc., Trans. 22: 117. 9.
simplex Robertson. Mich. to Mass., south to N. C. Ecology: Visits flowers of Achillea,
Ranunculus, Rubus, Solidago, Zizia.
Paralictus simplex Robertson, 1901. Canad. Ent. 33: 230. 9.

Genus SPHECODES Latreille

Sphecodes Latreille, 1804. Nouv. Dict. Hist. Nat., v. 24, p. 182.
Type-species: Sphex gibba Linnaeus. Monotypic.
Dichroa Mlliger, [1806]. Mag. Insektenk. 5: 39.
Type-species: Sphex gibba Linnaeus. Desig. by Sandhouse, 1943.
Sabulicola Verhoeff, 1890. Ent. Nachr. 16: 328.
Type-species: Sabulicola cirsii Verhoeff. Monotypic.
Drepanium Robertson, 1903. Ent. News 14: 103.
Type-species: Sphecodes falcifer Patton. Monotypic.
Proteraner Robertson, 1903. Ent. News 14: 103.
Type-species: Sphecodes ranunculi Robertson. Monotypic.
Sphecodium Robertson, 1903. Ent. News 14: 104.
Type-species: Sphecodes mandibularis Cresson. Orig. desig. (=Sphecodium
cressonti Robertson).
Machaeris Robertson, 1903. Ent. News 14: 104.
Type-species: Sphecodes stygius Robertson. Orig. desig.
Dialonia Robertson, 1903. Ent. News 14: 104.
Type-species: Sphecodes antennariae Robertson. Monotypic and orig. desig.
Sphecodes subg. Microsphecodes Eickwort and Stage, 1972. Kans. Ent. Soc., Jour. 45: 501.
Type-species: Sphecodes kathleenae Eickwort. Orig. desig.

This is a large and nearly worldwide genus of parasitic bees. While most of the species are
cleptoparasites in the nests of other Halictinae, some are parasitic in the nests of certain species
of Andrenidae and Colletidae. The adults visit a wide variety of flowers often in the company of
their pollen collecting hosts.

There are obviously several subgenera of the genus Sphecodes represented in the Nearctic
Region and some of the names cited above are available. However, until further studies are
made, it is not possible to establish the limits of the subgenera present in our area.

Taxonomy: Robertson, 1903. Ent. News 14: 103 (Illinois species). —Cockerell, 1907. Ann. and

Mag. Nat. Hist. (7) 20: 450-451 (key to large species). —Lovell and Cockerell, 1907. Psyche
14: 101 (Maine species). —Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 469-506
(eastern U.S. spp.). —Torchio, 1975. Kans. Ent. Soc., Jour. 48: 264-265, 269-275, 278-279,
figs. 15-18, 27 (life history, larvae, pupae).

antennariae Robertson. Wis. and Mich. to N. C.
Sphecodes antennariae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 63. 9.

aroniae Mitchell. Va., N.C.
Sphecodes aroniae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 474. 2, 6.

arroyanus Cockerell. Alta., N. D., Colo., N. Mex., Calif.
Sphecodes arroyanus Cockerell, 1904. Entomologist 37: 231. °.

Superfamily APOIDEA 1975

arvensiformis Cockerell. Colo., Utah, Oreg., Calif. Host: Halictus farinosus Sm.
Sphecodes arvensiformis Cockerell, 1904. In Viereck, Canad. Ent. 36: 232. 9.
Sphecodes levequeae Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 296. °.

Taxonomy: Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 191. d.
asclepiadis Cockerell. N. Mex.
Sphecodes asclepiadis Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 45. 2.
Republished by Cockerell, 1898. N. Mex. Univ., Bul. 1: 45. female.
atlantis Mitchell. Vt. and Mass., south to Fla., west to Tex., Ark., Tenn., Mich., Minn. Host:
Dialictus pilosus pilosus (Sm.).
Sphecodes atlantis Mitchell, 1956. Elisha Mitchell Sci. Soe., Jour. 72: 221. 2, d.
autumnalis Mitchell. Mich., Ont. and N. Y., south to N. C.
Sphecodes autumnalis Mitchell, 1956. Elisha Mitchell Sei. Soc., Jour. 72: 20% °, d.
banksii Lovell. Minn. to N. Y., south to Fla.
Sphecodes banksii Lovell, 1909. Ent. News 20: 416. °.

borealis Cockerell. Sask.

Sphecodes borealis Cockerell, 1937. Amer. Mus. Novitates 909: 2. 6.
brachycephalus Mitchell. N. C. to Fla. Host: Calliopsis andreniformis Sm?

Sphecodes brachycephalus Mitchell, 1956. Elisha Mitchell Sci. Soe., Jour. 72: 207. 2, d.
californicus Meyer. Calif.; Mexico.

Sphecodes californicus Meyer, 1922. Arch. Naturgesch. 88 (Abt. A, Heft. 8): 173. °.
carolinus Mitchell. Minn. to Maine, south to Tenn. and N. C.

Sphecodes carolinus Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 219. °.

castaneae Mitchell. Va. (Falls Church).
Sphecodes castaneae Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 480. °.

clematidis Robertson. Man. and Kans. to N.S., south to N. C.
Sphecodes clematidis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 320. ?, é.
Sphecodes obscurans Lovell and Cockerell, 1907. Psyche 14: 103. °.
Sphecodes australis Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 213. d.
columbiae Cockerell. Wash.
Sphecodes columbiae Cockerell, 1906. Canad. Ent. 38: 280. ?.
confertus Say. Minn. and Colo to N. S., Tex. and Ga.
Sphecodes confertus Say, 1837. Boston Jour. Nat. Hist. 1: 392.
Sphecodes falcifer Patton, 1880. Amer. Ent. 1: 280. 2, d.
Drepanum falciferum Robertson, 1903. Ent. News 14: 105. 2, d.
Sphecodes pithanus Lovell, 1909. Ent. News 20: 122. °.

Taxonomy: Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 317. d, °.
coronus Mitchell. Minn. to Que. and New England, south to N. C.
Sphecodes coronus Mitchell, 1956. Elisha Mitchell Sei. Soc., Jour. 72: 215. d.
Sphecodes canadensis Mitchell. 1956. Elisha Mitchell. Sci. Soe., Jour. 72: 218. d.
crawfordi Mitchell. N. C. (Bryson City).
Sphecodes crawfordi Mitchell, 1956. Elisha Mitchell, Sci. Soc., Jour. (PR PIE Oe
cressonii (Robertson). Minn. and Ont. to New England, south to Mo. and N. C:
Sphecodium cressonii Robertson, 1903. Ent. News 14: 106. ?, d.
Sphecodes nephelotus Lovell and Cockerell, 1907. Psyche 14: 106. d.
Sphecodes nubilus Lovell, 1909. Ent. News 20: 124, 417. Error for nephelotus.
Sphecodes heterurus Lovell, 1911. Ent. News 22: 212. 2.
davisii Robertson. Minn. and Ont. to N. Y. and New England. Predator: Philanthus solivagus
Say.
Sphecodes davisii Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 319. 3.
Taxonomy: Graenicher, 1910. Canad. Ent. 42: 159. d, °.
dichrous Smith. Minn. to N. S., south to Mo. and Ga. Host: Halictus rubicundus (Chr.).
Predator: Philanthus gibbosus (Fabr.).
Sphecodes dichroa Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 38. 2 (not male).
Sphecodes arvensis Patton, 1880. Amer. Ent. 1: 230. ?, d.

1976 Hymenoptera in America North of Mexico

2Halictus scabrosus Provancher, 1882. Nat. Canad. 13: 200. 6.

Sphecodes lautus Lovell and Cockerell, 1907. Psyche 14: 103. 2.

Sphecodes (Sphecodes) macfarlandi Viereck, 1909. Ent. News 20: 292. °.
eustictus Cockerell. Wash., Colo.

Sphecodes eustictus Cockerell, 1906. Canad. Ent. 38: 162. 2.
exaltus Mitchell. N. C. (Southern Pines).

Sphecodes exaltus Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 217. °.
fattigi Mitchell. Fla. (Gainesville).

Sphecodes fattigi Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 220. d.
fortior Cockerell. N. Mex., Ariz., Calif.

Sphecodes fortior Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 44. 9. Republished by

Cockerell, 1898. N. Mex. Univ., Bul. 1: 44. female.
Biology: Newberry, 1899. Psyche 9: 94.

fragariae Cockerell. Colo., N. Mex. Host: Perdita nuda Ckll.?

Sphecodes fragariae Cockerell, 1903. In Viereck, Amer. Ent. Soc., Trans. 29: 99.
galerus Lovell and Cockerell. N. H. and Vt. to Va.

Sphecodes galerus Lovell and Cockerell, 1907. Psyche 14: 106. 2.
heraclei heraclei Robertson. Ill. to New England south to Ala. and Fla.

Sphecodes heraclei Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 318. °.
heraclei ignitus Cockerell. Ala., Fla.

Sphecodes ignitus Cockerell, 1922. U. S. Natl. Mus., Proc. 60 (18): 13. d.

hesperellus Cockerell. Wash., Oreg., Calif.

Sphecodes (Sphecodes) hesperellus Cockerell, 1904. In Viereck, Canad. Ent. 36: 282. °.
hudsoni Cockerell. Hudson’s Bay, Canada.

Sphecodes hudsoni Cockerell, 1913. Canad. Ent. 45: 12. @.
hydrangeae Mitchell. Conn. to N. C.

Sphecodes hydrangeae Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 214. d.
illinoensis (Robertson). Minn. and Ont. to New England, south to N. C.

Machaeris illinoensis Robertson, 1903. Ent. News 14: 107. °.
johnsonii Lovell. Que. and N. B., south to N. C.

Sphecodes johnsonii Lovell, 1909. Ent. News 20: 122. °.
kincaidii Cockerell. Wash.

Sphecodes kincaidii Cockerell, 1898. Acad. Nat. Sci. Phila., Proc. 50: 56. ?.
knetschi Cockerell. Ill. to New England, south to N. C.

Sphecodes knetschi Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 186. 2, d.
lautipennis Cockerell. Sask., N. D., Mont., Wash., Colo., Calif.

Sphecodes lautipennis Cockerell, 1908. Entomologist 41: 60. d.
levis Lovell and Cockerell. Minn. and Ont. to Maine, south to Tenn. and N. C.

Sphecodes levis Lovell and Cockerell, 1907. Psyche 14: 105. 2, d.
mandibularis Cresson. Colo., Kans., N. Mex., Calif.

Sphecodes mandibularis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 250. 9.
manni Cockerell. Wash.

Sphecodes manni Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 68. °.
millsi Cockerell. Colo.

Sphecodes millsi Cockerell, 1919. Ent. News 30: 288. d.
minor Robertson. Alta. to Maine, south to N. C. and Miss.

Sphecodes minor Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 45. ?.
nigricans miguelensis Timberlake. Calif. (San Miguel Island).

Sphecodes nigricans miguelensis Timberlake, 1940. South. Calif. Acad. Sci., Bul. 39: 193. ¢.
nigricans nigricans Timberlake. Calif.

Sphecodes nigricans Timberlake, 1940. South. Calif. Acad. Sci., Bul. 39: 192. 6.

nigricorpus Mitchell. Conn. (Colebrook).
Sphecodes nigricorpus Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 220. d.

Superfamily APOIDEA 1977

nitidissimus Cockerell. Colo.

Sphecodes (Sphecodium) nitidissimus Cockerell, 1910. Canad. Ent. 42: 367. d.
olympicus Cockerell. Wash.

Sphecodes (Drepanium) Olympicus Cockerell, 1904. In Viereck, Canad. Ent. 36: 230. °.
paraplesius Lovell. R. I.

Sphecodes paraplesius Lovell, 1911. Ent. News 22: 212. °.
patruelis Cockerell. Wash. Predator: Philanthus crabroniformis Sm.

Sphecodes patruelis Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 61. 92, 6.
pecosensis pecosensis Cockerell. Colo., N. Mex.

Sphecodes pecosensis Cockerell, 1904. Entomologist 37: 5. 2.

Proteraner leptanthi Cockerell, 1904. Entomologist 37: 232. d.

Taxonomy: Cockerell, 1906. Canad. Ent. 38: 165. —Cockerell, 1909. Canad. Ent. 41: 219. 9, d.
—Timberlake, 1951. In Muesebeck, Krombein and Townes, U.S. Dept. Agr., Agr. Monog.
2: 1122 (synonymy).
pecosensis salicis Cockerell. Calif.
Sphecodes pecosensis salicis Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 547. °.
perlustrans Cockerell. N. Mex.
Sphecodes pertustrans Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 45. °.
Republished by Cockerell, 1898. N. Mex. Univ., Bul. 1: 45. female.
persimilis Lovell and Cockerell. Mich. to N. Y., R. I., Maine.
Sphecodes persimilis Lovell and Cockerell, 1907. Psyche 14: 103. °.
pimpinellae Robertson. Ill. to Kans. south to Va. and N. C. Host: Augochlora persimilis
(Vier.), A. striata (Prov.). .
Sphecodes pimpinellae Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 51. °.
Sphecodes distolus Lovell, 1909. Ent. News 20: 416. °.

Biology: Ordway, 1964. Kans. Ent. Soc., Jour. 37: 139-152, 5 figs. (parasitic relationships;
probable male).
politulus Cockerell. Sask.
Sphecodes politulus Cockerell, 1937. Amer. Mus. Novitates 909: 3. d.
prosphorus Lovell and Cockerell. Minn. to N. S., south to N. C.
Sphecodes prosphorus Lovell and Cockerell, 1907. Psyche 14: 404. 2, d.
prostygius Mitchell. N. H., Fla.
Sphecodes prostygius Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 499. °.
pulsatillae Cockerell. Colo.
Sphecodes hesperellus pulsatillae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 18: 75. ?.
pusillus Cockerell. Sask.
Sphecodes pusillus Cockerell, 1937. Amer. Mus. Novitates 909: 3. d.
pycnanthemi Robertson. IIl., Colo.
Sphecodes pycnanthemi Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 320. °.
ranunculi Robertson. Minn. to Maine, south to N. C., Kans., ?Ariz.
Sphecodes ranunculi Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 318. ?, d.
rhois (Cockerell). Colo., N. Mex., Ariz.; Mexico.
Proteraner rhois Cockerell, 1904. Entomologist 37: 233. d.
rohweri Cockerell. Colo., N. Mex.
Sphecodes (Machaeris) Rohweri Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 450. 2.
semicoloratus (Cockerell). N. Mex.
Halictus semicoloratus Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 168. °.
shawi Lovell. N. H.
Sphecodes shawi Lovell, 1911. Ent. News 22: 212. °.
smilacinae Robertson. Minn., IIl.
Sphecodes smilacinae Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 321. °.

solidaginis Cockerell. Sask.
Sphecodes solidaginis Cockerell, 1937. Amer. Mus. Novitates 909: 2. d.

1978 Hymenoptera in America North of Mexico

solonis Graenicher. Minn., Ont., and New England.
Sphecodes solonis Graenicher, 1910. Pub. Mus. City Milwaukee, Bul. 1: 229. 6, 9.
Sphecodes morsei Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 213. °.
Sphecodes packardi Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 216. ¢.
sophiae Cockerell. Colo., N. Mex., Ariz., Calif.
Sphecodes sophiae Cockerell, 1898 Denison Univ. Sci. Labs., Bul. 11: 44. 2. Republished by
Cockerell, 1898. N. Mex. Univ., Bul. 1: 44. female.
stygius Robertson. Minn. to Que. and New England, south to Fla.
Sphecodes stygius Robertson, 1893. Amer. Ent. Soc., Trans. 20: 145. 2, 6.

Taxonomy: Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 317. 3, 2.
sulcatulus Cockerell. Colo. Predator: Philanthus crabroniformis Sm.
Sphecodes sulcatulus Cockerell. 1906. Amer. Mus. Nat. Hist., Bul. 22: 426. °.
texana Cresson. Tex.
Sphecodes texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 249. 2, 6.
townesi Mitchell. Mich. and N. Y. to N. C.
Sphecodes townesi Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 212. 2.
trentonensis Cockerell. N. Y.
Sphecodes trentonensis Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11:.62. 2.
veganus Cockerell. N. Mex.
Sphecodes veganus Cockerell, 1904. Entomologist 37: 5. @.
washingtoni Cockerell. Wash.
Sphecodes (Machaeris) Washingtoni Cockerell, 1904. In Viereck, Canad. Ent. 36: 231. @.
wheeleri Mitchell. N. Y. and Conn., south to N. C., ?Kans. Probable male of pimpinellae Robt.
Sphecodes wheeleri Mitchell, 1956. Elisha Mitchell Sci. Soc., Jour. 72: 210. 2, d.

UNPLACED TAXA OF TRIBE HALICTINI
The following species cannot be placed as to genus from existing descriptions.

Halictus arctous Vachal, 1904. Soe. Hist. Arch. Correze, Bul. 26: 480. 3. B. C.

Parasphecodes Californica Provancher, 1896. Nat. Canad. 23: 8. 2. Calif.

Halictus crassicornis Kirby, 1837. In Richardson, Fauna Bor.-Amer., v. 4, p. 267. 2. Canada.

Halictus distinguendus Dalla Torre, 1896. Cat. Hym., v. 10, p. 60. N. name for Halictus
distinctus Provancher, 1882. Nat. Canad. 13: 200. male. ?Que. Preocc.

Family MELITTIDAE

With the exception of the subfamily Ctenoplectrinae which is restricted to the Ethiopian,
Oriental and Australian regions, this family is represented in North America by three subfami-
lies also present in the Old World. None of these is known to occur in the Neotropical Region
and only a few species of the Dasypodinae (Genus Hesperapis) are known to be present in
northern Mexico. The generic relationships within this latter subfamily are in need of clarifica-
tion especially those centering about the genus Hesperapis. On the basis of accumulated in-
trafloral information it appears that many, if not most, members of this family have evolved as
oligoleges of both annual and perennial plants.

Taxonomy: Rozen and McGinley, 1974. Amer. Mus. Novitates 2545: 1-31, figs. 1-82, tables 1, 2
(larvae).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 32-33, fig. 13 (Larrea visiting
spp.).

SUBFAMILY MELITTINAE
Genus MELITTA Kirby
Genus MELITTA Subgenus MELITTA Kirby

Melitta Kirby, 1802. Monographia Apum Angliae, v. 1, p. 117.
Type-species: Melitta tricincta Kirby. Desig. by Richards, 1935.
Cilissa Leach, 1812. In Brewster’s Edinburgh Encycel., v. 9, p. 154.

Superfamily APOIDEA 1979

Type-species: Andrena haemorrhoidalis Fabricius. Desig. by Westwood, 1840.

Kirbya Lepeletier, 1841. Hist. Nat. Ins. Hym., v. 2, p. 145. Preoce.
Type-species: Melitta tricincta Kirby. Desig. by Sandhouse, 1943.

Melitta subg. Brachycephalapis Viereck, 1909. Ent. Soc. Wash., Proc. 11: 47.
Type-species: Melitta californica Viereck. Monotypic and orig. desig.

americana (Smith). Maine to Ga., Miss. Pollen: Unknown, but visits flowers of Polycodiwm and
Rubus.
Cilissa americana Smith, 1853 Cat. Hym. Brit. Mus., v. 1, p. 128. 9.
Taxonomy: Cockerell, 1906. Psyche 13: 5. 2.

americaniformis Viereck. N. J.
Melitta americaniformis Viereck, 1909. Ent. Soc. Wash., Proc. 11: 50. 2.

wilmattae Cockerell. Ariz., Calif. Pollen: Unknown, but visits flowers of Coreopsis and Larrea.

Melitta wilmattae Cockerell, 1937. Amer. Mus. Novitates 948: 3. 9.
Genus MELITTA Subgenus DOLICHOCHILE Viereck

Dolichochile Viereck, 1909. Ent. Soc. Wash., Proce. 11: 49.
Type-species: Dolichochile melittoides Viereck. Monotypic and orig. desig.
melittoides (Viereck). Conn. to Ga., Tenn. Pollen: Unknown, but visits flowers of Polycodium,
Xolisma and Zenobia.
Dolichochile melittoides Viereck, 1909. Ent. Soc. Wash., Proc. 11: 49. 9.

Taxonomy: Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 692.
SuBFAMILY DASYPODINAE
Genus HESPERAPIS Cockerell
Revision: Cockerell, 1916. Psyche 23: 176-177.
Genus HESPERAPIS Subgenus HESPERAPIS Cockerell

Hesperapis Cockerell, 1898. Canad. Ent. 30: 147.

Type-species: Hesperapis elegantula Cockerell. Monotypic.
Zacesta Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 73.

Type-species: Zacesta rufipes Ashmead. Monotypic and orig. desig.

arida Michener. Ariz., south. Calif., desert. Pollen: Oligolectic on flowers of Larrea tridentata.
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 32-33 (floral relationships).
Hesperapis arida Michener, 1936. Ent. News 47: 183. 6, °.

elegantula Cockerell. N. Mex.
Hesperapis elegantula Cockerell, 1898. Canad. Ent. 30: 148. °.

rufipes (Ashmead). South. Calif. Pollen: Oligolectic on flowers of Hugelia.
Zacesta rufipes Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 73. d.
Taxonomy: Michener, 1936. Ent. News 47: 182. d, 2. —Michener, 1953. Kans. Univ. Sci. Bul.
35: 1040, figs. 281-287 (larva).

Genus HESPERAPIS Subgenus AMBLYAPIS Cockerell

Amblyapis Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 362.
Type-species: Halictoides ilicifoliae Cockerell. Monotypic and orig. desig.

ilicifoliae (Cockerell). Calif. Pollen: Oligolectic on flowers of Adenostoma.
Halictoides ilicifoliae Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 361. d.

Genus HESPERAPIS Subgenus PANURGOMIA Viereck

Panurgomia Viereck, 1909. Ent. Soc. Wash., Proc. 11: 48.
Type-species: Panurgomia fuchsi Viereck. Monotypic.

1980 Hymenoptera in America North of Mexico

arenicola Crawford. Ariz., Nev., Calif. deserts; Mexico. Pollen: Oligolectic on spring flowering
Compositae including but apparently not limited to Dyssodia, Encelia, Geraea,
Helianthus, and Rafinesquia.
Hesperapis arenicola Crawford, 1917. Ent. Soc. Wash., Proc. 19: 167. ¢.

carinata carinata Stevens. N. Dak., Idaho.
Hesperapis carinata Stevens, 1919. Canad. Ent. 51: 209. ¢.

Taxonomy: Stevens, 1921. Canad. Ent. 53: 65. 2.
carinata rodecki Cockerell. Colo.
Hesperapis carinata rodecki Cockerell, 1934. Canad. Ent. 66: 153. °.
eumarpha (Cockerell). Calif.
Parandrena eumarpha Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 187. d.
Hesperapis euwmorpha Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 70. Emend.

fuchsi (Viereck). Ariz.
Panurgomia fuchsi Viereck, 1909. Ent. Soc. Wash., Proc. 11: 49. 2.
Taxonomy: Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 671.
fulvipes Crawford. Calif., deserts. Pollen: Oligolectiec on flowers of Geraea canescens.
Hesperapis fulvipes Crawford, 1917. Ent. Soc. Wash., Proc. 19: 166. ¢.
Biology: Hurd, 1957. Kans. Ent. Soc., Jour. 30: 10 (seasonal synchronization with pollen
f source).
larreae Cockerell. Tex. to Utah and South. Calif. deserts; Mex. Pollen: Oligolectic on flowers of
Larrea tridentata.
Hesperapis larreae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 535. ¢.
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 33, fig. 13 (floral relationships).
laticeps Crawford. Calif. deserts and adjacent Nev. and Ariz. Pollen: Oligolectic on flowers of
Mentzelia.
Hesperapis laticeps Crawford, 1917. Ent. Soc. Wash., Proc. 19: 166. d.
leucura Cockerell. Calif.; Mexico (Baja California). Pollen: Possibly oligolectic on flowers of
Dalea, but is known to visit flowers of Cryptantha.
Hesperapis leucurus Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 44. d.
nitidula Cockerell. Calif.; Mexico (Baja California). Pollen: Oligolectic on flowers of Oenothera.

Hesperapis nitidulus Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 45. d.

Taxonomy: Cockerell, 1941. San Diego Soc. Nat. Hist., Trans. 9: 348. °.
oliviae (Cockerell). N. Mex.
Panurgus oliviae Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 149. 6.
parva Michener. Calif. deserts; Mexico. Pollen: Unknown, but visits flowers of Acacia,
Cryptantha, Eriogonum, Eschscholzia, Oenothera, and Phacelia.
Hesperapis parva Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 327. 6, 2.
pellucida Cockerell. Calif. Pollen: Oligolectic on flowers of Eschscholzia californica.
Hesperapis pellucidus Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 198. d.
regularis (Cresson). Calif. Pollen: Oligolectic on flowers of Clarkia.
Panurgus regularis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 62. 6.

Biology: Burdick and Torchio, 1959. Kans. Ent. Soc., Jour. 32: 83-87, 7 figs. —MacSwain,
Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 16-20 (floral relationships).
rhodocerata (Cockerell). N. Mex., Calif.
Panurgus rhodoceratus Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 148. 2, ¢.
semirudis Cockerell. Calif. Pollen: Appears to be oligolectic on vernal flowering Compositae
including Chaenactis, Coreopsis, Layia, and Malacothrix.
Hesperapis semirudis Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 24. 9.
wilmattae Cockerell. Calif., Ariz., Nev. deserts. Pollen: Chaenactis, Encelia, Malacothriz,
Oenothera, Rafinesquia, Sphaeralcea and Tetradymia, but also visits a wide variety of

Superfamily APOIDEA 1981

other flowers including Abronia, Astragalus, Baileya, Calycoseris, Cryptantha,
Eriophyllum, Eschscholzia, Geraea, Malva, Nama, and Phacelia for nectar.
Hesperapis wilmattae Cockerell, 1933. Pan-Pacifie Ent. 9: 26. 9.

Taxonomy: Michener, 1936. Ent. News 47: 184. 6.
SUBFAMILY MACROPIDINAE

Genus MACROPIS Panzer
Revision: Michener, 1938. Psyche 45: 133-135.

Genus MACROPIS Subgenus MACROPIS Panzer

Megilla Fabricius, 1804. Systema Piezatorum, p. 328.
Type-species: Megilla labiata Fabricius. Desig. by Westwood, 1840.
Macropis Panzer, 1809. Faunae Ins. German., h. 107, No. 16.
Type-species: Megilla labiata Fabricius. Monotypic.

This is the only subgenus present in North America, and all of the species are believed to be
oligolectic on flowers of Steironema.

ciliata Patton. Wis. to Que. and Maine south to Ga. Pollen: Unknown, but visits flowers of
Apocynum, Houstonia, and Hydrangea.
Macropis ciliata Patton, 1880. Ent. Monthly Mag. 17: 31. 9.
clypeata Swenk. Nebr.
Macropis clypeata Swenk, 1907. Ent. News 18: 293. 6.
longilingua Provancher. Que.
Macropis longilingua Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 424. 2.
nuda (Provancher). Colo., Mont., and south. Canada to N. Y., New England States and N. S.
Pollen: Unknown, but has been taken at flowers of Apocynum, Geranium, Rubus, and
Vaccinium.
Eucera nuda Provancher, 1882. Nat. Canad. 138: 174. 9.
Macropis (Macropis) morsei Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 388. ¢, 2.

Taxonomy: Mitchell, 1960. N. C. Agr. Expt. Sta. Tech. Bul. 141: 527 (tax. status, synonymy).
patellata Patton. Vt. to N. C., west to Nebr., and Mo. Pollen: Oligolectic on flowers of
Steironema.
Macropis patellata Patton, 1880. Ent. Monthly Mag. 17: 33. d.
steironematis opaca Michener. Wash.
Macropis steironematis opaca Michener, 1938. Psyche 45: 134. 6.
steironematis steironematis Robertson. Iowa and Mo. to Va., N. C., and Ga. Pollen:
Presumably oligolectic on flowers of Steironema, but has been taken at flowers of
Apocynum, Ceanothus, Melilotus, and Seriocarpus.
Macropis steironematis Robertson, 1891. Amer. Ent. Soc., Trans. 18: 68. 2, d.

Family MEGACHILIDAE

This is one of the largest families of bees and is represented by about an equal number of spe-
cies in each of the six zoogeographic regions. It is morphologically the most uniform and discrete
familial assemblage within the Apoidea. While many of the characteristics of this family are
shared with other families of bees and to a lesser extent with the Sphecoidea, the Megachilidae
also possess many characteristics in common with the Scolioidea.

The family is composed of three subfamilies, the Fideliinae and Lithurginae which are all pol-
len-collecting bees, and the Megachilinae which are predominantly pollen-collecting species, but
which contain several genera that are cleptoparasitic mainly in the nests of other megachilids.
The Fideliinae are found only in Chile and South Africa, but both of the other subfamilies are
nearly cosmopolitan and are well represented in America north of Mexico. Most of the pol-
len-collecting species do not make their own burrows, but appropriate a wide variety of
pre-existing burrows, holes, shells and other cavities or construct their nests either in the open

1982 Hymenoptera in America North of Mexico

attaching them to branches and so forth, or place them under stones, cow chips and so on. As a
consequence of these habits, many species readily accept artificial nesting devices, and this has
not only permitted detailed studies of their biology, but has also made possible the manipulation
and management of several species for use in the pollination of agricultural crops. However,
some species in certain genera (e.g., Megachile) and even some groups of species (e.g.,
Heteranthidium and Trachusa) do construct their own burrows. All of the pollen-collecting
Megachilidae use foreign materials in the construction of the cell walls. These materials include
leaves, plant down, leaf pulp, petals, resin, pebbles, mud, clay, and the like. The larvae spin tough
cocoons before pupation.

Apart from the cleptoparasitic species, many of the megachilids are highly restricted in their
intrafloral relationships and thus oligolecty is a relatively common phenomenon in this family.

Revision: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 5-232, 1 frontis., figs. 1-66,
tables 1-7 (eastern U. S. spp.).

Taxonomy: Rozen, 1973. Amer. Mus. Novitates 2527: 1-14, 22 figs. (immature stages).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 264-344, figs. (life histories, nests,
associates). —Eickwort, 1975. Evolution 29: 142-150, 1 fig., 2 tables (gregarious nesting,
and evolution of parasitism and sociality among megachilid bees).

Morphology: Pasteels and Pasteels, 1972. Acad. Roy. Belgique, Mem. Classe Sci. 18: 1-28, 3
pls. (cuticular hairs). —Pasteels and Pasteels, 1973. Tissue and Cell 5: 63-82
(morphogenetic fields of hairs on legs). —Pasteels and Pasteels 1974. Tissue and Cell 6:
65-83 (Stereoscan studies of abdominal scopae).

SUBFAMILY LITHURGINAE
Taxonomy: Rozen, 1973. Amer. Mus. Novitates 2527: 1-14, 22 figs. (immature stages).
Genus LITHURGE Latreille
Genus LITHURGE Subgenus LITHURGE Latreille

Lithurge Latreille, 1825. Fam. Nat. Regne Anim., p. 463.
Type-species: Andrena cornuta Fabricius. Monotypic.
Lithurgus Berthold, 1827. In Latreille, Nat. Fam. Thierr., p. 467. Emend.

Found in the Old World.
Genus LITHURGE Subgenus LITHURGOPSIS Fox

Lithurgopsis Fox, 1902. Ent. News 13: 138.
Type-species: Lithurgus apicalis Cresson. Orig. desig.

Revision: Mitchell, 1938. Psyche 45: 146-155. (Nearctic spp.).

apicalis apicalis (Cresson). S. Dak., Nebr., Kans., Tex., Wyo., Utah, Colo., N. Mex., Ariz., Nev.,
east. Calif. principally from Transit. Zone. Ecology: Nests have been found in holes of
an old, barkless, cottonwood log in Utah; it also is reported to construct its nests in the
doorways of homes in Arizona. Parasite: Anthrax cintalapa Cole. Pollen: Oligolectic,
stores pollen of Opuntia, but visits other flowers presumably for nectar including
Echinocactus, Encelia californica, Eriogonum, Sphaeralcea.

Lithurgus apicalis Cresson, 1875. In Wheeler, Rpt. Geog. Geol. Expl. and Surv. west of

100th Meridian, v. 5, p. 724. °.

Taxonomy: Rozen, 1973. Amer. Mus. Novitates 2527: 6-8, figs. 13-18 (larva).

Biology: Parker and Potter, 1973. Pan-Pacific Ent. 49: 294-299, 19 figs. (nest, larval habits,
parasite).
apicalis littoralis (Cockerell). South Tex. (Point Isabel; near Brownsville).
Lithurgus apicalis littoralis Cockerell, 1917. In W. P. Cockerell, N. Y. Ent. Soc. Jour. 25:
191 Gr

Superfamily APOIDEA 1983

apicalis opuntiae (Cockerell). Colo., N. Mex., Tex., Ariz., Calif.; north. Mexico, principally from
Sonoran Zone. Pollen: Oligolectic, presumably collects pollen from flowers of Opuntia
including O. echinocarpa, O. megacarpa, O. vaseyi, but visits other flowers for nectar.
Lithurgus apicalis var. opuntiae Cockerell, 1902. Ent. News 13: 182.
arizonensis (Cockerell). Ariz.
Lithurgus arizonensis Cockerell, 1937. South. Calif. Acad. Sci., Bul. 36: 108. °.
bruesi (Mitchell). Ill. to Tex.; Mexico.
Lithurgus bruesi Mitchell, 1927. Psyche 34: 104. ¢.
echinocacti (Cockerell). N. Mex., Ariz.; Mexico (Baja California and Sonora). Ecology: Nests in
rotten wood. Pollen: Presumably oligolectic, visits flowers of Echinocactus.
Lithurgus echinocacti Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 453. °.
gibbosus (Smith). N. C., to Fla., Tex. Pollen: Oligolectic, collects pollen from flowers of
Opuntia, but visits other flowers, presumably for nectar including Cirsium, Helianthus,
Ilex, Pontederia, Rudbeckia.
Lithurgus gibbosus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. M7. °.
Lithurgus compressus Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 147. ¢.

SUBFAMILY MEGACHILINAE

This is a large group of mostly pollen-collecting species which are present on all the continents
and many of the islands. The subfamily is composed of two tribes, the Anthidiini and the
Megachilini, both of which are well represented by numerous species in America north of Mex-
ico.

TRIBE ANTHIDIINI

Although this tribe is present throughout much of the world, it contains fewer species than
the Megachilini and, unlike that tribe, is represented by only a single species in Australia. Like
the Megachilini the pollen-collecting species use a wide variety of foreign materials in construct-
ing their nest cells, but are not known to use mud. While the majority of the Anthidiini are soli-
tary, some of the pollen-collecting species in certain genera (e. g., Heteranthidiwm and
Trachusa) live in colonies.

Revision: Michener, 1948. Amer. Mus. Novitates 1381: 1-29 (generic classification).

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 1-118, 229 figs., 41 maps
(Calif. spp.). —Pasteels, 1969. Soc. Roy. Ent. Belgique, Mem. 31: 1-148, 199 figs.
(classification). —Pasteels, 1972. Soc. Roy. Ent. Belgique, Bul. Ann. 108: 72-128, 107 figs.
(classification).

Biology: Custer and Hicks, 1927. Biol. Bul. 52: 258-277 (nesting habits).
Genus TRACHUSA Panzer

Trachusa Panzer, 1804. Faunae Ins. German., pt. 86, nos. 14-15.
Type-species: Apis byssina Panzer. Desig. by Sandhouse, 1943. (=Trachusa
serratulae Panzer).
Diphysis Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 307.
Type-species: Apis byssina Panzer. Monotypic. (=Diphysis pyrenaica Lepeletier).
Trachusomimus Popov, 1964. Rev. Ent. URSS 43: 406.
Type-species: Trachusa perdita Cockerell. Orig. desig.

The genus also occurs in the Palaearctic Region.
Revision: Michener, 1941. Pan-Pacific Ent. 17: 119-125. (Nearctie spp.).

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 5: 1041 (larva). —Thorp, 1963. Pan-Pacific
Ent. 39: 56-58 (Key to spp. of Trachusa). —Thorp, 1966. Kans. Ent. Soce., Jour. 39: 182
(generic status of Trachusomimus Popov). —Grigarick and Stange, 1968. Calif. Ins.
Survey, Bul. 9: 4-6, figs. 1, 129-134, map 1 (Calif. spp.).

gummifera Thorp. Calif. (Marin and San Francisco Counties).
Trachusa gummifera Thorp, 1963. Pan-Pacific Ent. 39: 56. 2, d.

1984 Hymenoptera in America North of Mexico

manni Crawford. South Ariz., north. Mexico. Parasite: Heterostelis manni Cwfd.?
Trachusa manni Crawford, 1917. Ent. Soe. Wash., Proc. 19: 167. 6.
perdita Cockerell. Calif. Parasite: Heterostelis hurdi Thorp, Nemognatha scutellaris LeC.
Pollen: Presumably polylectic, visits flowers of Brodiaea lutea, Clarkia cylindrica, C.
purpurea quadrivulnera, C. speciosa speciosa, Collinsia heterophylla, Convolvulus,
Eriodictyon californicum, Eriophyllum confertiflorum, Layia platyglossa campestris,
Lotus scoparius, Lupinus nanus, Penstemon antirrhinoides, P. heterophyllus, Salvia

columbariae.
Trachusa perdita Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 159. 6.

Biology: Michener, 1941. Pan-Pacific Ent. 17: 123 (nest). —Thorp, 1966. Kans. Ent. Soc., Jour.
39: 132 (parasite). —MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 49

(floral relationships).
Genus HETERANTHIDIUM Cockerell

Heteranthidium Cockerell, 1904, Ent. News 15: 292.
Type-species: Anthidium dorsale Lepeletier. Monotypie and orig. desig.

Taxonomy: Schwarz, 1926. Amer. Mus. Novitates 218: 1-15 (U.S. spp.). —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 23-31, figs. 8-10, table 1 (eastern U.S. spp.). —Snelling,
1966, Los Angeles County Mus. Contrib. Sci. 97: 1-8, 1 fig. (Key to U.S. spp.). —Grigarick

and Stange, 1968. Calif. Ins. Survey, Bul. 9: 6-10, figs. 111-128, maps 2-5 (Calif. spp.).

autumnale Snelling. Calif. (Desert Hot Springs); Mexico (Baja California). Pollen: Unknown,

but presumably visits flowers of Chrysothamnus or Haplopappus.
Heteranthidium autumnale Snelling, 1966. Los Angeles County Mus. Contrib. Sci. 97: 1,

figs ld, (2
Taxonomy: Snelling, 1975. Ent. Soc. Wash., Proc. 77: 90 (color variation in Baja Calif.

specimen).

bequaerti Schwarz. South. Calif. (Colorado Desert). Pollen: Oligolege of Dalea spinosa.
Heteranthidium bequaerti Schwarz, 1926. Amer. Mus. Novitates 218: 6. °.

cordaticeps Michener. Tex., N. Mex. Ariz. Pollen: Unknown, but visits flowers of Helianthus

annuus, also possibly Heterotheca subaxillaris.
Heteranthidium cordaticeps Michener, 1949. Kans. Ent. Soc., Jour. 22: 41. 9.

Taxonomy: Snelling, 1975. Ent. Soe. Wash., Proc. 77: 87-88, fig. 1 (male, geogr. and floral

records).
crassipes (Cresson). N. C. to Fla.

Anthidiwm crassipes Cresson, 1878. Amer. Ent. Soc., Trans. 7: 112. °.
dorsale (Lepeletier). N. J., N. C., Ga., Ala. Pollen: Unknown, but visits flowers of

Strophostyles.
Anthidium dorsale Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 384. d.
Anthidium harbecki Crawford, 1910. Ent. News 21: 456. °.

Taxonomy: Cresson, 1864. Ent. Soc. Phila., Proc. 2: 381. d.
fontemvitae Schwarz. N. C. to Fla. Pollen: Unknown, but visits flowers of Chrysopsis,

Kuhnistera.
Heteranthidium fontemvitae Schwarz, 1926. Amer. Mus. Novitates 218: 8. d, 9.
larreae (Cockerell). West. Tex. to south. Utah, Nev. and south. Calif. (Sonoran and Mojave
deserts); Mexico. Pollen: Oligolege of Larrea tridentata, but visits other flowers for
nectar including Baileya, Machaeranthera tortifolia, Salvia dorri, Senecio douglasii.
Anthidium larreae Cockerell, 1897. Canad. Ent. 29: 220. 2, do.
Biology: MacSwain, 1946. Pan-Pacific Ent. 22: 159. (nesting habits). —Hurd and Linsley,
1975. Smithsn. Contrib. Zool. 193: 34, fig. 14 (intrafloral relationships).

occidentale (Cresson). Colo., N. Mex.; Mexico (Puebla).
Anthidium occidentale Cresson, 1868. Amer. Ent. Soc., Trans. 1: 386. 2, 6.
ridingsii (Cresson). N. C. to Fla. and Tex. Pollen: Unknown, but visits flowers of Crotalaria,

Galactia, Hypericum, Melilotus, Rhus.
Anthidium ridingsii Cresson, 1878. Amer. Ent. Soc., Trans. 7: Proce. iii. °.

Superfamily APOIDEA 1985

timberlakei Schwarz. South. Oreg., Calif., west. Nev. Pollen: Polylectic, visits flowers of
Chaenactis douglasii, C. glabriuscula, Cirsium, Clarkia biloba, C. concinna, C.
cylindrica, C. gracilis, Coreopsis lanceolata, Encelia farinosa, Helianthus gracilentus,
Salvia mellifera.
Heteranthidium timberlakei Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 409. d, @.
Heteranthidium subtimberlakei Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 414. 9.

Taxonomy: Stephen and Torchio, 1961. Pan-Pacifie Ent. 37: 41-43 (as zebratum). —Snelling,
1966. Los Angeles County Mus. Contrib. Sci. 97: 5 (synonomy).

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 49 (floral
relationships).
zebratum (Cresson). Mich. south to Miss., west to S. Dak., Nebr., Colo. and N. Mex. Pollen:
Unknown, but visits flowers of Gaillardia, Helianthus.
Anthidium zebratum Cresson, 1872. Amer. Ent. Soc., Trans. 4: 270. d.
Protanthidiwm cockerelli Titus, 1902. Ent. News 13: 170. 6.
Anthidium (Protanthidium) Chippewaense Graenicher, 1910. Canad. Ent. 42: 157. d, 2.
Heteranthidium zebratum mississippi Michener, 1937. Amer. Midland Nat. 38: 449. 6, 9.

Taxonomy: Snelling, 1966. Los Angeles County Mus. Contrib. Sci. 97: 4-5 (synonymy).
Genus PARANTHIDIUM Cockerell and Cockerell

Genus PARANTHIDIUM Subgenus PARANTHIDIUM Cockerell and Cockerell

Anthidium subg. Paranthidium Cockerell and Cockerell, 1901. Ann. and Mag. Nat. Hist.
(7) 7: 50.
Type-species: Paranthidium jugatorium perpictum (Cockerell). Orig. desig.
(=Anthidium perpictum Cockerell).

Taxonomy: Schwarz, 1926. Amer. Mus. Novitates 226: 19-25 (N. Amer. spp.). —Moure, 1965.
Ent. Soc. Wash., Proce. 67: 29-31 (generic limits).
jugatorium butleri Snelling. Ariz. Pollen: Unknown, but visits flowers of Erigeron,
Helianthus.
Paranthidium jugatorium butleri Snelling, 1962. Pan-Pacifie Ent. 38: 225. 2, ¢.
jugatorium jugatorium (Say). N. Y., N. J., Ind., Minn., Wis., Mo., Nebr. Parasite: Dasymutilla
asopus bexar (Blake). Pollen: Unknown, but visits flowers of Helianthus strumosus.
Megachile jugatoria Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 352. °.
Stelis obesa Say, 1837. Boston Jour. Nat. Hist. 1: 398. d.

Biology: Michener, 1975. Kansas Ent. Soc., Jour. 48: 194-200 (nest, parasite).
jugatorium lepidum (Cresson). Ky. and Va., south to Ga. Pollen: Unknown, but visits flowers

of Helenium, Helianthus, Rudbeckia, Silphium.
Anthidium lepidum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 115. d.

jugatorium perpictum (Cockerell). Colo., N. Mex., Ariz.
Anthidium perpictum Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 63. 2.
Anthidium (Dianthidiwm) perpictum coloradense Swenk, 1914. Nebr. Univ., Studies 14:
Soar
Taxonomy: Cockerell, 1898. N. Mex. Univ., Bul. 1: 63. 9. —Cockerell, 1900. Ann. and Mag.
Nat. Hist. (7) 5: 413. 6, 2.

Biology: Banks, 1902. N. Y. Ent. Soc., Jour. 10: 213 (nesting habits).

Genus PARANTHIDIUM Subgenus MECANTHIDIUM Michener

Paranthidium subg. Mecanthidium Michener, 1942. N. Y. Ent. Soc., Jour. 50: 278.
Type-species: Paranthidium (Mecanthidium) sonorum Michener. Orig. desig.

Taxonomy: Moure, 1965. Ent. Soc. Wash., Proc. 67: 30 (systematic position).

sonorum Michener. Ariz., northern Mexico.
Paranthidium (Mecanthidium) sonorum Michener, 1942. N. Y. Ent. Soc., Jour. 50: 278. d,
Qe

1986 Hymenoptera in America North of Mexico

Genus ADANTHIDIUM Moure

Adanthidium Moure, 1965. Ent. Soc. Wash., Proc. 67: 29.
Type-species: Anthidium texanum Cresson. Orig. desig.

arizonicum (Rohwer). Ariz.
Dianthidium arizonicum Rohwer, 1916. Ent. Soc. Wash., Proe. 18: 192. 6.

Biology: Middleton, 1916. Ent. Soc. Wash., Proc. 18: 193 (nest).

texanum (Cresson). Kans., Ariz., Tex.
Anthidium texanum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 113. ¢.

Biology: Melander, 1902. Biol. Bul. 3: 27 (nest).
Genus ANTHIDIUM Fabricius

Genus ANTHIDIUM Subgenus ANTHIDIUM Fabricius

Anthidium Fabricius, 1804. Systema piezatorum, p. 364.
Type-species: Apis manicata Linnaeus. Desig. by Latreille, 1810.

Taxonomy: Swenk, 1913. Nebr. Univ., Studies 14: 9-18 (Nebr. spp.). —Cockerell, 1925. Calif.
Acad. Sci., Proc. (4) 14: 345-361 (western U. S. spp.). —Schwarz, 1927. Amer. Mus.
Novitates 252: 1-22; 253: 1-17 (N. Amer. spp.). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech.
Bul. 152: 9-13, figs. 1, 3, table 1 (eastern U. S. spp.). —Grigarick and Stange, 1968. Calif.
Ins. Survey, Bul. 9: 10-35, figs. 3, 7-100, 214, 217-220, maps 6-24 (Calif. spp.).

Biology: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 11 (summary of literature).

atripes Cresson. Tex., Colo., Nev., and Calif. Pollen: Apparently narrowly polylectic, visits
flowers of Astragalus including A. douglasii var. parishii, Lotus including L. strigosus
var. hirtellus.
Anthidium emarginatum var. atripes Cresson, 1879. Amer. Ent. Soc., Trans. 7: 205. 3.
Anthidium polingae Schwarz, 1931. N. Y. Ent. Soe., Jour. 39: 315. 2, 3.

Taxonomy: Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 388. 2. —Schwarz, 1937. N. Y. Ent. Soc.,
Jour. 45: 377.

banningense Cockerell. Wash., Utah, Oreg., Calif. Ecology: Nests in ground and uses

down-like nesting materials from Artemisia and Cirsium. Parasite: Chrysis
florissanticola Roh. Pollen: Apparently oligolectic on Phacelia including P. heterophylla,
P. imbricata, P. leucophylla, P. ramosissima, but also visits other flowers presumably
for nectar including Chaenactis glabriuscula, Clarkia amoena, Cryptantha, Lotus
davidsonii, Medicago, Melilotus, Oenothera, Physocarpus, Trifolium wormskioldii.

Anthidium banningense Cockerell, 1904. South. Calif. Acad. Sci. Proc. 3: 58. 3.

Anthidium plumarium Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 356. 6.

Anthidium longispinum Schwarz, 1927. Amer. Mus. Novitates 253: 6. 2.

Biology: Jaycox, 1967. Kans. Ent. Soc., Jour. 40: 565-570 (territoriality, pollen source).
clypeodentatum Swenk. Man. to B. C., Nebr., Colo., Calif. Pollen: Apparently narrowly
polylectic, presumably obtains pollen from Leguminosae including Astragalus douglasii
var. parishii, Lotus davidsonii, L. oblongiflorus, Lupinus, but also visits other flowers
for nectar including Ceanothus, Cryptantha intermedia, Phacelia frigida.
Anthidium clypeodentatum Swenk, 1914. Nebr. Univ., Studies 14: 12. 2.
Anthidium incurvatum Swenk, 1914. Nebr. Univ., Studies 14: 22. ¢.
Anthidium emarginatum bilineatum Schwarz, 1927. Amer. Mus. Novitates 252: 4. 6.
Anthidium clypeodentatum var. lutzi Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 380. 9, 6.

Taxonomy: Schwarz, 1928. Canad. Ent. 60: 215. —Schwarz, 1937. N. Y. Ent. Soc., Jour. 45:
380 (lutzi). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 15-16, figs. 16-18, 68,
70, 91, map 8 (synonymy).
cockerelli Schwarz. West. Tex. to south. Nev. and south. Calif.; Mexico (Baja California).
Pollen: Polylectic, visits flowers of Abronia villosa, Cercidium floridum, Chaenactis
carphoclinia, Dalea californica, D. emoryi, Encelia, Eschscholzia darwinensis, Geraea

Superfamily APOIDEA 1987

canescens, Larrea tridentata, Lupinus, Malacothrix, Palafoxia linearis, Phacelia
crenulata, P. distans, Trifolium, Oenothera clavaeformis var. auriantiaca.
Anthidium cockerelli Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 386. 6.

Taxonomy: Cockerell, 1937. Amer. Mus. Novitates 948: 6. °.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 35 (floral relationship with
Larrea).
collectum Huard. Calif. (including Channel Islands); Mexico (Baja California). Ecology: Nests
in abandoned burrows in the ground, and uses down from the leaves and along the stem
of Artemisia tridentata. Parasite: Chrysis cognata Bohart, C. coloradica Bohart, C.
tripartita Aaron, Diorys pomonae CkIl., D. productus cismontanicus Hurd,
Monodontomerus montivagus Ashm., Sphaeropthalma blakei (Fox), S. orestes (Fox), S.
unicolor (Cress.). Pollen: Polylectic, especially on Lotus, Phacelia, Cryptantha, but visits
a wide variety of flowers presumably for nectar and or pollen including Amsinckia
intermedia, Astragalus tener, Chaenactis glabriuscula, Cryptantha intermedia,
Dudleya, Encelia farinosa, Eriastrum pluriflorum, Erodium botrys, Gilia, Lotus
argyrophyllus, L. corniculatus, L. davidsonii, L. glaber, L. hamatus, L. scoparius, L.
strigosus var. hirtellus, L. subpinnatus, Lupinus bicolor, L. densiflorus, L. nanus,
Melilotus, Mimulus fremontii, Mirabilis laevis, Nemophila, Oenothera, Phacelia
cicutaria, P. distans, P. hispida, P. imbricata, P. ramosissima, P. tanacetifolia,
Pholisma racemosa, Platystemon californicus, Salix, Salvia columbariae, Sambucus,
Trichostema lanatum, T. parishii, Viguiera laciniata, V. multiflora.
Anthidium compactum Provancher, 1896. Nat. Canad. 23: 9. d. Preoce.
Anthidium collectum Huard, 1896. Nat. Canad. 23: 124. N. name.
Anthidium angelarum Titus, 1906. Ent. Soc. Wash., Proc. 7: 164. °.
Anthidium transversum Swenk, 1914. Nebr. Univ., Studies 14: 19. °.
Anthidium puncticaudum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 360. ¢.
Anthidium collectum bilderbacki Cockerell, 19388. San Diego Soc. Nat. Hist., Trans. 9: 38.
oe
Anthidium catalinense Cockerell, 1939. Calif. Acad. Sci., Proc. (4) 23: 433. d, 9.
Anthidium clementinum Cockerell, 1939. South. Calif. Acad. Sci., Bul. 38: 138. °.

Taxonomy: Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 57, 73. —Titus, 1906. Ent. Soc.
Wash., Proc. 7: 163. —Cockerell, 1941. Sixth Pacific Sci. Cong., Proc., v. 4, p. 289
(clementinum). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 16-19, figs. 19-21,
83, map 10 (synonymy).

Biology: Hicks, 1929. Canad. Ent. 61: 84 (nesting habits, copulation, nest construction,
sleeping sites). —Hurd, 1958. Calif. Univ. Pubs. Ent. 14: 285, 292-293 (parasites).
—Ferguson, 1962. Calif. Univ. Pubs. Ent. 27: 6-22 (parasites, nest).

dammersi Cockerell. Calif. (Colorado and Mojave deserts), Nev. Pollen: Unknown, but visits
flowers of Astragalus lentiginosus var. fremontii, Dalea fremontii, D. f. var. saundersii,
Phacelia distans.
Anthidium dammersi Cockerell, 1937. Amer. Mus. Novitates 948: 6. d, °.
edwardsii Cresson. Wash., Oreg., Calif., Idaho, Utah. Ecology: Nests in dead bamboo. Parasite:
Tricrania stansburyi Hald. Pollen: Polylectic, visits a wide variety of flowers including
Achillea millaefolium, Adenostegia pilosa, Astragalus, Bigelovia, Calochortus luteus,
Cirsium, Clarkia speciosa, Cleomella obtusifolia, Cordylanthus pilosa, C. tenuis, Cressa
cretica, Eriogonum gracile, Eriophyllum, Frankenia grandiflora, Heliotropium
curassavicum, Hemizonia, Lathyrus splendens, Lotus americanus, L. purshianus, L.
scoparius, Marrubium vulgare, Medicago sativa, Melilotus, Monardella lanceolata,
Navarretia heterodoxa, Penstemon heterophyllus, Phacelia distans, P. ramosissima,
Solidago californica, Stephanomeria exigua, Trichostema lanceolatum, T. laxum,
Trifolium obtusifolium, T. repens, T. variegatum, Verbena californica, V. lasiostachys,
Wislizenia refracta.
Anthidium edwardsii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 112. ¢.
Anthidium 3-cuspidum Provancher, 1896. Nat. Canad., 23: 10. d.
Anthidium tricuspidum Cockerell, 1904. South. Calif. Acad. Sci, Bul. 3: 57. Emend.
Anthidium hesperium Swenk, 1914. Nebr. Univ., Studies 14: 18. °.

1988 Hymenoptera in America North of Mexico

Anthidium depressum Schwarz, 1927. Amer. Mus. Novitates 253: 4. ¢.

Taxonomy: Titus 1906. Ent. Soe. Wash. Proc. 7: 163. —Grigarick and Stange. 1968. Calif. Ins.
Survey, Bul. 9: 19-21, figs. 3, 49-51, 79, map. 12 (synonymy).
emarginatum (Say). B. C. to Calif., east to Kans., Nebr. and N. Mex. Ecology: Uses down from
stems of Cirsium undatum for nest construction in Colo. Parasite: Leucospis affinis
Say, Monodontomerus montivagus Ashm., Nemognatha dubia LeC., N. lutea LeC.,
Physocephala texana (Will.). Pollen: Apparently polylectic, visits flowers of Arenaria,
Astragalus bolanderi, Clarkia pulchella, Cryptantha intermedia, Dalea fremontii,
Erigeron foliosus var. stenophyllus, Eriodictyon, Hackelia jessicae, Lotus argyraeus
multicaulis, L. scoparius, Lupinus lyallii, L. superbus, Penstemon, Phacelia californica,
P. superbus, P. frigida, P. heterophylla, P. ramosissima, Symphoricarpos.
Megachile emarginata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 352. 9.
Anthidium atrifrons Cresson, 1868. Amer. Ent. Soc., Trans. 1: 387. @.
Anthidium atriventre Cresson, 1878. Amer. Ent. Soc., Trans. 7: Proe. iii. 2.
Anthidium saxorum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 57. ¢.
Anthidium collectum var. ultrapictum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 73.
3.
Anthidium titusi Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 58. d.
Anthidium bernardinum var. aridum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 58.
3.
Anthidium astragali Swenk, 1914. Nebr. Univ., Studies 14: 16. d, 2.
Anthidium fresnoense Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 347. @.
Anthidium angulatum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 357. ¢.
Anthidium hamatum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 358. d.
Anthidium spinosum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 359. ¢.
Anthidium lucidum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 361. 6.
Anthidium rhodophorum Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 623. °@.
Anthidium sculleni Schwarz, 1930. N. Y. Ent. Soe., Jour. 38: 10. ¢.

Taxonomy: Cresson, 1864. Ent. Soe. Phila., Proc. 2: 374. 2, 6. —Cockerell, 1904. South. Calif.
Acad. Sci., Bul. 3: 72. d (saxorwm). —Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 73.
6 (tituwsi). —Cockerell, 1904. South. Calif. Acad. Sei., Bul. 3: 76. —Schwarz, 1928. N. Y.
Ent. Soc., Jour. 36: 372 (bernardinum var. aridum). —Grigarick and Stange, 1968. Calif.
Ins. Survey, Bul. 9: 21-22, figs. 34-36, 87, 95, map 13 (synonymy).

Biology: Hicks, 1926. Colo. Univ., Studies 15: 47 (collection of nest material).

. jocosum Cresson. Mont., Colo., Calif.; Mexico (Baja California and Sonora). Pollen: Apparently
polylectic, visits flowers of Ceanothus, Chorizanthe staticoides, Cryptantha intermedia,
Eriastrum plurifolium, Ericameria monactis, Eriodictyon, Eriogonum fasciculatum, E.
f. polifolium, E. inflatum, Gutierrezia californica, Haplopappus cooperi, Hemizonia,
Larrea tridentata, Lasthenia, Lotus scoparius, Marrubium vulgare, Oenothera,
Penstemon, Phacelia ramosissima, Salvia carduacea, S. pachyphylla, Trifoliwm repens.

Anthidium jocosum Cresson, 1878 Amer. Ent. Soc., Trans. 7: 111. ¢.
Anthidium xanthognathum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 347. 9.
Anthidiwm fontis Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 348. d, 2.

Taxonomy: Schwarz, 1927. Amer. Mus. Novitates 277: 5. °.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool..193: 35 (floral relationships).
maculifrons Smith. Ill., Tenn. and Va., south to Fla., west to Ariz.; Mexico. Pollen: Unknown,
but visits flowers of Afzelia, Asclepias, Bidens, Coreopsis, Cracca, Crotalaria, Galactia,
Galax, Phaseolus, Psoralea.
Anthidium maculifrons Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 214. 9.
Anthidium cognatum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 109. , d.

Biology: Hungerford and Williams, 1912. Ent. News 23: 256.

maculosum Cresson. S. Dak., Colo., and Tex., west to Oreg. and Calif.; Mexico. Ecology: Nests
in elderberry stems. Parasite: Chrysis tripartita Aaron, Dioxys aurifuscus (Titus),
Leucospis affinis Say. Pollen: Apparently polylectic, visits flowers of Antirrhinum,

Superfamily APOIDEA = 1989

Arctostaphylos, Aster, Astragalus douglasii var. parishii, Chrysopsis villosa, Cirsium
vulgare, Cordylanthus nevinii, Corethrogyne filaginifolia, Eriodictyon trichocalyx,
Grindelia camporum, Haplopappus linearifolius, Helianthus, Hulsea callicarpha, Lotus
argophyllus, L. corniculatus, L. davidsonit, L. humistratus, L. nevadensis, L. scoparius,
L. strigosus var. hirtellus, Phacelia ramosissima, Senecio ionophyllus, Solidago
californica, Stachys albens, S. bullata, S. pycnantha, Trichostema laxum, Verbena
hastata, V. lasiostachys.

Anthidium maculatum Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 216. ¢. Preoce.

Anthidium maculosum Cresson, 1878. Amer. Ent. Soe., Trans. 7: 110. 2.

Anthidium lupinellum Cockerell, 1904. South. Calif. Acad. Sei., Bul. 3: 58. d.

Anthidium americanum Friese, 1911. Das Tierreich, v. 28, p. 395. N. name.

Taxonomy: Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 412. 6. —Cockerell, 1905. Amer.
Ent. Soc., Trans. 31: 336. —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 23-25,
figs. 7, 9, 71, map 15.

Biology: Parker and Bohart, 1966. Pan-Pacific Ent. 42: 95 (nest, parasite). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 269-272, pl. 17, figs. 79, 80 (nest, supersedure, nest
architecture, life history). —Horning, 1971. Ent. Soc. Wash., Proc. 73: 43 (nest, parasite).

manicatum Linnaeus. N. Y. (Ludlowville); Argentina, Brazil, Uruguay and Canary Is.; Europe.
Ecology: Nests readily in a wide variety of movable objects including trap-nests.
Adventive from Europe. Pollen: Polylectic, visits a wide variety of flowers for nectar
and pollen in Europe and South America, but so far has only been taken at the flowers
of Caryopteris clandonensis in N. Y.
Apis manicata Linnaeus, 1758. Syst. Nat., Ed. 10, p. 577.
Apis uncata Schrank 1802. Fauna Boica 2(2): 379.

Taxonomy: Moure and Urban, 1964. II Congr. Latino-Amer. Zool., An. 1: 96-102, figs. 1-5 (S.
Amer. distr.). —Jaycox, 1967. Kans. Ent. Soc., Jour. 40: 124-126 (N. Amer. distr.).

Biology: Green, 1921. Ent. Soe. London, Proc., p. 1xxii-lxxiii (territoriality). —Ward, 1928.
Entomologist 61: 267-272 (territoriality). —Haas, 1960. Ztschr. Tierpsychol. 17: 402-416
(territoriality). —Pechuman, 1967. N. Y. Ent. Soc. 75: 68-73 (behavior).

montivagum Cresson. Colo.
Anthidium montivagum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 110. 2, ¢.
mormonum Cresson. B. C., Idaho, Mont. and Nebr., south to Calif., Ariz. and N. Mex.; Mexico

(Baja California). Ecology: Nests in deserted beetle burrows in old yucca floral scapes
and oak stumps, uses down gathered from hairy leaves and stems of Lepidospartum
squamatum for nest construction. Parasite: Chelynia leucotricha Ckll.? Pollen:
Apparently polylectic, visits flowers of Astragalus, Ceanothus parviflorus,
Chrysothamnus viscidiflorus, Cryptantha intermedia, Eriodictyon, Eriogonum
fasciculatum, Grindelia, Lotus andersoni, L. argophyllus, L. davidsonii, L. nevadensis,
L. oblongifolius, L. scoparius, L. strigosus var. hirtellus, Lupinus lobbii, Penstemon
spectabilis, Phacelia ciliata, P. heterophylla, P. imbricata, P. pringeli, P. ramosissima,
Rubus leucodermis, Stenotopsis linearifolius, Trifolium repens.

Anthidium mormonum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 110. ¢.

Anthidium blanditum Cresson, 1879. Amer. Ent. Soe., Trans. 7: 206. °.

Anthidium pondrewm Titus, 1902. Ent. News 13: 169. ¢.

Anthidium pecosense Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 58. ¢.

Anthidium bernardinum var. wilsoni Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 58.
3.

Anthidium bernardinum var. fragariellum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3:
58. 6.

Anthidium blanditum praedentatum Cockerell, 1907. Entomologist 40: 99. °.

Anthidium poudreum(!) Cockerell, 1911. U. S. Natl. Mus., Proc. 40: 248. d (2 misdet.).

Anthidiuwm wallisi Cockerell, 1913. Canad. Ent. 45: 13. 2.

Anthidium nebrascense Swenk, 1914. Nebr. Univ., Studies 14: 14. d, @.

Anthidium praedentatum trianguliferum Swenk, 1914. Nebr. Univ., Studies 14: 18. 9.

Anthidium flavicaudum Cockerell, 1925. Calif. Acad. Sei., Proc. (4) 14: 359. d.

Anthidium wyomingense Schwarz, 1927. Amer. Mus. Novitates 252: 20. °.

1990 Hymenoptera in America North of Mexico

Anthidium mormonum hicksi Schwarz, 1934. Amer. Mus. Novitates 743: 4. 2.
Anthidium wallisi var. wallowana Schwarz, 1940. Amer. Mus. Novitates 1058: 5. @.

Taxonomy: Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 74. ¢ (pecosense). —Cockerell,
1904. South. Calif. Acad. Sci., Bul. 3: 75, 76. ¢, 2 ? —Cockerell, 1911. U. S. Natl. Mus.,
Proce. 40: 248. ¢, 2 (pecosense). —Schwarz, 1927. Amer. Mus. Novitates 252: 7
(nebrascense). —Schwarz, 1928. Canad. Ent. 60: 213 (nebrascense). —Schwarz, 1930. N. Y.
Ent. Soc., Jour. 38: 9 (nebrascense). —Schwarz, 1940. Amer. Mus. Novitates 1058: 6
(nebrascense). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 25-27, figs 43-45,
80, 92, map 16 (synonymy).

Biology: Hicks, 1929. Ent. News 40: 105 (nesting habits).

pallidiclypeum Jaycox. Calif. Pollen: Unknown, but visits flowers of Ceanothus, Collinsia
bicolor, Delphinium parryi var. seditiossum, Lotus scoparius, Trifolium variegatum.
Anthidium pallidiclypeum Jaycox, 1963. Pan-Pacific Ent. 39: 267, 5 figs. d, 2.
palliventre Cresson. Utah, B. C., Oreg., Calif. Ecology: Excavates nest in sand. Pollen:
Unknown, but visits flowers of Abronia maritima, Cakile edentula, Dudleya, Horkelia,
Lathyrus littoralis, Lotus eriophorus, L. heermannii, Phacelia distans, P. ramosissima.
Anthidium palliventre Cresson, 1878. Amer. Ent. Soc., Trans. 7: 114. 9.
Anthidium pallidiventre Dalla Torre, 1896. Cat. Hym., v. 10, p. 468. Emend.
Anthidium californicum Cresson, 1879. Amer. Ent. Soc., Trans. 7: 206. 6.
Anthidium palliventre vanduzeei Cockerell 1937. Pan-Pacific Ent. 13: 150. 2, ¢.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 28-29, figs. 28-30, 85, map
18 (synonymy).
Biology: Hicks, 1928. Pan-Pacific Ent. 5: 51 (nest excavation).
palmarum Cockerell. West. Tex to south. Calif. and Nev.; Mexico (Baja California). Ecology:
Found in burrow of dead floral scape of Yucca whipplei. Pollen: Possibly oligolectic on
Phacelia including P. affinis, P. cicutaria, P. ciliata, P. crenulata, P. distans, P. hispida,
P. ramosissima, but visits other flowers including Acamptopappus sphaerocephalus,
Amsinckia, Aster tortifolius, Astragalus, Bebbia juncea, Dalea schottii, Encelia
virginiensis actoni, Lotus argophyllus, Lotus scoparius, Oenothera, Senecio douglasii,
Sphaeralcea, Stenotopsis linearifolius.
Anthidium palmarum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 59. ¢.
Anthidium palmarum micheneri Schwarz, 1957. Kans. Ent. Soc., Jour. 30: 132. 6, 9.

Taxonomy: Snelling, 1962. Pan-Pacific Ent. 38: 226-227 (distribution and floral records,
palmarum micheneri). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 29-30, figs.
64-66, 69, 74, map 19 (synonymy).

paroselae Cockerell. Tex. to south. Calif.; Mexico (Sonora). Ecology: Nests in hard sand; two

flight periods occur, one in the spring and one in the autumn. Parasite: Dioxys
productus subruber (Ckll.)? Pollen: Polylectic, visits flowers of Baileya multiradiata,
Bebbia juncea, Croton californicus, Cryptantha barbigera, Dalea scoparia, Helianthus
petiolaris var. canescens, Larrea tridentata, Malacothrix, Melilotus, Palafoxia linearis,
Pulchea sericea, Prosopis.

Anthidium paroselae Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 62. 2.

Anthidium parosetae(!) Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 302.

Taxonomy: Cockerell, 1898. N. Mex. Univ., Bul. 1: 62. 2. —Cockerell, 1900. Ann. and Mag.
Nat. Hist. (7) 5: 412. 2, 6. —Schwarz, 1929 (1928). N. Y. Ent. Soc., Jour. 36: 384. 2, 6.
—Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 30, figs. 10-12, 73, 100, map. 20.

Biology: Newberry, 1900. Psyche 9: 94 (nest, parasite). —Hurd and Linsley, 1975. Smithsn.
Contrib. Zool. 193: 35 (floral relationships).
placitum Cresson. Idaho and Wyo., south to Calif., Ariz. and N. Mex. Pollen: Apparently

polylectic, although current data suggests a predilection for the flowers of Cordylanthus
including C. filifolius, C. nevini, C. rigidus brevibracteatus, but also visits flowers of
Chrysopsis villosa, Cryptantha intermedia, Eriodictyon trichocalyx, Eriogonum
subscaposum, Gutierrezia californica, Monardella, Solidago confinis.

Anthidiuwm placitum Cresson, 1879. Amer. Ent. Soc., Trans. 7: 206. 2.

Anthidium bernardinum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 58. 6.

Superfamily APOIDEA 1991

Anthidium hesperium dentipygum Swenk, 1914. Nebr. Univ., Studies 14: 19. °.

Anthidium permaculatum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 349. 9.

Anthidium bernardinum mesaverdense Schwarz, 1927. Amer. Mus. Novitates 252: 15. 6,
OF

Anthidium niveumtarsum Schwarz, 1927. Amer. Mus. Novitates 252: 18. d, 2.

Taxonomy: Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 74. ¢. —Schwarz, 1928. N. Y. Ent.
Soe., Jour. 36: 378. 2, 3d (permaculatum). —Sechwarz, 1937. N. Y. Ent. Soc.; Jour. 45: 382.
—Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 30-32, figs. 46-48, 78, 93, map 21
(synonymy).

porterae Cockerell. B. C., Mont., Wyo., Nebr., Colo., Ariz., N. Mex., Tex. Parasite: Chrysis
lauta Cress. Pollen: Unknown, but visits flowers of Asclepias, Erigeron, Penstemon
coloradensis, P. occidentata, Petalostemon, Phacelia distans, Psilostrophe.
Anthidium porterae Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 411. d, °.
Anthidium porterae var. amabile Cockerell, 1904. Entomologist 37: 7. 6.
Anthidium porterae personulatum Cockerell, 1907. Canad. Ent. 39: 135. 2, 6.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 247. —Custer and Hicks, 1927. Biol. Bul. 52:
258-267 (nesting habits). —Custer, 1928. Ent. News 39: 123 (nesting habits).
psoraleae Robertson. Colo. and N. Dak., east to Ill. and Mich.; although it has been
questionably listed from B. C., Calif. and Ariz., there are no recent records west of Colo.
Pollen: Unknown, but visits flowers of Astragalus, Lobelia, Psoralea, Trifolium,
Verbena.
Anthidium psoraleae Robertson, 1902. Canad. Ent. 34: 322. 9, d.

Taxonomy: Schwarz, 1928. Canad. Ent. 60: 214. —Mitchell, 1962. N. C. Agr. Exp. Sta. Tech.
Bul. 152: 11-18, fig. 3 (redescription).
rodecki Schwarz. Colo., Nev. (Washoe Co.).
Anthidium rodecki Schwarz, 1934. Amer. Mus. Novitates 748: 1. d, 2.
sonorense Cockerell. Ariz., south. Nev. and south. Calif.; Mexico (Baja California and Sonora).
Pollen: Possibly an oligolege of Larrea tridentata, but a female has been collected at the
flowers of Prosopis.
Anthidium sonorense Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 91. d.
Anthidium sonorense productum Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 92. °.
Anthidium rohweri Schwarz, 1927. Amer. Mus. Novitates 253: 7. 6.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 32, figs. 13-15, 72, 90, 99,
map 22 (synonymy).
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 35 (floral relationships).
tenuiflorae Cockerell. N. W. T., Alta., Sask., Mont., Wyo., S. Dak., south to Calif., Ariz., N.
Mex., and Nebr. Ecology: A nest was constructed between two rocks. Pollen: Unknown,
but visits flowers of Arenaria kingii, Aster foliaceus, Astragalus, Epilobium
adenocaulum var. parishii, E. californicum, Gormania obtusata, Horkelia fusca, Lotus
argophyllus, L. davidsonii, Lupinus confertus, L. lyallii lobbii, Penstemon
secundiflorus, Phacelia frigida, P. heterophylla, Potentilla glandulosa, P. gracilis,
Solidago confinis, Streptanthus tortuosus, Trifolium oliganthum.
Anthidium tenuiflorae Cockerell, 1907. Canad. Ent. 39: 135. 2, d.
Anthidium tenuiflorae yukonense Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 18: 622. d,
25
Taxonomy: Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 379 (as tenuiflorae yukonense).
—Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 32-33, figs. 31-33, 88, 96, map 23
(synonymy).
Biology: Hicks, 1926. Colo. Univ., Studies 15: 217, 248 (nest).
utahense Swenk. B. C., Mont. and Utah, south to Calif., Nev. and Ariz. Ecology: Nests in
ground in holes and crevices frequently using cavities made by other insects and

animals as well as artificial nest cavities which offer vertical holes at ground level.
Parasite: Diorys productus (Cress.)? Pollen: Polylectic, stores pollen of Phacelia and

1992 Hymenoptera in America North of Mexico

Melilotus, but visits a wide variety of flowers, some of which may also serve as pollen
sources including Antirrhinum coulterianum, Artemisia, Aster, Astragalus bolanderi,
Brodiaea laxa, Calycadenia multiglandulosa, Calyptridiwm umbellatum, Ceanothus,
Chaenactis glabriuscula, Chamaebatia foliolosa, Cirsium vulgare, Clarkia amoena, C.
cylindrica, C. speciosa speciosa, C. unguiculata, Croton californicus, Cryptantha
intermedia, C. muricata, eriastrum plurifolium, Eriodictyon californicum, Eriogonum
fasciculatum, E. f. polifolium, Grindelia, Hemizonia lobbii, Horkelia bolanderi, H. b.
var. clevelandii, H. b. var. parryi, Lotus argophyllus, L. davidsonii, L. hamatus, L.
humistratus, L. nevadensis, L. nuttallianus, L. purshianus, L. scoparius, L. strigosus
var. hirtellus, Lupinus bicolor, L. nanus, Melilotus alba, Oenothera, Penstemon,
Phacelia ramosissima.

Anthidium utahense Swenk, 1914. Nebr. Univ., Studies 14: 23. 6, °.

Anthidium sagittipictum Swenk, 1914. Nebr. Univ., Studies 14: 20. 9.

Anthidium divisum Cockerell, 1925. Calif. Acad. Sei., Proc. (4) 14: 350. @.

Anthidium divisum nanuluwm Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 350. 9.

Anthidium divisum ornatifrons Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 350. °.

Anthidium brachyurum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 357. 6.

Taxonomy: Schwarz, 1930. N. Y. Ent. Soe., Jour. 38: 10 (divisuwm). —Grigarick and Stange,
1968. Calif. Ins. Surv., Bul. 9: 33-35, figs. 31-33, 88, 96, map 23 (synonymy).

Biology: Jaycox, 1966. Pan-Pacifie Ent. 42: 18-20 (nest, parasite). —Jaycox, 1967. Kans. Ent.
Soc., Jour. 40: 569 (interspecific relationships).

Genus CALLANTHIDIUM Cockerell

Callanthidium Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 365.
Type-species: Anthidium illustre Cresson. Orig. desig.
formosum (Cresson). Mont., Colo., Nev., Oreg., Calif., Ariz. (Oak Creek Canyon). Pollen:

Unknown, but visits flowers of Astragalus, Castilleia, Clarkia cylindrica, Lotus
argophyllus, Monardella linoides, Medicago sativa, Phacelia.

Anthidium formosum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 112. d.

Anthidium conspicuum Cresson, 1879. Amer. Ent. Soc., Trans. 7: 207. 9°.

Anthidium illustre var. consonum Cresson, 1879. Amer. Ent. Soc., Trans. 7: 207. °.

Dianthidium balli Titus, 1902. Ent. News 13: 170. 2.

Callanthidium formosum pratense Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 366. d.

Taxonomy: Cockerell, 1909. Ent. News 20: 262. —Grigarick and Stange, 1968. Calif. Ins.
Survey, Bul. 9: 36, figs. 101-105, map 25 (geogr. and floral records).
illustre (Cresson). Utah, Oreg., Calif., Nev., Ariz., N. Mex.; Mexico (Baja California). Ecology:
Nests in dead, floral seape of Yucca whipplei and in stump of oak. Parasite: Diorys
aurifuscus (Titus)?, Nemognatha scutellaris LeC., Sphaeropthalma unicolor (Cress.).
Pollen: Polylectic, stores pollen of Lotus scoparius in southern Calif., visits a wide
variety of flowers including Artemisia californica, Astragalus douglasii, Calochortus,
Cirsium, Castilleia plagiotoma, Clarkia, Collinsia, Cordylanthus nevinii, Cryptantha
intermedia, Eriodictyon crassifolium, E. trichocalyx, E. t. var. lanatum, Helianthus,
Lathyrus odoratus, Lotus argophyllus, L. oblongifolius, L. scoparius, Lupinus albifrons,
Malacothamnus arcuatus, Medicago sativa, Penstemon, Phacelia distans, P.
grandiflora, P. heterophylla, P. imbricata, P. ramosissima, Turricula parryi.
Anthidium illustre Cresson, 1879. Amer. Ent. Soc., Trans. 7: 206. °.
Anthidium serranum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 24. ¢.

Taxonomy: Cockerell, 1909. Ent. News 20: 262. ¢. —Schwarz, 1940. Amer. Mus. Novitates
1058: 3. —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 36-38, figs. 4, 106-110, map
26 (geogr. and floral records).

Biology: Johnson, 1904. Ent. News 15: 284 (nest). —Hicks, 1929. Canad. Ent. 61: 1-8 (nest,
parasite). —Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 96 (nest).

Genus DIANTHIDIUM Cockerell
Anthidium subg. Dianthidiwm Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 412.

Superfamily APOIDEA 1993

Type-species: Dianthidium sayi Cockerell. Orig. desig. (=Anthidium curvatum
Cockerell).

Taxonomy: Schwarz, 1926. Amer. Mus. Novitates 226: 1-15 (N. Amer. spp.). —Timberlake,
1943. N. Y. Ent. Soe., Jour. 51: 71-109 (racial differentiation of Nearctic spp.).
—Timberlake, 1948 N. Y. Ent. Soc., Jour. 56: 149-153 (additions and corrections).
—Timberlake, 1949. Pan-Pacific Ent. 25: 129-132 (Baja Calif. spp.). —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 152: 15-18, figs. 1, 5, table 1 (eastern U. S. spp.). —Grigarick and
Stange, 1968. Calif. Ins. Survey, Bul. 9: 38-58, figs. 5, 138-203, 221-224, maps 27-39 (Calif.
Spp.)-

Biology: Hurd and Linsley, 1950. N. Y. Ent. Soe., Jour. 58: 247-250 (parasites and inquilines).
—Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 39 (summary of literature).
—Clement, 1976. Wasmann Jour. Biol. 34: 9-10 (summary of literature).

concinnum (Cresson). Kans., Colo., Tex., Ariz. Ecology: Nests have been reported from
underside of stone and in dead branch of Chinese elm.
Anthidium concinnum Cresson, 1872. Amer. Ent. Soc., Trans. 4: 270. @, d.

Biology: Hungerford and Williams, 1912. Ent. News 23: 256 (nest). —Schwarz, 1928. N. Y.
Ent. Soc., Jour. 36: 408-409 (nest). —Fischer, 1951. Kans. Ent. Soc. Jour. 24: 46, figs. 1, 2
(nest).

cressonii (Dalla Torre). Utah, Colo., Nev., Ariz., N. Mex.
Anthidium venustum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 113. 2. Preoce.
Anthidiuwm cressonii Dalla Torre, 1896. Cat. Hym., v. 10, p. 458. N. name.
curvatum curvatum (Smith). N. C. to Fla., west to Mo. and Ark. Pollen: Unknown, but visits
flowers of Aster, Chrysopsis, Helenium, Koellia, Psoralea.
Anthidium curvatum Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 215. 9.
curvatum sayi Cockerell. Alta., Idaho, Utah, Colo. and Kans., south to east. Calif., Ariz. and
Tex. Ecology: Nests in aggregations in the soil, uses resin from stems and leaves of
Helianthus petiolaris for nest construction. Parasite: Dasymutilla asopus bexar (Blake),
Eusapyga rubripes Cress., Monodontomerus montivagus Ashm., Rhydinofoenus
pattersonae Melander and Brues, Sphaeropthalma uro (Blake), Villa (Anthrax) sp.
Megachile interrupta Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 350. d, °.
Preoce.
Dianthidium Sayi Cockerell, 1907. Canad. Ent. 39: 136. N. name.

Taxonomy: Michener, 1953. Kans. Univ. Sci., Bul. 35: 1045, figs. 180-133 (larva). —Grigarick
and Stange, 1968. Calif. Ins. Surv., Bul. 9: 41-42, figs. 171-178, 178, 197, map 27 (tax.
characters, geogr. records).

Biology: Hungerford and Williams, 1912. Ent. News 23: 256 (nest, as curvatum). —Hicks,
1926. Amer. Nat. 9: 199 (nest). —Hicks, 1926. Colo. Univ., Studies 15: 249 (nest, parasite).
—Custer and Hicks, 1927. Biol. Bul. 52: 268 (nest) —Custer, 1928. Ent. News 39: 128
(nest). —Mickel, 1928. U. S. Natl. Mus., Bul. 143: 61 (parasite). —Fischer, 1951. Kans. Ent.
Soe., Jour. 24: 47-49 (nest).

curvatum xerophilum Cockerell. N. Mex., Ariz. (Apache Pass).
Dianthidium Sayi xerophilum Cockerell, 1907. Canad. Ent. 39: 136. °.
desertorum Timberlake. South. Calif. (Colorado Desert). Pollen: Unknown, but visits flowers
of Hyptis emory?.
Dianthidium desertorum Timberlake. 1943. N. Y. Ent. Soc., Jour. 51: 84. d, &.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 42, figs. 168-170, 184, 196,

map 28 (tax. characters, geogr. and floral records).
discors Timberlake. West Tex., Ariz.
Dianthidium discors Timberlake, 1948. N. Y. Ent. Soc., Jour. 56: 149. 9.
dubium dilectum Timberlake. Calif. (Central Coast and Peninsular ranges). Ecology: Nests
made of gravel and resin have been found attached to a leaf and on exposed clay shales.
Parasite: Amobia floridensis (Twn.), Chrysis coerulans Fabr., Sapyga nevadica Cress.,
Toxophora pellucida Coq. Pollen: Apparently polylectic, visits flowers of Achillea

1994 Hymenoptera in America North of Mexico

millefolium, Calochortus weedii, Chenopodium, Cirsium vulgare, Convolvulus,
Cordylanthus pilosus, Corethrogyne, Dicentra, Encelia californica, Erigeron,
Eriodictyon trichocalyx, E. t. var. lanatum, E. parryi, Eriogonum elegans, E.
fasciculatum, Helianthus gracilentus, Hesperochiron californicum, Lepidium
virginicum, Lotus americanus, L. humistratus, L. scoparius, Monardella villosa,
Penstemon, Phacelia ramosissima, Stachys pycnantha, Stephanomeria cichoriacea,
Trichostema parishii, Turricula parryi, Verbena lasiostachys, Viguiera multiflora.
Dianthidium dubium dilectum Timberlake, 1948. N. Y. Ent. Soc., Jour. 56: 152. 2, d.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 42-43, map 2 (tax.
characters, geogr. and floral records).

Biology: Hurd and Linsley, 1950. N. Y. Ent. Soc., Jour. 58: 247, 248 (nest, parasite).
dubium dubium Schwarz. South. Oreg. and north. Calif. (Coast Range, north of San Francisco;
Cascade Range; and west. slope of Sierra Nevada Mts.). Ecology: Nests are made of
resin and gravel attached to vegetation or rocks. Pollen: Apparently polylectic, visits
flowers of Bigelovia, Chamaebatia foliolosa, Clarkia biloba, C. cylindrica, C.
dudleyana, C. unguiculata, Collinsia tinctoria, Eriodictyon, Eriogonum latifoliwm
nudum, Gayophytum nuttallii, Lessingia leptoclada, Lotus americanus, L. nevadensis,
L. purshianus, Monardella lanceolata, Marrubium vulgare, Penstemon spectabilis,
Stachys ajugoides, Stephanomeria virgata, Trichostema lanceolatum, T. laxum,
Trifolium, Vicia.
Dianthidium dubium Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 405. 3, °.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 42, 43-44, figs. 5, 138-140,
185, 195, map 29 (tax. characters, geogr. and floral records).
dubium mecrackenae Timberlake. Calif. (east slope, Sierra Nevada Mts.). Pollen: Unknown,

but visits flowers of Eriastrum densifolium.
Dianthidium consimile mecrackenae Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 104. °,

3.
Dianthidium plenum convictorum Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 108. °.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 44-45, map 29 (tax.
characters, geogr. and floral records).
floridiense Schwarz. Fla. Ecology: Nests in borings. Pollen: Unknown but visits flowers of
Bidens, Sabal.
Dianthidium floridiense Schwarz, 1926. Amer. Mus. Novitates 226: 5. d, °.
Dianthidium floridense(!) Michener, 1951. In Muesebeck, Krombein and Townes, U. S.
Dept. Agr., Agr. Monog. 2: 1143.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 16-17 (redescription).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 272-273 (nest architecture, life
history).
heterulkei fraternum Timberlake. Tex., N. Mex., Ariz., Nev. Parasite: Anthrax irroratus Say.
Dianthidium fraternum fraternum Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 92. d, ¢.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 273-274 (nest architecture, life
history, parasite).
heterulkei heterulkei Schwarz. Calif. (Sierra Nevada Mts.), Oreg., Nev., Utah, Wyo. Ecology:
Constructs nests in natural depressions within voleanic outcroppings and possibly uses
lodgepole pine as source of resin. Parasite: Monodontomerus clementi Grissell. Pollen:
Polylectic, is known to collect pollen from flowers of Eriophyllum integrifolium,
Trifolium longipes, but also visits flowers of Aster, Chaenactis glabriuscula lanosa,
Chrysothamnus, Haplopappus suffruticosa, Solidago.
Dianthidium heterulkei Schwarz, 1940. Amer. Mus. Novitates 1058: 6. d, 9.
Dianthidium heterulkei var. cornucopiana Schwarz, 1940. Amer. Mus. Novitates 1058: 8. d.
Dianthidium fraternum hirtulum Timberlake, 1943. N. Y. Ent. Soe., Jour. 51: 94. d, °.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 45-46, figs. 159-161, 181,
194, map 30 (geogr. and floral records). —Clement, 1976. Wasmann Jour. Biol. 34: 16-18,

fig. 2 (larva).

Superfamily APOIDEA 1995

Biology: Clement, 1976. Wasmann Jour. Biol. 34: 9-22, 2 figs. (nest site, nest architecture, life
history, mating, floral relationships, parasite).
implicatum Timberlake. Tex. to south. Calif. and Nev.; Mexico (Chihuahua). Ecology: A pebble
and resin nest attached to a twig of Dalea. Pollen: Unknown, but visits flowers of
Cevallia, Gutierrezia microcephala.
Dianthidium implicatum Timberlake, 1948. N. Y. Ent. Soc., Jour. 56: 150. 9.

Taxonomy: Grigarick and Stange, 1964. Pan-Pacifie Ent. 40: 152. (male). —Grigarick and
Stange, 1968. Calif. Ins. Survey, Bul. 9: 46, figs. 150-152, 174, 190, map 31 (geogr. and floral
record).

marshi Grigarick and Stange. Calif., Ariz. (Santa Catalina Mts.). Pollen: Unknown, but visits
flowers of Bebbia, Eriastrum densifolium.
Dianthidium marshi Grigarick and Stange, 1964. Pan-Pacific Ent. 40: 149, figs. 1, 2, 3, 10,
See

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 46-47, figs. 147-149,
186-187, 201, map 32 (geogr. and floral records).
parkeri Grigarick and Stange. Ariz., Tex.; Mexico (Sonora, Sinaloa, Durango). Pollen:
Unknown, but visits flowers of Verbesina encelioides.
Dianthidium parkeri Grigarick and Stange, 1964. Pan-Pacifie Ent. 40: 150, figs. 4, 5, 6, 12.
OQe
parvum parvum (Cresson). N. Mex., west to Colo., Utah, Nev. and Calif.; Mexico. Pollen:
Unknown, but visits flowers of Chrysothamnus, Grindelia serrulata, Hyptis emoryi,
Palafoxia linearis, Senecio douglasii.
Anthidium parvum Cresson, 1878. Amer. Ent. Soe., Trans. 7: 114. 2, d.
Dianthidium profugum Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 90. °.
Dianthidium parvum heteropoda Schwarz, 1934. Amer. Mus. Novitates 7438: 4. °.
Dianthidium parvum basingeri Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 97. 6.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 47-48, figs. 153-155, 183,
188, 190, map 33 (synonymy, geogr. and floral records).
parvum schwarzi Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Asclepias

erosa, Chrysothamnus nauseosus mohavensis, Corethrogyne bernardina, Croton
californicus, Encelia californica, Erigeron foliosus var. stenophyllus, Eriogonum
fasciculatum polifolium, E. gracile, Grindelia, Gutierrezia californica, G. microcephala,
G. sarothrae, Heterotheca grandiflora, Lepidospartum squamatum, Lotus scoparius,
Senecio douglasii, Stephanomeria exigua.

Dianthidium parvum schwarzi Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 96. 2, d.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 48, map 33 (geogr. and
floral records).
platyurum mohavense Timberlake. Calif. Pollen: Unknown, but visits flowers of Erigeron,

Eriogonum fasciculatum.
Dianthidium platyurum mohavense Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 87. ?.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 49, map 34 (geogr. and
floral records).
platyurum platyurum Cockerell. Tex., Ariz., west. Nev., Calif.; Mexico (Baja California).
Pollen: Unknown, but visits flowers of Bebbia juncea, Encelia californica, Erigeron
foliosus var. stenophyllus, Eriastrum densifolium, E. plurifolium, E. virgatum,
Eriogonum, Gutierrezia californica, G. sarothrae, Heterotheca grandiflora, Lotus
scoparius, Malacothrix, Phacelia ramosissima, Senecio douglasti, Stephanomeria
exigua, S. virgata.
Dianthidium platywrum Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 90. d, °.
Dianthidium parvum baculifrons Cockerell, 1925. Calif. Acad. Sci., Proce. (4) 14: 365. °.
Dianthidium ulkei riparii Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 400. d, °.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 49-50, figs. 165-167, 180,
200, map 34 (geogr. and floral records).

1996 Hymenoptera in America North of Mexico

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 274-275 (nest architecture, life
history).
plenum Timberlake. Oreg., Calif. and west. Nev. Ecology: A resin and gravel nest attached in
crotch of apple tree. Pollen: Unknown, but visits flowers of Chaenactis glabriuscula
lanosa, Monardella lanceolata, Phacelia.
Dianthidium plenum plenum Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 106. d, 2.
Dianthidium plenum williamsi Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 107. d.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 50-51, figs. 144-146, 175,
193, 225, map 35 (synonymy, geogr. and floral records).
pudicum consimile (Ashmead). Ariz., Calif.; Mexico (Baja California). Ecology: Constructs

nest of resin and large grains of sand or gravel, either in the crotches of terminal
branches of shrubs or in depressions or angles of stones or boulders lying on the
ground. Parasite: Eusapyga verticalis Cress., Leucospis affinis Say, Microdontomerus
anthidii (Ashm.), Monodontomerus montivagus Ashm., Sapyga minor Roberts. Pollen:
Unknown, but visits flowers of Cirsium vulgare, Corethrogyne, Cryptantha intermedia,
Encelia californica, Eriastrum densifolium, E. plurifolium, Eriodictyon crassifolium,
E. trichocalyx var. lanatum, Eriogonum fasciculatum, E. inflatum, E. latifoliwm
nudum, Grindelia camporum, G. robusta, G. stricta procumbens, Gutierrezia
californica, G. microcephala, G. sarothrae, Haplopappus, Krameria grayi, Lonicera
hispidula var. vacillans, Lotus glaber, L. scoparius, Melilotus indica, Monardella
villosa, Phacelia distans, P. ramosissima, Salvia, Stachys pycnantha, Stephanomeria
cichoriacea, S. exigua, Trichostema lanceolatum. Predator: Trichodes ornatus ornatus
Say, T. o. tenellus LeC.

Anthidium consimile Ashmead, 1896. Ent. News 7: 25 “2” = 6.

Dianthidium provancheri Titus, 1906. Ent. Soc. Wash., Proce. 7: 164. d.

Dianthidium plenum australe Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 108. ?.

Dianthidium macswaini Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 109. @.

Dianthidium pudicum peninsulare Timberlake, 1949. Pan-Pacific Ent. 25: 130. 3.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 51-52, map 36 (synonymy,
geogr. and floral records).

Biology: Davidson, 1896. Ent. News 7: 22 (nest, parasites). —Hicks, 1934. Colo. Univ. Studies
21: 265 (nest, parasites). —Linsley and MacSwain, 1942. Ent. Soc. Amer., Ann. 36: 589
(parasite). —Hurd and Linsley, 1950. N. Y. Ent. Soc. 58: 248, 249 (parasites). —Grigarick
and Stange, 1968. Calif. Ins. Survey, Bul. 9: 52 (summary of literature).

pudicum pudicum (Cresson). B. C. and Alta., south to Ariz. and Calif. Ecology: Constructs

nest of resin and gravel either in crotches of small trees or shrubs or on underside of
rocks. Parasite: Eusapyga proxima (Cress.), EZ. rubripes (Cress.), Monodontomerus
montivagus Ashm., Sapyga sp., Zonitis sp. Pollen: Unknown, but visits flowers of Aster,
Chaenactis glabriuscula lanosa, Chrysothamnus, Cirsium vulgare, Cleome lutea,
Eriastrum densifolium, Gormania obtusata, Medicago sativa, Oenothera clavaeformis
cruciformis, Streptanthus tortwosus, Symphoricarpos.

Anthidiwm pudicum Cresson, 1879. Amer. Ent. Soc., Trans. 7: 208. d.

Anthidiwm pudens Cresson, 1879. Amer. Ent. Soc., Trans. 7: 208. °.

Dianthidium pudicum decorum Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 100. 2, d.

Dianthidium pudicum inyoense Timberlake, 1943. N. Y. Ent. Soe., Jour. 51: 101. 2, d.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Surv., Bul. 9: 53-54, figs. 141-143, 176, 191,
map 36 (synonymy, geogr. and floral records).

Biology: Hicks, 1927. Psyche 34: 193. (nest, parasites). —Hicks, 1931. Canad. Ent. 63: 173
(nest). —Hicks, 1934. Colo. Univ. Studies 21: 268 (nest, parasites). —Roberts, 19338. Kans.
Ent. Soc., Jour. 61: 91 (parasite). —Hurd and Linsley, 1950. N. Y. Ent. Soc., Jour. 58: 248
(parasites, as pudicum decorum). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9:
53 (summary of literature). —Clement and Rust, 1974. Pan-Pacifie Ent. 50: 87-89, 1 fig.
(nest).

simile (Cresson). Minn. to Ont. and Maine, south to Ga. Ecology: Reared from partly rotted
log. Pollen: Unknown, but visits flowers of Aster, Clethra.
Anthidium simile Cresson, 1864. Ent. Soc. Phila., Proc. 2: 378. 2, ¢.

Superfamily APOIDEA 1997

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 17-18, fig. 5 (redescript.).

Biology: Fischer, 1951. Kans. Ent. Soc., Jour. 24: 46-47 (nest).
singulare (Cresson). Calif., Nev. Ecology: Constructs nest of resin and pebbles on side or face

of rocks. Pollen: Unknown, but visits flowers of Chaenactis glabriuscula lanosa,
Chrysanthemum, Chrysothamnus, Cirsium californicum, Erysimum asperum, Lupinus
parishu, Perideridia, Solidago californica, Viguiera multiradiata.

Anthidium singulare Cresson, 1879. Amer. Ent. Soc., Trans. 7: 207. 9.

Dianthidium singulare var. perluteum Cockerell and Cockerell, 1904. In Cockerell, South.
Calif. Acad. Sci., Bul. 3: 23. °.

Dianthidium singulare melanognathum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 363.
o.

Taxonomy: Cockerell, 1925. Calif. Acad. Sci. Proc. (4) 14: 363. d, 2 (singulare perluteum).
—Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 54, figs. 135-137, 177, 202-203, map
37 (synonymy, geogr. and floral records).

Biology: Michener, 1935. Pan-Pacific Ent. 11: 23-24 (nest). —Timberlake, 1943. N. Y. Ent.
Soc., Jour. 51: 75 (nest). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 54
(summary of literature).

subparvum Swenk. B. C., Idaho and Mont., south to Calif., Nev., Utah and Wyo. Pollen:

Possibly oligolege of Compositae, visits flowers of Aster alpigenus, A. andersonii, A.
canescens, A. foliaceus, Chaenactis glabriuscula, Chaetopappa aurea, Chenopodium,
Chrysothamnus viscidiflorus, Clematis pauciflora, Corethrogyne bernardina, Erigeron
divergens, Erigeron foliosus, E. f. var. stenophyllus, Eriogonum fasciculatum,
Grindelia, Gutierrezia californica, G. sarothrae, Haplopappus suffruticosus, Linum,
Lotus strigosus var. hirtellus, Machaeranthera tephrodes, Phacelia, Ranunculus,
Senecio douglasii, Solidago.

Dianthidium subparvum Swenk, 1914. Nebr. Univ., Studies 14: 30. 2, 6.

Dianthidium semiparvum Schwarz, 1926. Amer. Mus. Novitates 226: 12. 3.

Dianthidium semiparvum gallatinae Schwarz, 1927. Amer. Mus. Novitates 277: 6. 6, °.

Dianthidium parvum var. swenki Schwarz, 1928. N. Y. Ent. Soe., Jour. 36: 402. 3 (2
misdet.).

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 54-56, figs. 156-158, 182,
189, 192, map 38 (synonymy, geogr. and floral records).
subrufulum Timberlake. Mo., Ark., Tex.
Dianthidium subrufulum Timberlake, 1943. N. Y. Ent. Soc., Jour. 51: 78. 6, 2.

ulkei cooleyi Schwarz. Mont.
Dianthidium ulkei cooleyi Schwarz, 1927. Amer. Mus. Novitates 277: 7. °.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 58 (taxonomic status).

ulkei perterritum Cockerell. Nebr., N. Mex., Ariz. Ecology: Nests in borings, constructs nest
using resin and pebbles. Parasite: Nemognatha nigripennis LeC.
Dianthidium ulkei perterritum Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 191. 9.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 275-276, pl. 17, fig. 81 (nest
architecture, supersedure, life history, parasite).

ulkei ulkei (Cresson). B. C., Idaho, Mont. and S. Dak. south to Calif., Nev., Utah, N. Mex. and
Nebr. Ecology: Nests in abandoned nests of other aculeates, cavities or short tunnels in
the ground, and trap nests. Parasite: Husapyga verticalis (Cress.). Pollen: Possibly an
oligolege of Compositae, but visits a wide variety of flowers including Achillea
millefolium, Aster adscendens, A. canescens, Calycadenia multiglandulosa, Centaurea,
Chaetopappa aurea, Chrysopsis fastigiata, C. villosa, Chrysothamnus nauseosus, C.
viscidiflorus, C. v. typicus, Cleomella serrulata, Cordylanthus filifolius, Encelia
californica, Eriastrum densifolium, Erigeron divergens, E. foliosus var. stenophyllus,
Eriodictyon, Eriogonum fasciculatum, E. subscaposum, E. vimineum, Gilia capitata,
Grindelia camporum, G. serrulata, Haplopappus arborescens, H. bloomeri, Helenium
bigelovii, Helianthus petiolaris, Heterotheca grandiflora, Lessingia leptoclada, Lotus
davidsonii, L. scoparius, Machaeranthera tephrodes, Medicago sativa, Melilotus alba,

1998 Hymenoptera in America North of Mexico

Monardella laceolata, Ranunculus, Senecio douglasii, S. ionophyllus, Solidago

californica, Stephanomeria californica, S. cichoracea, S. virgata, Vicia cracca, Viguiera
multiflora.

Anthidium ulkei Cresson, 1878. Amer. Ent. Soc., Trans. 7: 115. °.
Anthidium davidsoni Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 5. ¢.
Dianthidium ulkei reductum Timberlake, 1943. N. Y. Ent. Soce., Jour. 51: 82. 2, 6.

Taxonomy: Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 56-58, figs., 162-164, 179,
198, map 39 (synonymy, geogr. and floral records).

Biology: Hicks, 1926. Colo. Univ., Studies 15: 250 (nest). —Hicks, 1933. Ent. News 44: 75

(nest). —Parker and Bohart, 1966. Pan-Pacifie Ent. 42: 96. —Grigarick and Stange, 1968
Calif. Ins. Survey, Bul. 9: 58 (summary of literature).

Genus ANTHIDIELLUM Cockerell

Anthidium subg. Anthidiellum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 3.
Type-species: Anthidiuwm strigatum Panzer. Orig. desig.

Anthidiwm subg. Ceri-anthidium Friese, 1923. Die Europ. Bienen, p. 304.
Type-species: Anthidium strigatum Panzer. Desig. by Cockerell, 1925.

Cerianthidium Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 361. Emend.

Revision: Schwarz, 1926. Amer. Mus. Novitates 226: 15-19 (N. Amer. spp.)

Taxonomy: Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 369-418, 23 figs. (status of spp. and
subspp.). —Schwarz, 1957. Kans. Ent. Soc., Jour. 30: 137 (key to spp. of notatwm complex).
—Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 18-23, figs. 1, 6-7, table 1 (eastern U.
S. spp.). —Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 58-62, figs. 204-211, maps
40-41 (Calif. spp.).

ehrhorni (Cockerell). Ariz., Nev., Calif.; Mexico (Baja California). Ecology: Constructs a single
celled nest of resin attached to twigs or flat pieces of wood. Pollen: Apparently
polylectic, visits flowers of Asclepias, Baileya, Chaenactis fremontii, Dalea schottii,
Encelia, Eriogonum inflatwm, Geraea canescens, Larrea tridentata, Lotus scoparius,
Phacelia distans, P. ramosissima, Pluchea sericea, Prosopis, Stephanomeria virgata,
Trixis californica.
Anthidium (Dianthidium) ehrhorni Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 414. 6.

Taxonomy: Schwarz, 1929 (1928). N. Y. Ent. Soc., Jour. 36: 392. ?. —Michener, 1953. Kans.
Univ. Sci. Bul. 35: 1045, figs. 119, 123, 124, 127 (larva). —Grigarick and Stange, 1968. Calif.
Ins. Survey, Bul. 9: 59-60, figs. 208-211, map 40 (geogr. and floral records).

Biology: Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 397 (nest). —Grigarick and Stange, 1968.

Calif. Ins. Survey, Bul. 9: 60, fig. 227 (nest). —Hurd and Linsley, 1975. Smithsn. Contrib.
Zool. 193: 35-36 (floral relationships).

notatum boreale (Robertson). Ill., Nebr. Pollen: Unknown, but visits flowers of Verbena
stricta.

Dianthidium boreale Robertson, 1902. Canad. Ent. 34: 323. 6.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 20-21 (redescription).
notatum gilense (Cockerell). Colo., Ariz., N. Mex., Tex.
Anthidiwm gilense Cockerell, 1897. Canad. Ent. 29: 222. °.
Taxonomy: Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 413. d.

notatum notatum (Latreille). Mass. to III., south to Fla. and Miss. Pollen: Apparently
polylectic, visits flowers of Afzelia, Baptisia, Chrysopsis, Desmodium, Erigeron,

Eupatorium, Galactia, Helianthus, Melilotus, Monarda, Phaseolus, Psoralea, Solidago,
Trifolium.

Anthidium notatum Latreille, 1809. Mus. Nat. Hist., Ann. 13: 48, 231. ¢.

Taxonomy: Cresson, 1864. Ent. Soc. Phila., Proc. 2: 376. 2, d6. —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 19-20, fig. 6 (redescript., floral records).

Superfamily APOIDEA 1999

notatum robertsoni (Cockerell). B. C., Idaho, Colo., Utah, Oreg., Calif., Nev., Ariz.; Mexico

(Baja California). Pollen: Polylectic, visits a wide variety of flowers including Agastache,
Amorpha californica, Aster adscendens, Astragalus, Chrysothamnus, Clarkia biloba, C.
elegans, C. rhomboidea, Cordylanthus pilosus, C. rigidus, Clematis pauciflora,
Cryptantha intermedia, Dalea, Eriodictyon, Eriogonum fasciculatum, E. latifolium
nudum, Grindelia, Gutierrezia californica, Heliotropium curassavicum, Hemizonia
fasciculata, Lippia, Lonicera hispidula var. vacillans, Lotus americanus, L.
argophyllus, L. nevadensis, L. oblongifolius, L. purshianus, L. scoparius, Marrubium
vulgare, Melilotus alba, Monardella lanceolata, Penstemon, Phacelia distans, P.
ramosissima, Salvia, Solidago californica, Stellaria longipes, Stephanomeria exigua,
Trichostema lanceolatum, Wislizenia refracta.

Dianthidium robertsoni Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 4. 2, ¢.

Anthidiellum robertsoni race citrinellum Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14:
362. d.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1046, figs. 121, 122, 128 (larva).
—Grigarick and Stange, 1968. Calif. Ins. Survey, Bul. 9: 60-62, figs. 204-207, 228, map. 41
(geogr. and floral records).

Biology: Leech, 1947. Ent. Soc. Brit. Col., Proc. 44: 39, 2 figs. (nest). —Linsley, 1962. Ent.

Soe. Amer., Ann. 55: 159-160, figs. 6-7 (sleep). —Grigarick and Stange, 1968. Calif. Ins.
Survey, Bul. 9: 62, fig. 228 (nest, summary of literature).
notatum rufimaculatum Schwarz. Fla. Pollen: Unknown, but visits flowers of Rhus.
Anthidiellum notatum var. rufimaculatum Schwarz, 1926. Amer. Mus. Novitates 226: 15.
Gime

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 21 (redescription).

Biology: Schwarz, 1928. N. Y. Ent. Soc., Jour. 36: 395, pl. 12, figs. 1A, 1B (nest).

perplexum (Smith). N. C. to Fla. Pollen: Unknown, but visits flowers of Afzelia, Amorpha,
Bidens, Chrysopsis, Desmodium, Erigeron, Hypericum, Ilex, Kuhnistera, Linaria,
Monarda, Ocimum, Polygonum, Psoralea, Ptilimnium, Rhus, Solidago.
Anthidium perplexum Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 214. 2, 6.

Taxonomy: Cresson, 1864. Ent. Soc. Phila., Proc. 2: 377. 2, o.

Genus HETEROSTELIS Timberlake

Stelis subg. Heterostelis Timberlake, 1941. N. Y. Ent. Soe., Jour. 49: 125.
Type-species: Stelis anthidioides Timberlake. Orig. desig.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 33-35, fig. 11, table 2 (eastern
U.S. spp.). —Thorp, 1966. Kans. Ent. Soc., Jour. 39: 131-146, 16 figs. (N. Amer. spp.).
—Pasteels, 1968. Nat. Canad. 95: 1057 (tax. status).

anthidioides (Timberlake). Calif.
Stelis (Heterostelis) anthidioides Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 123. 2.

Taxonomy: Thorp, 1966. Kans. Ent. Soc., Jour. 39: 134, 141-142, fig. 8 (tax. characters, geogr.
records).
australis australis (Cresson). N. J. to Fla.
Stelis australis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 92. 9.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 33. 2 (redescription).
—Thorp, 1966 Kans. Ent. Soc., Jour. 39: 136, figs. (tax. characters of male and female,
geogr. and floral records).

australis floridensis Mitchell. Fla.
Heterostelis australis floridensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 33,
fig mle enor

Taxonomy: Thorp, 1966. Kans. Ent. Soc., Jour. 39: 136-137 (tax. characters).

grossa Mitchell. Fla. (Gainesville), Ala. (Ft. Morgan).
Heterostelis grossa Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 35. °.

2000 Hymenoptera in America North of Mexico

Taxonomy: Thorp, 1966. Kans. Ent. Soc., Jour. 39: 134, 137-138, fig. 5 (tax. characters, geogr.
record).

hurdi Thorp. Calif. Host: Trachusa perdita Ckll.
Heterostelis hurdi Thorp, 1966. Kans. Ent. Soc., Jour. 39: 142, figs. 3, 9, 11, 13-16. 2, 6.

Taxonomy: Thorp, 1966. Kans. Ent. Soc., Jour. 39: 143-144, figs. 14-16 (cocoon, larva, pupa).

Biology: Thorp, 1966. Kans. Ent. Soc., Jour. 39: 132, 145 (nest, parasite).

manni (Crawford). Ariz. Host: Trachusa manni Cwfd.?
Stelis manni Crawford, 1917. Ent. Soe. Wash., Proc. 19: 168. °.

Taxonomy: Thorp, 1966. Kans. Ent. Soc., Jour. 39: 134, 140-141, fig. 7 (tax. characters).

texana Thorp. Tex. (Austin).
Heterostelis texana Thorp, 1966. Kans. Ent. Soc., Jour. 39: 135, figs 2, 6, 12. ¢.

Genus STELIS Panzer

Insofar as known the species of this genus are cleptoparasites, chiefly in the nests of pol-
len-collecting bees of the tribe Megachilini (Ashmeadiella, Chalicodoma, Heriades, Hoplitis,
Osmia, Proteriades and Robertsonella). However, one of our species has been reared from a
nest of the genus Anthidiwm and some extralimital species of Stelis have been shown to be clep-
toparasites in the nests of Ceratina and Euglossa.

Taxonomy: Cockerell, 1898. Entomologist 31: 166-167 (U.S. spp.). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 35-47 figs. 12-14, table 2 (eastern U.S. spp.). —Rozen, 1966. N.
Y. Ent. Soe., Jour. 74: 84-91, 18 figs. (immature stages). —Pasteels, 1968. Nat. Canad. 95:
1055-1063 (tax. status, affinities and origin).

Genus STELIS Subgenus STELIS Panzer

Stelis Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 246.

Type-species: Apis punctulatissima Kirby. Monotypic. (=Apis aterrima Panzer).
Gyrodroma Klug, 1807. Mag. Insektenk. 6: 198.

Type-species: Apis punctulatissima Kirby. Monotypic. (=Apis aterrima Panzer).
Gymnus Spinola, 1808. Insectorum Liguriae, v. 2, p. 9.

Type-species: Apis punctulatissima Kirby. Monotypic. (=A pis aterrima Panzer).
Ceraplastes Gistel, 1848. Naturgesch. Thierr. f. hoh. Schul., p. x. Proposed unnecessarily to

replace Stelis.

This subgenus does not occur in North America.
Genus STELIS Subgenus PROTOSTELIS Friese

Stelis subg. Protostelis Friese, 1895. Die Bienen Europas v. 1, p. 25.
Type-species: Stelis freygessneri Friese. Desig. by Popov, 1938.

arizonensis Swenk. Ariz.
Stelis (Microstelis) arizonensis Swenk, 1915. Nebr. Univ., Studies 15: 189. °, ¢.

costalis costalis Cresson. Va. and Ky., south to Fla. and Tex.
Stelis costalis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 274. 2, d.
Stelis louisae Cockerell, 1911. U. S. Natl. Mus., Proc. 40: 247. ¢.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 37-38, figs 12-13
(redescription).
costalis floridana Graenicher. Fla. Host: A resin-using anthidiine bee or a species of
Chalicodoma subg. Chelostomoides.
Stelis floridana Graenicher, 1928. Ent. News 39: 282. 2, d.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 38-39 (redescription).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 481-482 (cocoon, host).

laticincta Cresson. Calif.
Stelis laticincta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 92. 2.

Taxonomy: Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 3. d.

superfamily APOIDEA 2001

perpulchra Crawford. Ariz., Calif., Mexico (Baja California).
Stelis perpulchra Crawford, 1916. Insecutor Inscitiae Menstruus 4: 107. é.

Taxonomy: Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 89. 2, ¢.
rudbeckiarum Cockerell. N. Mex., Ariz.
Stelis rudbeckiarum Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 3. 2.
Stelis rudbeckiana(!) Snow, 1906. Kans. Acad. Sci., Trans. 20: 138.

Genus STELIS Subgenus MICROSTELIS Robertson

Microstelis Robertson, 1903. Amer. Ent. Soc., Trans. 29: 170.
Type-species: Stelis lateralis Cresson. Orig. desig.
coarctatus Crawford. Kans., Nev. (Elko County). Host: Proteriades shoshone Parker.
Stelis coarctatus Crawford, 1916. Insecutor Inscitiae Menstruus 4: 105. 2.

Biology: Parker, 1976. Pan-Pacifie Ent. 52: 78 (host).

crassiceps Cockerell. Colo. Host: Hoplitis producta (Cress.).
Stelis crassiceps Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 18: 625. o.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 246 (host).
lateralis Cresson. Maine and Ont. to N. Dak., south to Ga. and Tex. Host: Hoplitis cylindrica
(Cress.), H. pilosifrons (Cress.), H. producta (Cress.). Predator: Philanthus pulcher

Dalla Torre.
Stelis lateralis Cresson, 1864. Ent. Soc. Phila., Proc. 2: 410. 9.

Taxonomy: Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 48. ¢, 2. —Michener, 1953, Kans.
Univ. Sci. Bul. 35: 1048, figs. 114-116, 118 (larva). —Mitchell, 1962. N. C. Agr. Expt. Sta.
Tech. Bul. 152: 42-43, fig. 14 (redescription). —Rozen, 1966. N. Y. Ent. Soc., Jour. 74: 86-89,
figs. 9-10 (larva).

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 153 (as sexmaculatus). —Swenk,
1914. Nebr. Univ., Studies, 14: 5 (host). —Hicks, 1926. Colo. Univ., Studies 15: 217 (life
history, host). —Rau, 1928. Psyche 35: 100 (life history, host). —Michener, 1955. Kans. Ent.
Soc., Jour. 28: 83-86 (life history, host).

maculata (Provancher). B. C.
Heriades maculatum Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 323. 2.

Taxonomy: Titus, 1906. Ent. Soc. Wash., Proc. 7: 162 (tax. characters).
plena (Provancher). Ont. (Ottawa).
Heriades plenum Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 425. 6.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 43 (status).

sexmaculata Ashmead. Calif., Nev., ?Mo., ?Colo. Host: Hoplitis brachyodonta (Ckll.), H. colei
(Cwfd.), H. grinnelli Ckll., H. hypocrita (Ckll.), H. producta gracilis (Mich.), H. producta
producta (Cress.), Osmia glauca (Fowler). Parasite: Epistenia sp., Leucospis affinis Say,
Microdontomerus anthidii (Ashm.).
Stelis 6-maculata Ashmead, 1896. Ent. News 7: 218. °.
Stelis sexmaculata Cockerell, 1898. Entomologist 31: 167. Emend.

Taxonomy: Cockerell, 1934. Amer. Mus. Novitates 732: 6. d. —Rust and Clement, 1972.
Kans. Ent. Soe., Jour. 45: 526 (cocoon, host).

Biology: Davidson, 1896. Ent. News 7: 218 (host). —Davidson, 1897 South. Calif. Acad. Sci.,
Proce. 1: 4 (life history, hosts). —Hicks, 1926. Colo. Univ., Studies 15: 217 (life history,
hosts). —Rau, 1928. Psyche 35: 100 (Colo. and Mo. records may refer to lateralis).
—Parker and Bohart, 1966. Pan-Pacific Ent. 42: 98 (parasites).

vernalis Mitchell. Pa., N. C., Mich., Minn. Host: Heriades carinata Cress.
Stelis (Microstelis) vernalis Mitchell, 1962. N C. Agr. Expt. Sta. Tech. Bul. 152: 43. 2, ¢.

Biology: Matthews, 1965. Amer. Ent. Inst., Contrib. 1: 24-25 (behavior, host).
Genus STELIS Subgenus PAVOSTELIS Sladen
Stelis subg. Pavostelis Sladen. 1916. Canad. Ent. 48: 313.

2002 Hymenoptera in America North of Mexico

Type-species: Stelis montana Cresson. Monotypic.

anthracina Timberlake. Calif.
Stelis (Pavostelis) anthracina Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 125. ¢.

ater Mitchell. Fla.
Stelis (Pavostelis) ater Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 39. 2.

callura Cockerell. Colo., Utah.
Stelis callura Cockerell, 1925. Calif. Acad. Sei., Proc. (4) 14: 212. 6.

carnifex Cockerell. Nev., Calif.
Stelis carnifex Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 769. °.

diversicolor Crawford. Tex. to Tenn. and N. C.
Stelis diversicolor Crawford, 1916. Insecutor Inscitiae Menstruus 4: 106. ¢.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 39-40, fig. 14 (redescription).

fremonti Cockerell. Oreg.
Stelis fremonti Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 213. ?.

montana Cresson. Alta., B. C., Wyo., Oreg., Colo., Utah, Ariz., Calif. Host: Osmia montana
Cress., O. texana Cress.
Stelis montana Cresson, 1864. Ent. Soc. Phila., Proc. 3: 39. °.
Biology: Rust, 1974. Wasmann Jour. Biol. 32: 80 (host).

seneciophila Cockerell. Colo.
Stelis seneciophila Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 329. 2.

Genus STELIS Subgenus CHELYNIA Provancher

Chelynia Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 322.
Type-species: Chelynia labiata Provancher. Monotypic.

Taxonomy: Cockerell, 1898. Entomologist 31: 166-167. —Cockerell, 1936. Canad. Ent. 68: 275
(Key to U.S. spp.). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 36 (tax. status).

calliphorina (Cockerell). Calif.
Chelynia calliphorina Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 769. °.
chlorocyanea (Cockerell). Calif. Host: Osmia nigrifrons Ckll. Parasite: Dibrachys sp.
Chelynia chlorocyanea Cockerell, 1925. Calif. Acad. Sci., Proe. (4) 14: 208. °.

Taxonomy: Rust and Thorp, 1973. Kans. Ent. Soc., Jour. 46: 548-562, 26 figs. 2 tables
(immature stages).

Biology: Rust and Thorp, 1973. Kans. Ent. Soc., Jour. 46: 548-562 (life history, hosts). —Rust,
Thorp and Torchio, 1974. Nat. Hist., Jour. 8: 44 (host).
cusackae (Cockerell). Colo.
Chelynia cusackae Cockerell, 1910. Ent. News 21: 270. 2.
depressa Timberlake. Calif.
Stelis (Chelynia) depressa Timberlake, 1941. N. Y. Ent. Soe., Jour. 49: 127. 9, d.
elegans Cresson. B. C., Colo., Ariz.
Stelis elegans Cresson, 1864. Ent. Soc. Phila., Proc. 2: 411. 5 (2 misdet.).
foederalis Smith. N. B. to Minn., south Ga. and Colo. Host: Hoplitis cylindrica (Cress.), Osmia
atriventris Cress.
Stelis foederalis Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 275. 2, d.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 45, figs. 12-13 (redescription).
fragariella (Cockerell). Calif.
Chelynia fragariella Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 209. d.
franciscana (Cockerell). Calif.
Chelynia franciscana Cockerell, 1925. Calif. Acad. Sci., Proce. (4) 14: 207. 2.
holocyanea (Cockerell). Calif.
Chelynia holocyanea Cockerell, 1925. Calif. Acad. Sci., Proe. (4) 14: 209. 2.

idahoensis Swenk. Idaho.
Stelis (Chelynia) idahoensis Swenk, 1914. Nebr. Univ., Studies 14: 6. °.

Superfamily APOIDEA 2003

Taxonomy: Cockerell, 1936. Canad. Ent. 68: 275 (tax. characters).

interrupta Cresson. Nev.
Stelis interrupta Cresson, 1897. Amer. Ent. Soc., Trans. 7: 205. °.

labiata (Provancher). Que. to Wis., south to N. C. and Tex.? Host: Hoplitis anthocopoides
(Schenck), H. cylindrica (Cress.), H. pilosifrons (Cress.), H. producta (Cress.), H.
simplex (Cress.).
Chelynia labiata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 322. 2, d.
¢Stelis (Microstelis) birkmanni Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 29. d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 40-42, fig. 13 (redescription,
synonymy).
Biology: Eickwort, 1973. Search, Cornell Univ. Agr. Expt. Sta. 3: 27 (host relationships).
leucotricha (Cockerell). Calif. Host: Anthidiwm mormonum Cress.
Chelynia leucotricha Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 208. 2, d.
Biology: Hicks, 1929. Ent. News 40: 108. (host).

linsleyi Timberlake. Calif.
Stelis (Chelynia) linsleyi Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 129. °.

melanotricha (Cockerell). Colo.
Chelynia melanotricha Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 626. 2.

michiganensis Mitchell. Mich. (Luce Co.).
Stelis michiganensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 45. °.

monticola Cresson. B. C., Idaho, Wyo., Colo. Host: Hoplitis fulgida Cress.
Stelis? monticola Cresson, 1878. Amer. Ent. Soc., Trans. 7: 94. 9.
Biology: Hicks, 1926. Colo. Univ., Studies 15: 243 (host).
nitida Cresson. Canada, N. Y., Wis., N. C.
Stelis? nitida Cresson, 1878. Amer. Ent. Soc., Trans. 7: 92. °.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 46-47 (redescription).
nitidula (Cockerell). Calif.
Chelynia nitidula Cockerell, 1925. Calif. Acad. Sei., Proc. (4) 14: 210. d.
Taxonomy: Cockerell, 1936. Canad. Ent. 68: 274 (tax. characters).
pavonina (Cockerell). Wyo., Colo. :
Chelynia pavonina Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 339. 6.
Taxonomy: Cockerell, 1922. Amer. Mus. Novitates 40: 7. d, °.
pulchra Crawford. Nebr., Colo.
Stelis pulchra Crawford, 1902. Canad. Ent. 34: 289. ?.
semirubra reducta Snelling. Calif. (Tuolumne and Mariposa counties).
Stelis (Chelynia) semirubra reducta Snelling, 1962. Pan-Pacifie Ent. 38: 227. 2, 6.
semirubra semirubra Timberlake. Calif.
Stelis (Chelynia) semirubra Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 127. °.
subcaerulea Cresson. Wyo., Nev., Calif.
Stelis? subcaerulea Cresson, 1878. Amer. Ent. Soc., Trans. 7: 938. ¢.
Taxonomy: Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 768. d, 2.
subemarginata Cresson. N. B., Ont., Man., Wis., N. Dak., Wyo., Idaho, Colo. Host: Hoplitis
albifrons (Kirby), H. cylindrica (Cress.), Osmia proxima Cress., O. simillima Sm.
Stelis? subemarginata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 9372
Taxonomy: Swenk, 1915. Nebr. Univ., Studies 15: 191. ¢. —Mitchell, 1962. N. C. Agr. Expt.
Sta. Tech. Bul. 152: 47 (redescription).

Biology: Graenicher, 1935. Ent. Soc. Amer., Ann. 28: 300 (host). —Fye, 1965. Canad. Ent. 97:
865 (hosts).

subglauca (Cockerell). Wash.
Chelynia subglauca Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 210. 6.

2004 Hymenoptera in America North of Mexico

Genus STELIS Subgenus MELANOSTELIS Ashmead

Melanostelis Ashmead, 1898. Psyche 8: 283.
Type-species: Chelynia rubi Cockerell. Monotypic and orig. desig. (=Melanostelis
betheli Ashmead).

ricardonis (Cockerell). B. C.
Chelynia ricardonis Cockerell, 1912. Canad. Ent. 44: 293. 2.

rubi Cockerell. B. C., Alta., Wash., Calif., Colo.
Stelis rubi Cockerell, 1898. Entomologist 31: 167. °.
Melanostelis betheli Ashmead, 1898. Psyche 8: 283. 2

Taxonomy: Swenk, 1914. Nebr. Univ. Studies 14: 8. 9, d.

Genus STELIS Subgenus STELIDIUM Robertson

Stelidium Robertson, 1902. Canad. Ent. 24: 323.
Type-species: Stelidium trypetinum Robertson. Monotypic and orig. desig.

ashmeadiellae Timberlake. Calif. Host: Ashmeadiella californica (Ashm.).
Stelis (Stelidium) ashmeadiellae Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 133. 2, d.
Biology: Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 134 (host).

nyssonoides (Brues). Tex.
Melanostelis nyssonoides Brues, 1903. Ent. News 14: 84. 2.

palmarum Timberlake. Calif.
Stelis (Stelidium) palmarwm Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 134. °.

permaculata Cockerell. Colo., N. Mex. Host: Heriades carinata Cress.?
Stelis lateralis var. permaculata Cockerell, 1898. Entomologist 31: 167. d.
Biology: Hicks, 1927. Ent. News 38: 297 (host).

robertsoni Timberlake. Calif.
Stelis (Stelidium) robertsoni Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 136. 2, ¢.

trypetina (Robertson). IIl., Ont., Va.
Stelidium trypetinum Robertson, 1902. Canad. Ent. 34: 323. 2.
Stelis (Stelidium) ontariana Sladen, 1916. Canad. Ent. 48: 312. 92, 6.

Taxonomy: Cockerell, 1922. Canad. Ent. 54: 143 (tax. characters). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 44-45 (redescription).

Genus STELIS Subgenus STELIDINA Timberlake

Stelis subg. Stelidina Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 125.
Type-species: Stelis hemirhoda Linsley. Orig. desig.
Stelidella(!) Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 133.
Revision: Linsley, 19389. Ent. News 50: 255.
acutiventris Linsley. Calif.
Stelis (Stelidium) acutiventris Linsley, 1939. Ent. News 50: 252. °.
cockerelli (Hicks). Calif.
Herbstiella cockerelli Hicks, 1933. Amer. Mus. Novitates 616: 1. °.

hemirhoda Linsley. Calif.
Stelis (Stelidium) hemirhoda Linsley, 1939. Ent. News 50: 250. °, d.

micheneri Linsley. Calif.

Stelis (Stelidium) micheneri Linsley, 1939. Ent. News 50: 253. 2, d.
nigriventris Timberlake. Calif.

Stelis (Stelidina) nigriventris Timberlake, 1941. N. Y. Ent. Soe., Jour. 49: 182. 2.

trichopyga Timberlake. Calif.
Stelis (Stelidina) trichopyga Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 180. 2.

Superfamily APOIDEA 2005

UNPLACED TAXON OF STELIS

The following species can neither be recognized as to subgenus nor associated with any of the
described species of Stelis in North America, but is a member of the genus Stelis.

Andrena nigrita Fabricius, 1775. Systema Ent., p. 377. “Habitat in America.”

NOMEN NUDUM IN STELIS PANZER

Stelis foxi Smith, 1910 (1909). N. J. State Mus., Ann. Rpt., p. 695.

Genus DIOXYS Lepeletier and Serville

Dioxys Lepeletier and Serville, 1825. Eneycel. Meth., Dict. Ins., v. 10, p. 109.
Type-species: Trachusa cincta Jurine. Monotypic.
Paradioxys Moesary, 1894. Termes. Fuzetek, v. 17, p. 35.
Type-species: Dioxys pannonica Mocsary. Monotypic.
Hoplopasites Ashmead, 1898. Psyche 8: 284.
Type-species: Phileremus? productus Cresson. Monotypic and orig. desig.
Chrysopheon Titus, 1901. Canad. Ent. 33: 256.
Type-species: Chrysopheon aurifuscus Titus. Monotypic.

Revision: Hurd, 1958. Calif. Univ. Pubs. Ent. 14: 275-302, 28 figs., 4 maps. (Nearctic spp.).

Taxonomy: Cockerell, 1928. Colo. Univ., Studies 16: 111 (synopsis). —Rozen, 1967. N. Y. Ent.
Soce., Jour. 75: 236-248, 31 figs. (immature stages).

Biology: Hurd, 1958. Calif. Univ. Pubs. Ent. 14: 276 (floral and host relationships).
aurifuscus (Titus). Colo., Idaho, Calif., Oreg. Host: Anthidium maculosum Cress.?,
Callanthidium illustre Cress.
Chrysopheon aurifuscus Titus, 1901. Canad. Ent. 33: 256. °.
Dioxys fulvohirta Ducke, 1909. Rev. de Ent. 27: 44. 2.

Taxonomy: Horning, 1966. Ent. Soc. Wash., Proc. 68: 157 (geogr. records).

Biology: Cockerell, 1909. Canad. Ent. 41: 334 (host). —Hicks, 1929. Canad. Ent. 61: 8 (host).
—Horning, 1971. Ent. Soe. Wash., Proc. 73: 43 (host).
pacificus melanogaster Hurd. South. Calif. (San Bernardino, San Gabriel and San Jacinto
Mts.).
Dioxys pacificus melanogaster Hurd, 1958. Calif. Univ. Pubs. Ent. 14: 296. 2, d.
pacificus pacificus Cockerell. Calif. (except mts. of southern Calif.), Oreg., Nev., Utah, Wyo.
Dioxys pacificus Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 286. °.
pomonae pomonae Cockerell. Idaho, Utah, Colo., N. Mex., Ariz., Oreg. and Calif. (except
Sonoran Desert). Host: Anthidium collectum Huard, Chalicodoma subewilis (Ckll.),
Osmia nifoata Ckll., O. nigrobarbata CkIl.
Dioxys pomonae Cockerell, 1910. Canad. Ent. 42: 169. d.
Dioxys phaceliae Cockerell, 1911. Amer. Ent. Soc., Trans. 37: 235. 2.
Dioxys catalinensis Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 148. ¢.

Taxonomy: Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 285. 2. —Gittins, 1959. Brooklyn
Ent. Soc., Bul. 54: 135 (geogr. records). —Horning, 1966. Ent. Soc. Wash., Pro¢. 68: 157
(floral and geogr. records). —Rozen, 1967. N. Y. Ent. Soc., Jour. 75: 236-248, 31 figs.
(immature stages).

Biology: Rozen and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 197-203, 7 figs. (life history,
host).
pomonae timberlakei Hurd. Southern Calif., deserts.
Dioxys pomonae timberlakei Hurd, 1958. Calif. Univ. Pubs. Ent. 14: 285. 2, d.
productus cismontanicus Hurd. Pacific slope of Calif.; Mexico (Baja California). Host:
Anthidium collectum Huard.
Dioxys productus cismontanicus Hurd, 1958. Calif. Univ. Pubs. Ent. 14: 290. 9, d.
productus productus (Cresson). Great Basin of Calif., Nev., Wash., Oreg., Utah. Host:
Anthidium utahense Swenk.
Phileremus? productus Cresson, 1879. Amer. Ent. Soc., Trans. 7: 203. 2.

2006 Hymenoptera in America North of Mexico

Taxonomy: Ducke, 1909. Rev. de Ent. 27: 44 (tax. characters). —Horning, 1966. Ent. Soe.
Wash., Proc. 68: 157 (geogr. record). —Rozen, 1967. N. Y. Ent. Soc., Jour. 75: 2, 36, 238-239,
240, 242, figs. 9-15 (larva, as productus productus?).

Biology: Jaycox, 1966. Pan-Pacific Ent. 42: 18-20 (habits, host).
productus subruber (Cockerell). Calif. deserts, Ariz., N. Mex., western Tex. Host: Anthidiwm
paroselae Ckll.?
Phileremus productus var. subruber Cockerell, 1898. N. Mex. Univ., Bul. 1: 60. 2.
Dioxys marti Cockerell, 1902. Ann. and Mag. Nat. Hist. (7) 9: 233. 2 (= male).

Biology: Newberry, 1900. Psyche 9: 94 (host).

rohweri Cockerell. Colo. (Troublesome).
Dioxys rohweri Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 329. 6.

TRIBE MEGACHILINI

The tribe Megachilini, which contains the well known leaf-cutter and mason bees, is
represented by numerous species throughout much of the world. Although some of these bees
are cleptoparasites, mainly in the nests of other Megachilini, most members of this tribe are pol-
len-collecting bees, and like the Anthidiini make use of a wide variety of foreign materials for
the construction of the cells in their nests. However, unlike the Anthidiini some of these bees
utilize mud for cell construction, but none is known to use plant down for this purpose. Some of
the species live in colonies (¢.g., certain Chalicodoma and Osmia) which are communal and even
possibly quasisocial, but the majority of the species are solitary. Most of the species appropriate
a wide variety of pre-existing holes and cavities of all sorts in which to make their nests, but
some of them do excavate their own tunnels in the ground. In some classifications the tribe
Megachilini is divided into two subtribes, the Osmiini (arolium present) and the Megachilini
(arolium absent).

Taxonomy: Michener, 1941. Amer. Midland Nat. 26: 147-167 (partial generic revision).
—Michener, 1944. Amer. Nat. 78: 257-266, 1 fig. (distribution of osmiine bees of the N.
Amer. deserts). —Hurd and Michener, 1955. Calif. Ins. Survey Bul. 3: 1-248, 141 figs., 112
maps (Calif. spp. except of Osmia, Megachile, and Coelioxys). —Michener and Sokal, 1957.
Evolution 11: 130-162, 15 figs., 4 tables (genera of Hoplitis complex). —Sokal and
Michener, 1958. Kans. Univ. Sci. Bul. 38: 1409-1438 (statistical evaluation of systematic
relationships in the Hoplitis complex). —Michener, 1962. N. Y. Ent. Soc., Jour. 70: 17-29
(classification of bees commonly placed in the genus Megachile). —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 152: 48-232, figs. 1, 15-66, tables 3-7 (eastern U. S. spp.).
—Rubin, 1966. Syst. Zool. 15: 169-182, 3 tables (homogeneous groups in the Hoplitis
complex). —Michener and Sokal, 1966. Ent. Soc. Amer., Ann. 59: 1211-1217, 5 figs.
(phenetic similarities in the Hoplitis complex). —Sokal and Michener, 1967. Linn. Soc.
London, Proe. 178: 59-74 (effects of different numerical techniques on the phenetic
classification of the Hoplitis complex).

Biology: Eickwort, 1975. Evolution 29: 142-150, 1 fig., 2 tables (evolution of parasitism and
sociality in Hoplitis).

Genus HERIADES Spinola

The bees of this genus nest in small holes, such as those made by emerging beetles in old logs,
dead branches, and pine cones. Insofar as known, the species are polylectic and have rather long
seasons of flight.

Revision: Michener, 1938. Ent. Soc. Amer., Ann. 31: 514-531 (Amer. spp.). —Hurd and
Michener, 1955. Calif. Ins. Survey, Bul. 3: 11-24, pls. 3, 12, figs. 7-16, maps 2-7 (Calif. spp.).
—Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 48-52, figs. 1, 15, table 3 (eastern U.
S. spp.).

Genus HERIADES Subgenus HERIADES Spinola

Heriades Spinola, 1808. Insectorum Liguriae, v. 2, p. 7.
Type-species: Apis truncorum Linnaeus. Desig. by Latreille, 1810.
Trypetes Schenck, 1859. Nassau. Ver. f. Naturk. Jahrb. 14: 32. Preoce.

Superfamily APOIDEA 2007

Type-species: Apis truncorum Linnaeus. Monotypice.

This subgenus is not found in America.
Genus HERIADES Subgenus NEOTRYPETES Robertson

Neotrypetes Robertson, 1918. Ent. News 29: 92.
Type-species: Megachile variolosa Cresson. Monotypic and orig. desig. (=Trypetes
productus Robertson).

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 68-69 (rev. key to spp.).
crucifera Cockerell. N. Mex., Ariz.
Heriades crucifera Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 137. ¢.
leavitti Crawford. N. B. and Maine, south to Fla., west to Tex. and Wyo. Pollen: Polylectie,
visits a wide variety of flowers including Anthemis cotulla, Bidens, Chrysopsis,
Erigeron, Galax, Helenium, Ilex, Jussiaea, Melilotus officinalis, Oenothera, Polygonum,
Senecio, Solidago, Stokesia.
Heriades leavitti Crawford, 1913. Canad. Ent. 45: 270. 6.
Heriades crawfordi Graenicher, 1928. Ent. News 39: 281. 2, 6.
*Neotrypetes truncatus Robertson, 1929. Flowers and Insects, p. 131. 2, d.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 278-279 (nest architecture, life
history).
micheneri Timberlake. Ariz.
Heriades (Neotrypetes) micheneri Timberlake, 1947. Pan-Pacific Ent. 23: 26. 6, °.
micropthalma Michener. Deserts of west. Tex., N. Mex., Ariz., Utah. Pollen: Apparently
polylectic, visits flowers of Melilotus officinalis, Monarda, Petalostemon, Tetradymia
canescens.
Heriades micropthalma Michener, 1954. Kans. Ent. Soc., Jour. 27: 66, 2 figs. 2, d.
texana Michener. Tex., Ariz.
Heriades terana Michener, 1938. Ent. Soc. Amer., Ann. 31: 517. °.
variolosa purpurascens Cockerell. South. Tex. to Yucatan.
Heriades carinata purpurascens Cockerell, 1931. Ann. and Mag. Nat. Hist. (10) 8: 543. °.

Taxonomy: Michener, 1949. Kans. Ent. Soc., Jour. 22: 43.
variolosa variolosa (Cresson). Ont. and Maine, south to Fla., west to B. C., Wash., Oreg., Utah,

Colo., N. Mex. and Tex; Mexico (northern). Ecology: Nests in twigs of sumac. Pollen:
Polylectic, principally Compositae, but visits a wide variety of flowers including
Amorpha, Apocynum, Asclepias, Aster, Berteroa, Bidens, Blephilia, Brassica,
Ceanothus, Cirsium, Coreopsis, Erigeron philadelphicus, Grindelia, Gutierrezia
sarothrae, Helianthus, Heliopsis helianthoides, Houstonia, Malva silvestris, Melilotus
alba, Mentha canadensis, Monarda, Parthenium, Prunus, Ratibida, Senecio, Solidago
rigida, Thelesperma gracile, Toxicodendron, Trifolium, Vernonia.

Megachile variolosa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 270. °.

Heriades odontophora Schletterer, 1889. Zool. Jahrb., Abt. f. System., Geog. u. Biol. Tiere
4: 679. 9.

Heriades asteris Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 135. ¢.

Trypetes barbatus Robertson, 1903. Amer. Ent. Soc., Trans. 29: 171. ¢ (2 misdet.).

Trypetes productus Robertson, 1905. Canad. Ent. 37: 236. d.

Biology: Fischer, 1955. Canad. Ent. 87: 33-36 (nest).
Genus HERIADES Subgenus PHYSOSTETHA Michener

Heriades subg. Physostetha Michener, 1938. Ent. Soe. Amer., Ann. 31: 523.
Type-species: Heriades carinata Cresson. Orig. desig.
carinata Cresson. N. B. and Que. to Fla., west to B. C., Oreg., Utah, N. Mex., and Tex.
Parasite: Leucospis affinis Say, Melittobia chalybui Ashm., Sapyga louwist Krombein,
Stelis permaculata Ckll., S. vernalis Mitchell. Pollen: Polylectic, visits a wide variety of
flowers including Apocynum, Aronia, Asclepias syriaca, Cichorium intybus, Cleome

2008 Hymenoptera in America North of Mexico

serrulata, Erigeron, Fayopyrum, Helianthus, Ilex, Medicago sativa, Melilotus alba, M.
officinalis, Mentha canadensis, Monarda fistulosa, Parthenocissus quinquefolia,
Penstemon, Petalostemon candidus, Polygonum scandens, Potentilla, Rhus glabra, R.
typhina, Rubus, Senecio, Solidago, Trifolium, Vicia.

Heriades carinatum Cresson, 1864. Ent. Soc. Phila., Proc. 2: 388. 2, ¢.

Heriades glomerans Schletterer, 1889. Zool. Jahrb., Abt. f. System., Geog. u. Biol. Tiere 4:
681. 2.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7): 39-40 (nest). —Matthews, 1965. Amer.
Ent. Inst. Contrib. 1: 1-33, 23 figs., 7 tables (nest, life history, immature stages, parasites).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 279-282 (nest architecture, life
history, supersedure).

cressoni Michener. Nebr. to N. Mex., west to B. C. and Calif. Pollen: Polylectic, visits a wide
variety of flowers including Achillea lanulosa, Aster adscendens delectabilis, A.
canescens, A. parishii, Calycadenia multiglandulosa, Chrysopsis, Chrysothamnus
pumilus, C. viridulus, Chamaenerion angustifolium, Cirsium californicum, Erigeron
foliosus var. stenophyllus, Eriogonum elongatum, E. latifoliwm var. nudum,
Eriophyllum confertiflorum, Gayophytum ramosissimum, Geranium richardsoni,
Gutierrezia californica, Haplopappus gracilis, Helenium bigelovii, Heteromeles
arbutifolia, Potentilla gracilis, P. glandulosa, Senecio ionophyllus, Solidago californica,
S. confinis, S. petradoria.
Heriades (Physostetha) cressoni Michener, 1938. Ent. Soc. Amer., Ann. 31: 529. ?, 6.
gracilior Cockerell. Colo., N. Mex., Ariz., Nev. and south. Calif. Pollen: Unknown, but visits
flowers of Convolvulus arvensis, Opuntia.
Heriades gracilior Cockerell, 1897, Ann. and Mag. Nat. Hist. (6) 20: 1388. 6, 2.
occidentalis Michener. Oreg., Calif. Ecology: Nests in abandoned beetle burrows in cones of
Pinus attenuata. Pollen: Polylectic, visits a wide variety of flowers including
Adenostoma fasciculatum, Angelica tomentosa, Asclepias fascicularis, Baccharis
douglasti, Cirsium vulgaris, Cordylanthus, Cryptantha intermedia, Eriogonum
fasciculatum, Eriogonum latifolium var. nudum, Eriodictyon trichocalyx, Eriophyllum
lanatum, Eryngium aristatulum, Gnaphalium thermale, Haplopappus squarrosus,
Helianthum scoparius, Helianthus gracilentus, Hemizonia paniculata, Heteromeles
arbutifolia, Lonicera interrupta, Lotus, Nolina parryi, Opuntia occidentalis, Phacelia
imbricata, Rhus integrifolia, Sidalcea malvaeflora, Solidago californica, S. occidentalis,
Stephanomeria, Swertia parryi, Tetradymia canescens, Verbena lasiostachys.
Heriades (Physostetha) occidentalis Michener, 1938. Ent. Soc. Amer., Ann. 31: 525. 9, 6.
timberlakei Michener. Ariz., Nev., N. Mex., Colo. Pollen: Unknown, but visits flowers of
Euphorbia albomarginata.
Heriades (Physostetha) timberlakei Michener, 1938. Ent. Soc. Amer., Ann. 31: 527. 6, @.

Genus PROCHELOSTOMA Robertson

Prochelostoma Robertson, 1903. Amer. Ent. Soc., Trans. 29: 167.
Type-species: Heriades philadelphi Robertson. Monotypic and orig. desig.

Revision: Michener, 1938. Ent. News 69: 131.
philadelphi (Robertson). N. Y. to Ga., west to Mich., Ill., Kans. and Miss. Ecology: Nests in
deserted anobiid borings and trap-nests. Parasite: Melittobia chalybii Ashm., Pyemotes
ventricosus (Newport). Pollen: Unknown, but visits flowers of Capsella, Crataegus,
Ellisia, Geraniwm, Hydrophyllum, Ilex, Philadelphicus, Rubus. Predator: Trogoderma

ornatum Say.
Heriades philadelphi Robertson, 1891. Amer. Ent. Soc., Trans. 18: 64. 2, 6.

Biology: Krombein, 1959. Ent. News 70: 135-136 (nest, mating). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 276-278, pl. 17, fig. 86 (nest architecture, life history,
supersedure, parasites).

Genus CHELOSTOMA Latreille
Chelostoma Latreille, 1809. Gen. Crust. Ins., v. 4, p. 16.

Superfamily APOIDEA 2009

Type-species: Apis florisomnis Linnaeus. Monotypic. (=Apis mawillosa Linnaeus).
Gyrodroma Thomson, 1872. Hym. Scand., v. 2, p. 259. Preoce.
Type-species: Heriades nigricornis Nylander. Desig. by Cockerell, 1925.

Nearctic species of this genus visit flowers of Hydrophyllaceae, especially the genera
Phacelia and Eriodictyon, and possibly collect pollen exclusively from these flowers.

Revision: Michener, 1938. Pan-Pacific Ent. 14: 36-45.
bernardinum Michener. South. Calif. Pollen: Unknown, but visits flowers of Nemophila
integrifolia, N. menziesii, N. spatulata, Phacelia davidsonii.
Chelostoma bernardinum Michener, 1938. Pan-Pacific Ent. 14: 40. 2, 3.
californicum Cresson. Calif.; Mexico (Baja California). Pollen: Apparently oligolectic on
flowers of Phacelia including P. ciliata, P. davidsonii, P. distans, P. hispida, P.
imbricata, P. platyloba, P. ramosissima, but visits other flowers presumably for nectar
including Cryptantha intermedia, C. flaccida, Eriodictyon californicum, E. trichocalyx,
Hesperochiron californicum, Layia platyglossa, Nemophila menziesii, N. integrifolia,
Plagiobothrys nothofulvus, Salvia carnosa.
Chelostoma californicum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 108. d.
Heriades albicinctum Provancher, 1895. Nat. Canad. 22: 190. 2, 6.
Heriades odontura Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 139. 6.
Robertsonella dolichosoma Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 456. 2.

cockerelli Michener. South. Calif. Pollen: Apparently oligolectic on flowers of Eriodictyon
including EF. californicum, E. crassifolium, E. trichocalyx, but visits other flowers
including Clarkia, Cryptantha intermedia, Gilia exilis, Lotus scoparius, Mimulus
fremontii, Nama parryi, Navarretia heterodoxa, Penstemon, Phacelia distans, P.
tanacetifolia, Trichostema lanatum, T. parishii, Verbena lasiostachys, Viguiera
deltoides.
Chelostoma minutum cockerelli Michener, 1938. Pan-Pacific Ent. 14: 48. 6, 9.
incisulum Michener. Calif. Pollen: Apparently oligolectic on flowers of Phacelia including P.
distans, P. platyloba, but also visits flowers of Ceanothus, Cryptantha, Gilia tricolor,
Hesperochiron californicus, Lasthenia chrysostoma, Linanthus aureus.
Chelostoma minutum incisulum Michener, 1938. Pan-Pacifie Ent. 14: 44. d.

marginatum incisuloides Michener. Cent. Calif. Pollen: Apparently oligolectic on flowers of
' Phacelia including P. platyloba, but also visits other flowers including Amsinckia,
Cryptantha, Eriodictyon californicum, Nemophila pulchella, Penstemon laetus.
Chelostoma marginatum incisuloides Michener, 1954. Kans. Ent. Soc., Jour. 27: 69. 6, @.
marginatum marginatum Michener. South Calif. Pollen: Apparently oligolectic on flowers of
Phacelia including Phacelia davidsonii, P. distans, P. hispida, P. minor, but also visits
other flowers including Allium parvum, Cryptantha intermedia, Eriodictyon
crassifolium, Lomatium dasycarpum, Nemophila pulchella, Rhamnus crocea, Rhus
trilobata, Salvia columbariae, S. mellifera.
Chelostoma minutum marginatum Michener, 1938. Pan-Pacific Ent. 14: 44. 6, 9.
minutum Crawford. Wash., Oreg., Utah, Nev., Calif. (Sierra Nevada and San Bernardino Mts.),
Idaho. Pollen: Presumably oligolectic on Phacelia including P. californica, P. davidsonii,
P. distans, P. heterophylla, P. imbricata, P. linearis, P. ramosissima, but also visits
other flowers including Alliwm parvum, Calochortus, Cryptantha, Draperia systyla,
Erigeron divergens, Eriodictyon crassifolium, E. trichocalyx, Eriophyllum lanatum,
Erysimum asperum, Heuchera micrantha, Lepidium virginicum, Lotus argophyllus,
Mimulus fremontii, Nemophila menziesii, Penstemon spectabilis, Polemonium,
Potentilla glandulosa, Sidalcea malvaeflora.
Chelostoma minuta Crawford, 1916. Insecutor Inscitiae Menstruus 4: 102. °.
phaceliae Michener. Wash., Oreg., Calif., Nev. Pollen: Apparently oligolectic on Phacelia
including P. californica, P. distans, P. heterophylla, P. hispida P. imbricata, P. linearis,
P. ramosissima, P. tanacetifolia, but also visits other flowers including Allium
dichlamydium, Brodiaea laxa, Calochortus, Clarkia dudleyana, C. pulchella, C.
rhomboidea, Cryptantha intermedia, Erigeron foliosus var. stenophyllus, Eriodictyon
californicum, E. crassifolium, Gilia exilis, Horkelia bolanderi, Lepechinia calcycina,

2010 Hymenoptera in America North of Mexico

Lotus scoparius, Mimulus fremonti, Monardella lanceolata, Nama parryi, Nasturtium
officinale, Polemonium, Potentilla glandulosa, Sidalcea malvaeflora.
Chelostoma phaceliae Michener, 1938. Pan-Pacifie Ent. 14: 38. 3, 2
tetramerum Michener. North. Calif. Pollen: Unknown, but visits flowers of a yellow Mimulus.
Chelostoma tetramerum Michener, 1942. Ent. News 53: 47. 6.

Genus CHELOSTOMOPSIS Cockerell

Chelostomopsis Cockerell, 1925. Calif. Acad. Scei., Proc. (4) 14: 205.
Type-species: Chelynia rubifloris Cockerell. Orig. desig.
Raphidostoma Cockerell, 1936. Pan-Pacifie Ent. 12: 133.
Type-species: Chelynia rubifloris Cockerell. Monotypic and orig. desig.
(=Raphidostoma ceanothi Cockerell).

Revision: Michener, 1938. Ent. News 49: 127.

Taxonomy: Popov, 1961. Zool. Zhur. Akad. Nauk 40: 359-370, 3 figs. (tax. status, tax,
relationships).
rubifloris (Cockerell). Wash., Oreg., Calif., Ariz. Ecology: Has been reared from cones of

Pinus attenuata. Pollen: Polylectic, visits a wide variety of flowers including Alliwm
parvum, Agoseris, Arbutus menziesii, Arctostaphylos drupacea, A. mariposa, A.
nevadensis, A. patula, Arenaria douglasii, Calochortus luteus, C. nudus, Castilleia,
Ceanothus cyneatus, C. integerrimus, C. parviflorus, Cercocarpus ledifolius,
Chamaebatia foliolosa, Chorizanthe staticoides, Clarkia unguiculata, Collinsia bicolor,
Convolvulus occidentalis, Cryptantha denticulata, C. intermedia, C. lepida, Delphinium,
Eriastrum, Eriodictyon californicum, Frasera californica, Fragaria californica, Gilia
exilis, G. tricolor, Grindelia, Haplopappus linearifolius, Helianthus, Hesperochiron,
Horkelia, Lepechinia, calycina, Limnanthes douglasii, Lotus argophyllus, L.
corniculatus, L. davidsonii, L. glaber, L. scoparius, Lupinus bicolor, L. micranthus, L.
nanus, Melilotus, Mimulus fremontii, M. primuloides, M. suksdorfii, Montia perfoliata,
Nama parryi, Navarretia heterodoxa, Nemophila exilis, N. heterophylla, N. integrifolia,
N. maculata, N. menziesii, Penstemon breviflorus, P. heterophyllus, Phacelia
brachyloba, P. californica, P. davidsonii, P. heterophylla, Plagiobothrys nothofulvous,
Plectritis ciliosa, Potentilla glandulosa, Prunus ilicifolia, P. subcordata, Radicula
nasturtium- -aquaticum, Ranunculus, Rhamnus californica, R. crocea, Rhus trilobata,
Ribes roezlii, Salvia carnosa, S. columbariae, S. mellifera, Satureja douglasii, Sedum
stenopetalum, Senecio, Stenotopsis linearifolius, Streptanthus tortuosus, Thysanocarpus
curvipes, Trifolium gracilentum, T. melananthum, T. microcephalum, T. tridentatum,
Verbena lasiostachys, V. prostrata, Viola pedunculata, V. purpurea, Wyethia.

Chelynia rubifloris Cockerell, 1898. Canad. Ent. 30: 50. °.

Chelostoma rubifloris edwardsii Cockerell, 1916. Entomologist 49: 157. 2.

Raphidostoma ceanothi Cockerell, 1936. Pan-Pacific Ent. 12: 134. 6.

Genus PROTERIADES Titus

The bees of this genus appear to collect pollen almost exclusively from flowers of the
boraginaceous genus Cryptantha. Some of the species make their nests in the ground while
others construct them in preexisting holes in wood above ground.

Revision: Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 387-440, 16 figs.

Taxonomy: Hurd and Michener, 1955. Calif. Ins. Survey, Bul. 3: 129-153, pls. 9, 21-22, figs.
92-111, maps 63-76 (Calif. spp.). —Michener and Sokal, 1957. Evolution 11: 158 (subgeneric
assignments). —Rubin, 1966. Syst. Zool. 15: 176-182, tables 1-3 (phenetic classification).
—Michener and Sokal, 1966. Ent. Soc. Amer., Ann. 59: 1211-1217, 5 figs. (phenetic
similarities among the species of the Hoplitis complex).

Genus PROTERIADES Subgenus CEPHALAPIS Cockerell

Chelostoma subg. Cephalapis Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 23.
Type-species: Chelostoma (Cephalapis) jacintanum Cockerell. Monotypic and orig.
desig.

Superfamily APOIDEA 2011

jacintana (Cockerell). Calif., Oreg. Pollen: Oligolege of Cryptantha including C. intermedia, C.
micrantha, C. m. var. lepida, C. muricata var. denticulata, but also visits flowers of

Ceanothus and Eriogonum for nectar.
Chelostoma jacintanum Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 23. 6.

Genus PROTERIADES Subgenus PROTERIADES Titus

Proteriades Titus, 1904. N. Y. Ent. Soe., Jour. 12: 25.
Type-species: Heriades semirubra Cockerell. Monotypic.
basingeri Timberlake and Michener. Calif. (Riverside). Pollen: Cryptantha intermedia.
Proteriades basingeri Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 413. 6, °.
bidenticauda Timberlake and Michener. Calif. (Inyo and Kern Cos.). Pollen: Cryptantha sp.,
but also visits flowers of Mentzelia presumably for nectar.
Proteriades bidenticauda Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 420.,
figss4 epee
boharti Timberlake and Michener. Calif. (Carrville, Trinity Co.).
Proteriades boharti Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 429, figs. 2,
14. d.
caudex Timberlake and Michener. Calif. (San Jacinto Mts.). Pollen: Oligolege of Cryptantha
intermedia, C. lepida, C. micrantha.
Proteriades caudex Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 418, figs. 3,
155)
cryptanthae Timberlake and Michener. South. Calif. Pollen: Cryptantha intermedia.
Proteriades cryptanthae Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 416. 6,
oF

deserticola Timberlake and Michener. South. Calif. Ecology: Nests in burrows in ground.
Pollen: Cryptantha angustifolia, C. barbigera.
Proteriades deserticola Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 430. d, °.
evansi Michener. Calif. (Sierra Nevada Mts.). Pollen: Cryptantha flaccida.
Proteriades evansi Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 92. °.
hamulicornis Timberlake and Michener. Calif. (Inyo Co.). Pollen: Cryptantha spp.
Proteriades hamulicornis Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 425,
figs. 5, 13. 5, @.
nanula nanula Timberlake and Michener. South. Calif. lowlands. Pollen: Cryptantha
intermedia.
Proteriades nanula nanula Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 409.
Oy Qe
nanula sparsa Timberlake and Michener. Centr. Calif., south. Calif. mts. Pollen: Cryptantha
angustifolia, C. flaccida, C. micrantha var. lepida.
Proteriades nanula sparsa Timberlake and Michener, 1950. Kans. Univ. Sci. Bub. 33: 410.
Sy Ie
nigrella attonita Michener. Calif. (Inyo Co.). Pollen: Cryptantha racemosa.
Proteriades nigrella attonita Michener, 1954. Kans. Ent. Soc., Jour. 27: 73. 2, ¢.
nigrella nigrella Michener. South. Calif. Pollen: Cryptantha angustifolia.
Proteriades nigrella nigrella Michener, 1954. Kans. Ent. Soc., Jour. 27: 73. 2.
palmarum (Cockerell). South. Calif. Pollen: Cryptantha angustifolia, C. barbigera.
Osmia palmarum Cockerell, 1935. Pan-Pacific Ent. 11: 48. °.
Proteriades nigra Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 405. °.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 71. 6. —Hurd and Michener, 1955.
Calif. Ins. Survey, Bul. 3: 139 (synonymy).
pygmaea Timberlake and Michener. South. Calif. Pollen: Oligolege of Cryptantha including C.
angustifolia, C. barbigera, but also visits flowers of Eriogonwm and Nama demissum.
Proteriades pygmaea Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 411, figs. 6,
I, Gin Se
reducta Timberlake and Michener. Calif. (Riverside Co.). Pollen: Cryptantha intermedia.
Proteriades reducta Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 414. d, °.

2012 Hymenoptera in America North of Mexico

seminigra seminigra Timberlake and Michener. South. Calif. Pollen: Cryptantha intermedia.
Proteriades seminigra seminigra Timberlake and Michener, 1950. Kans. Univ. Sei. Bul. 33:
403. 3, 2.
seminigra yosemitensis Timberlake and Michener. Cent. Calif. Pollen: Cryptantha sp.
Proteriades seminigra yosemitensis Timberlake and Michener, 1950. Kans. Univ. Sci. Bul.
33: 404. d, 2.
semirubra (Cockerell). Coastal south. and cent. Calif. Pollen: Cryptantha intermedia, but also
visits Eriophyllum multicaule for nectar.
Heriades semirubra Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 198. ¢.
shoshone Parker. Nev. (Elko County). Ecology: Nests in elderberry trap-stems. Parasite:
Stelis coarctatus Cwfd. Pollen: Stores pollen of Cryptantha.
Proteriades shoshone Parker, 1976. Pan-Pacific Ent. 52: 73, figs. 1-8. 6, 2.

Biology: Parker, 1976. Pan-Pacifie Ent. 52: 75-79, figs. 9-14 (nest, sex ratio, cocoon,
supersedure, parasite).
similis Timberlake and Michener. South. Calif. Pollen: Cryptantha intermedia.
Proteriades similis Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 406. 2.
tristis Michener. South. Calif. Pollen: Cryptantha intermedia, C. micrantha, C. muricata.
Proteriades tristis Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 92. °.

truicauda Timberlake and Michener. Calif. (San Jacinto Mts.). Pollen: Cryptantha micrantha.
Proteriades truicauda Timberlake and Michener, 1950. Kans. Univ. Sci. Bul. 33: 428. d, °.

Genus PROTERIADES Subgenus XEROSMIA Michener

Anthocopa subg. Xerosmia Michener, 1943 Ent. Soc. Amer., Ann. 36: 81.
Type-species: Osmia xerophila Cockerell. Orig. desig.
xerophila (Cockerell). South. Calif. Ecology: Reared from old nests of Anthophora linsleyi
Timberlake dug from ground. Parasite: Nemognatha scutellaris Say. Pollen: Oligolege of
Cryptantha including C. barbigera, but also visits flowers of Larrea tridentata and
Malva parviflora for nectar. Predator: Cymatodera.
Osmia xerophila Cockerell, 1935. Pan-Pacifie Ent. 11: 45. 2, d.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1055, figs. 146-148, 150 (larva).
Biology: Linsley and MacSwain, 1942. Amer. Midland Nat. 27: 408 (nest).

Genus PROTERIADES Subgenus PENTERIADES Michener and Sokal

Proteriades subg. Penteriades Michener and Sokal, 1957. Evolution 11: 158.
Type-species: Osmia remotula Cockerell. Orig. desig.
incanescens incanescens (Cockerell). Ariz.
Hoplitina incanescens Cockerell, 1922. Amer. Mus. Novitates 40: 7. °.
incanescens nevadensis Timberlake and Michener. S. Dak., Wyo., Nev., Calif. Pollen:
Cryptantha bradburiana.
Proteriades incanescens nevadensis Timberlake and Michener, 1950. Kans. Univ. Sci. Bul.

33: 399. 2.
Taxonomy: Michener, 1954. Kans. Ent. Soc. Jour. 27: 71. ¢. —LaBerge, 1973. Ent. News 84:
160. (geog. and floral recs.). —Tepedino, 1974. Ent. News 85: 146 (geogr. record).

incanescens tota Michener. Calif. (Tuolumne Co.).
Proteriades incanescens tota Michener, 1954. Kans. Ent. Soc., Jour. 27: 72. 2.

remotula (Cockerell). Calif. Ecology: Observed entering hole in the ground. Pollen: Oligolege
of Cryptantha including C. intermedia, C. micrantha, but also visits flowers of
Asclepias and Eriophyllum confertiflorum for nectar.
Osmia remotula Cockerell, 1910. Canad. Ent. 42: 170. @.

Taxonomy: Snelling, 1962. Pan-Pacific Ent. 38: 228-229 (geogr. and floral records).
Genus PROTERIADES Subgenus HOPLITINA Cockerell
Hoplitella Cockerell, 1910. Canad. Ent. 42: 169. Preoce.

Superfamily APOIDEA J 2013

Type-species: Ashmeadiella howardi Cockerell. Monotypic. (=Hoplitella pentamera
Cockerell).
Hoplitina Cockerell, 1913. Canad. Ent. 45: 34. Proposed to replace Hoplitella Cockerell.
bullifacies (Michener). East. Calif., desert. Parasite: Anthrax irroratus Say, Leucospis affinis
Say, Sphaeropthalma sp., Stelis sp. Pollen: Presumably Cryptantha, but is known to
visit flowers of Phacelia including P. aff. fremontii. Predator: Trichodes oriatus Say.
Hoplitis (Hoplitina) bullifacies Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 274. 2 (6
misdet.).

Taxonomy: Michener, 1954. Pan-Pacific Ent. 30: 37-38. d, °.
bunocephala (Michener). Cent. Calif. Parasite: Chrysura pacifica (Say). Pollen: Unknown, but
visits flowers of Lotus including L. subpinnatus.
Hoplitis (Hoplitina) bunocephala Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 276. °,
3.

howardi (Cockerell). South. and cent. Calif. Parasite: Stelis sp. Pollen: Unknown, but appears
to visit principally the flowers of Lotus including L. glaber, L. scoparius, L. strigosus,
but has also been taken at flowers of Cryptantha intermedia, Eriophyllum
confertiflorum, and Salvia.
Ashmeadiella howardi Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 22. 2 (¢ misdet.).
Hoplitella pentamera Cockerell, 1910. Canad. Ent. 42: 169. ¢.
Hoplitina hesperia Crawford, 1916. Insecutor Inscitiae Menstruus 4: 103. °.
linsdalei (Michener). Cent. Calif.
Hoplitis (Hoplitina) linsdalei Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 277. d.
mazourka (Michener). Calif. (Inyo Co.). Pollen: Unknown, but visits flowers of Cryptantha,
Gilia, Tetradymia.
Hoplitis mazourka Michener, 1954. Pan-Pacific Ent. 30: 38. d.
mojavensis (Michener). South. Calif. Pollen: Unknown, but visits flowers of Cryptantha
barbigera, Lasthenia chrysostoma, Nama demissum, Phacelia fremontii.
Hoplitis (Hoplitina) mojavensis Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 275. 2.

Genus PROTERIADES Subgenus ACROSMIA Michener

Hoplitis subg. Acrosmia Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 298.
Type-species: Hoplitis plagiostoma Michener. Orig. desig.
laevibullata (Michener). Calif., Utah. Pollen: Unknown, but visits flowers of Calyptridium
umbellatum, Nemophila, Phacelia.
Anthocopa (Eremosmia) laevibullata Michoner! 1943. Ent. Soc. Amer., Ann. 36: 68. 2.
Hoplitis (Acrosmia) perissocera Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 299. ¢.

Taxonomy: Michener, 1954. Pan-Pacific Ent. 30: 40 (synonymy).
plagiostoma (Michener). Calif. (Sierra Nevada Mts.), Oreg.
Hoplitis (Acrosmia) plagiostoma Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 298. 6.
rufina (Michener). Calif., Oreg., Utah. Pollen: Unknown, but visits flowers of Nemophila, Viola

purpurea.
Hoplitis rufina Michener, 1954. Pan-Pacifie Ent. 30: 40. 2, d.

Genus HOPLITIS Klug

Revision: Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 261-317 (N. Amer. spp. with
references to biological works).

Taxonomy: Hurd and Michener, 1955. Calif. Ins. Survey, Bul. 3: 44-94, pls. 7, 14-18, figs. 5,
51-70, maps 17-39 (Calif. spp.). —Michener and Sokal, 1957. Evolution 11: 130- 162 (phenetic
classification of the Hoplitis complex). —Sokal and Michener, 1958. Kans. Univ. Sci. Bul.
38: 1409-1438 (systematic relationships within the Hoplitis complex). —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 56-68, figs. 19-25, table 3 (eastern U. S. spp.).
—Michener, 1966. Entomologist 99: 146 (status of Anthocopa and Hoplitis). Tales 1966.
Syst. Zool. 15: 176-182, tables 1-3 (phenetic classification of the Hoplitis complex). *
—Michener and Sokal, 1966. Ent. Soc. Amer., Ann. 59: 1211-1217, 5 figs. (phenetic

2014 Hymenoptera in America North of Mexico

similarities among the species of the Hoplitis complex). —Michener, 1968. Ent. South.
Africa, Jour. 31: 337 (status of Anthocopa and Hoplitis).

Genus HOPLITIS Subgenus HOPLITIS Klug

Hoplitis Klug, 1807. Mag. Insektenk. 6: 225.
Type-species: Apis adunca Panzer. Monotypic.
Osmia subg. Ctenosmia Thomson, 1872. Hym. Scand., v. 2, p. 233.
Type-species: Apis adunca Panzer. Desig. by Michener, 1941.
anthocopoides (Schenck). N. Y. (Albany and adjacent counties); Europe. Ecology: Builds
mortar and pebble nests on exposed rock surfaces. Adventive from Europe. Parasite:
Acroricnus stylator (Thunb.), Anthrax irroratus Say, Chrysis coerulans Fabr.,
Melittobia chalybii Ashm., Monodontomerus montivagus Ashm., Stelis labiata (Prov.).
Pollen: Oligolectic on Echium vulgare, but in Europe also collects pollen from Anchusa
officinalis.
Osmia spinolae Schenck, 1851. Nassau. Ver. f. Naturk. Jahrb. 7: 68. 2. Preoce.
Osmia anthocopoides Schenck, 1853. Nassau. Ver. f. Naturk. Jahrb. 9: 181. N. name.
Osmia caementaria Gerstaecker, 1869. Stettin. Ent. Ztg. 30: 339. 2, d.

Taxonomy: Eickwort, 1970. Psyche 77: 190-201, 11figs. (synonymy, redescription, systematic
position, distribution).

Biology: Eickwort, 1970. Psyche 77: 196-199 (floral relationships, summary of life history).
—Eickwort, 1973. Search, Cornell Univ. Agr. Expt. Sta. 3: 1-31, 55 figs., 3 tables (life
history, nest architecture, immature stages, parasites). —Eickwort, 1975. Evolution 29:
142-150, 1 fig., 2 tables (gregarious nesting and evolution of parasitism and sociality among
megachilid bees). —Eickwort, 1975. Ztschr. Tierpsychol. 3: 237-254, 8 figs, 1 table
(nest-building behavior).

Morphology: Schmidt and Stockton, 1971. Ent. News 82: 275-277 (analysis of DNA in sperm).

Genus HOPLITIS Subgenus ANDRONICUS Cresson

Andronicus Cresson, 1864. Ent. Soe. Phila., Proc. 2: 384.
Type-species: Andronicus cylindricus Cresson. Montypic.
cylindrica (Cresson). Que. and N. S. to Fla., west to N. W. T., B. C., Colo., Tex. Ecology: Nests

in dry pithy stems including trap-nests. Parasite: Melittobia chalybii Ashm., Stelis
foederalis Smith, S. labiata Prov., S. lateralis Cress., Tricrania stansburyi Hald. Pollen:
Polylectic, visits flowers of many families, principally Leguminosae, Compositae, and
Labiatae, including Amorpha, Blephilia, Celastrus, Dianthera, Hydrophyllum, Lobelia,
Petalostemon, Rubus, Senecio, Specularia, Trifolium, Verbena, Veronica.

Andronicus cylindricus Cresson, 1864. Ent. Soc. Phila., Proc. 2: 384. d.

Osmia spoliata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 327. °.

Hopilitis monardae Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 363. 2.

Hoplitis hicksi Cockerell, 1932. Brooklyn Ent. Soc., Bul. 27: 203. °.

Biology: Hicks, 1926 Colo. Univ. Studies 15: 217 (nest). —Fye, 1965. Canadian Ent. 97:
871-872, fig. 2 (life history, parasite). —Medler, 1967. Kans. Ent. Soc., Jour. 40: 187-140, fig.
1 (nest architecture, life history, parasites).

Genus HOPLITIS Subgenus MONUMETHA Cresson

Monumetha Cresson, 1864. Ent. Soe. Phila., Proc. 2: 387.
Type-species: Chelostoma albifrons Kirby. Desig. by Titus, 1904. (=Monumetha
argentifrons Cresson).
Chlorosmia Sladen, 1916. Canad. Ent. 48: 270.
Type-species: Osmia fulgida Cresson. Monotypic.

Taxonomy: Michener, 1936. Amer. Mus. Novitates 875: 29 (key).

Superfamily APOIDEA 2015

albifrons albifrons (Kirby). N.S., to N. Y., west to B. C., north to N. W. T. and Alaska.
Parasite: Melittobia chalybii Ashm., Stelis subemarginata Cress. Pollen: Unknown, but
visits flowers of Phacelia.
Chelostoma albifrons Kirby, 1837. In Richardson, Fauna Bor.-Amer., v. 4, p. 270. 6.
Monumetha borealis Cresson, 1864. Ent. Soe. Phila., Proc. 2: 388. °.
Monumetha obsoleta Cresson, 1864. Ent. Soe. Phila., Proc. 2: 388. d.
Megachile oblonga Provancher, 1882. Nat. Canad. 13: 230. 9.

Taxonomy: Michener, 1947. Evolution, 1: 172.

Biology: Fye, 1965. Canad. Ent. 97: 867-871, tables 2-4 (nest, nest architecture, parasite).

albifrons argentifrons (Cresson). Alta. south to N. Mex. and Ariz., east to Nebr. (Sioux Co.),
west to B. C., Wash. and Oreg., thence south along Sierra Nev. in Calif. Pollen:
Apparently polylectic, visits a wide variety of flowers including Allium, Apocynum,
Arctostaphylos nevadensis, Astragalus, Cirsium, Clarkia pulchella, C. unguiculata, C.
williamsoni, C. xantiana, Cleome serrulata, Erigeron, Eriodictyon californicum,
Geranium fremontii, Gilia, Gormania obtusata, Lotus, Mertensia franciscana, Mirabilis
laevis, Nama rothrockii, Opulaster, Opuntia, Pedicularis groenlandica, Penstemon
newberryi, Phacelia frigida, P. glandulosa, P. leucophylla, Phyllodoce brewer,
Potentilla glandulosa, Rosa, Senecio, Spiraea sorbifolia, Streptanthus tortuosus.

Monumetha argentifrons Cresson, 1864. Ent. Soe. Phila., Proc. 2: 387. d.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 51 (floral
relationships).
albifrons maura (Cresson). Southwest. Oreg., Calif. (west of Sierra Nevada Mts.). Ecology: A

female has been collected carrying a pulpy, green mass, apparently consisting of
macerated leaf tissue. Parasite: Stelis sp., Tricrania stansburyi Hald. Pollen:
Apparently polylectic, visits a wide variety of flowers including Apocynum
androsaemifolium, Brodiaea lutea, Chamaebatia foliolosa, Clarkia biloba, C. brewer, C.
concinna, C. cylindrica, C. dudleyana, C. elegans, C. gracilis albicaulis, C. rhomboidea,
C. rubicunda, C. speciosa nitens, C. unguiculata, Cirsium, Convolulus, Cordylanthus
pilosus, Cryptantha intermedia, Eriodictyon trichocalyx, Eriogonum, Gilia capitata,
Gormania obtusata, Iris missouriensis, Helianthella californica, Lotus davidsonii, L.
scoparius, Lonicera interrupta, Mimulus, Mirabilis laevis, Nemophila integrifolia, N.
menziesii, Penstemon, Phacelia californica, P. distans, P. heterophylla, P. ramosissima,
Phyllodoce breweri, Ranunculus, Stanleya pinnata.

Osmia maura Cresson, 1878. Amer. Ent. Soc., Trans. 7: 104. 9.

Andronicus hesperius Cockerell, 1903. South. Calif. Acad. Sci., Bul. 2: 35. d.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 51 (floral
relationships).
fulgida fulgida (Cresson). Alaska and Yukon, south to Oreg., Ariz. and N. Mex. Ecology: Nests

in abandoned beetle burrows in wood of various sorts. Parasite: Aritranis imitator
ruficauda Townes, Melittobia sp., Sapyga aculeata Cress., Sphaeropthalma sp., Stelis
monticola Cress., Stelis sp., Tricrania stansburyi Hald. Pollen: Apparently polylectic,
visits flowers of Allium diehlii, Clarkia pulchella, Convolvulus, Fragaria vesca,
Gentiana parryi, Geranium caespitosum, Phacelia leucophylla, P. linearis, Potentilla,
Prunus melanocarpa, Ranunculus, Rosa, Rubus deliciosus. Predator: Trichodes
ornatus Say.

Osmia fulgida Cresson, 1864. Ent. Soe. Phila., Proc. 3: 34. °.

Osmia viridis Cresson, 1864. Ent. Soc. Phila., Proc. 3: 35. °.

Biology: Clement and Rust, 1976. Pan-Pacifie Ent. 52: 114-116, figs. 3, 4, tables 1, 2 (nest
architecture, cell provisions, development, parasites).

fulgida platyura (Cockerell). South. Oreg., Calif. Parasite: Anthrax irroratus Say, Lewcospis
affinis Say, Sapyga aculeata Cress. Pollen: Apparently polylectic, visits flowers of
Allium parvum, Astragalus douglasii, Calochortus nudus, Centromadia, Chamaebatia
foliolosa, Clarkia cylindrica, C. dudleyana, C. purpurea quadrivulnera, C. rhomboidea,
C. unguiculata, Chaenactis glabriuscula, Collinsia heterophylla, C. tinctoria, C. torreyi
wrightii, Cryptantha intermedia, Eriodictyon californicum, E. crassifolium, Fragaria

2016 Hymenoptera in America North of Mexico

californica, Gilia capitata, Haplopappus linearifolius, Hemizonia, Hesperochiron
californicus, Lappula, Lotus subpinnatus, Lupinus bicolor, L. formosus, L. nanus,
Marrubium vulgare, Mentzelia albicaulis, Mimulus, Nemophila integrifolia, N.
maculata, N. menziesii, Phacelia alba, P. ciliata, P. davidsonii, P. distans, P. frigida, P.
heterophylla, P. ramosissima, P. tanacetifolia, Potentilla glandulosa, Rhus trilobata,
Salix, Sidalcea malvaeflora, Sphaeralcea fasciculatum, Stachys californica, Taraxacum,
Thysanocarpus curvipes, Trichostema parishii, Trifolium.

Osmia platywra Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 765. 36, °.

Osmia lawae Michener, 1936. Amer. Mus. Novitates 875: 29. 6, 2.

Biology: Clement and Rust, 1976. Pan-Pacifie Ent. 52: 114-116, figs 3, 4, tables 1, 2 (nest
architecture, cell provisions, development)
louisae (Cockerell). B. C., Wash., Oreg., Idaho, north. Calif. and Utah.
Osmia louisae Cockerell, 1934. Amer. Mus. Novitates 679: 14. °.
viridimicans (Cockerell). Wash., west Oreg., Calif. (Sierra Nevada Mts.).
Osmia viridimicans Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 334. °.

Genus HOPLITIS Subgenus DASYOSMIA Michener

Hoplitis subg. Dasyosmia Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 294.
Type-species: Alcidamea biscutellae Cockerell. Orig. desig.
biscutellae (Cockerell). Tex., N. Mex., Ariz., Utah, Nev., south. Calif.; deserts. Ecology: Nest

has been observed in a bank. Parasite: Anthrax irroratus Say, Nemognatha macswaini
Enns, N. nigripennis LeC., Stelis sp. Pollen: Oligolege of Larrea tridentata, but also
visits other flowers for nectar including Aster abatus, Encelia farinosa, Geraea
canescens, Helianthus niveus, Hyptis emoryi, Prosopis, Salvia, Sphaeralcea, ambigua,
Stenotopsis linearifolius. Predator: Cymatodera sp., Trichodes ornatus Say.

Alcidamea biscutellae Cockerell, 1897. Ann. and Mag., Nat. Hist. (6) 19: 400. ¢.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 36, tables 2, 5, 7, 9, 11-15 (floral
relationship with Larrea).
paroselae Michener. East. Calif., desert. Pollen: Unknown, but visits flowers of Dalea

fremontii, D. polyadenia.
Hoplitis (Dasyosmia) paroselae Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 296. d, 2.

Genus HOPLITIS Subgenus CYRTOSMIA Michener

Hoplitis subg. Cyrtosmia Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 292.
Type-species: Osmia hypocrita Cockerell. Orig. desig. and monotypic.
hypocrita (Cockerell). B. C. to Calif., Nev., Utah., Colo. and Ariz. Ecology: Nests in dead dry

stems and also trap-nests of various sorts. Parasite: Aritranis imitator ruficauda
Townes, Eurytoma sp., Nemognatha scutellaris LeC., Sapyga aculeata Cress., Stelis
seemaculata Ashm., Stelis sp., Tricrania stansburyi Hald. Pollen: Polylectic, visits a
wide variety of flowers including Amsinckia douglasiana, Astragalus antisellii, A.
goniatus, A. pomonensis, A. parishii, A. tener, Brodiaea capitata, Brassica campestris,
Eriodictyon californicum, Grindelia, Lathyrus, Lotus crassifolius, L. glaber, L.
scoparius, Lupinus paynei, Medicago sativa, Penstemon leonardi, Salix, Salvia carnosa,
Sambucus, Stanleya pinnata, Trifolium, Vicia.

Osmia hypocrita Cockerell, 1906. Canad. Ent. 38: 160. °.

Biology: Hicks, 1926. Colo. Univ. Studies 15: 217 (nest). —Clement and Rust, 1976.
Pan-Pacific Ent. 52: 111-114, figs. 1, 2, tables 1, 2 (nest architecture, cell provisions,
development, parasite)

Genus HOPLITIS Subgenus ALCIDAMEA Cresson

Alcidamea Cresson, 1864. Ent. Soc. Phila., Proc. 2: 385.
Type-species: Alcidamea producta Cresson. Desig. by Michener, 1941.
Autochelostoma Sladen, 1916. Canad. Ent. 48: 270.
Type-species: Alcidamea producta Cresson. Monotypic. (=Autochelostoma
canadensis Sladen).

Superfamily APOIDEA 2017

The species of this subgenus, whose habits are known, nest in the pithy cores of dead stems or
canes of various plants.
Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1051, figs. 142-145, 149 (larva).
brachyodonta (Cockerell). South. Calif. Ecology: Uses plant material sometimes supplemented
with pebbles in nest construction. Parasite: Stelis sewmaculata Ashm. Pollen: Unknown,
but visits flowers of Chaenactis glabriuscula, Heliotropium curassavicum.
Alicidamea brachyodonta Cockerell, 1933. Ent. News 44: 205. d.
colei (Crawford). South. Nev., south. Calif. Parasite: Nemognatha scutellaris LeC., Stelis
sexmaculata Ashm. Pollen: Possibly an oligolege of Eriodictyon including LE.
californicum, E. crassifolium, E. trichocalyx, but also visits other flowers including
Collinsia tinctoria, Eriastrum virgatum, Lotus, Nama demissum.
Alcidamea colei Crawford, 1916. Ent. Soc. Wash., Proc. 18: 127. d.
elongaticeps Michener. Calif. (Mojave and east. deserts). Pollen: Unknown, but visits flowers
of Dalea fremontii, Mimulus.
Hoplitis (Alcidamea) elongaticeps Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 290. °,
é.
grinnelli grinnelli (Cockerell). Ariz., Calif., north. Mexico (Baja California and Sonora).
Parasite: Nemognatha scutellaris LeC., Stelis seemaculata Ashm. Pollen: Polylectic,
visits a wide variety of flowers including Amsinckia intermedia, Astragalus fremontii,
Ceanothus, Chaenactis, Chorizanthe staticoides, Collinsia torreyi, Cryptantha
intermedia, C. racemosa, Dalea fremontii, Eriastrum virgatum, Erigeron stenophyllus,
Eriogonum fasciculatum, Eucnide urens, Larrea tridentata, Lotus glaber, L. scoparius,
L. strigosus, L. subpinnatus, Lupinus concinnus, Marrubium vulgare, Melilotus indica,
Mentzelia, Mesembryanthemum edule, Mimulus, Phacelia distans, P. platyloba, P.
ramosissima, Prosopis pubescens, Rhamnus crocea, Salvia mellifera, Sphaeralcea
ambigua, Trifolium.
Hoplitis grinnelli Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 22. 2.
grinnelli septentrionalis Michener. B. C., Wash., Oreg., Idaho, Utah. Pollen: Unknown, but
visits flowers of Linwm, Penstemon cyananthus, Phacelia linearis.
Hoplitis grinnelli septentrionalis Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 291. d,
De
pilosifrons (Cresson). Que. and Mass. to Fla., west to Alta., Colo., and Tex. Ecology: Excavates
pith in dead stem of Helianthus tuberosus for nest. Parasite: Stelis labiata Cress., S.
lateralis (Prov.). Pollen: Stores pollen of Amorpha fruticosa and Melilotus officinalis,
but also visits other flowers for nectar and or pollen including Barbarea, Blephilia,
Cardamine, Chrysanthemum, Coreopsis, Desmodium, Dianthera, Erigeron, Geranium,
Gillenia, Heracleum, Houstonia, Hydrophyllum, Krigia, Lepachys, Linaria, Lobelia,
Lythrum, Malva, Nepeta, Oenothera, Oxalis, Pedicularis, Penstemon, Petalostemon,
Polymonium, Potentilla, Psoralea, Pycnanthemum, Radicula, Rubus, Scutellaria,
Senecio, Specularia, Stachys, Taenidia, Tephrosia, Teucrium, Trifolium, Verbena,
Veronica, Vicia, Zizia.
Alcidamea pilosifrons Cresson, 1864. Ent. Soc. Phila., Proc. 2: 386. d.
Hoplitis graceae Cockerell, 1923. Ann. and Mag. Nat. Hist. (9) 11: 268. °.
Alcidamea mucronata Cockerell, 1934. Brooklyn Ent. Soc., Bul. 29: 18. d.

Biology: Michener, 1955. Kans. Ent. Soc., Jour. 28: 81-83 (nest, life history, parasite).

producta bernardina Michener. South. Calif. Mts. (overlapping range of producta gracilis).
Pollen: Apparently polylectic, visits a wide variety of flowers including Astragalus
parishii, Collinsia, Convolvulus, Dicentra chrysantha, Gilia exilis, Helianthus,
Hesperochiron, Lotus scoparius, Mesembryanthemum crystallinum, Mimulus fremontii,
Penstemon cordifolius, P. grinnelli, P. heterophyllus, P. spectabilis, Phacelia.

Hoplitis producta bernardina Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 287. ¢, °.

producta gracilis (Michener). Oreg., Calif. Nev. Ecology: Nests in dead, pithy stems and
trap-nests of various sorts. Parasite: Aprostocetus sp., Chrysura sonorensis (Cam.),
Eurytoma stigmi Ashm., Sapyga aculeata Cress., Stelis secmaculata Ashm. Pollen:
Apparently polylectic, visits a wide variety of flowers including Allium parvum, Aster

2018

Hymenoptera in America North of Mexico

foliaceus, Astragalus bolanderi, Brodiaea capitata, Calyptridium umbellatum, Carduus
tenuiflorus, Castilleia, Chamaebatia foliolosa, Cirsium vulgare, Clarkia amoena, C.
biloba, C. dudleyana, C. elegans, C. gracilis albicaulis, C. purpurea, C. rhomboidea, C.
unguiculata, C. williamsonii, Collinsia parviflora, C. tinctoria, C. torreyi, Convolvulus,
Cryptantha, Dudleya, Hugelia brauntonit, Eriodictyon californicum, E. trichocalyzx,
Gayophytum diffusum, Gilia exilis, Glycyrrhiza lepidota, Gormania obtusata,
Hesperochiron, Heterogaura heterandra, Kelloggia galioides, Lepechinia calycina,
Lessingia germanorum, Lotus argophyllus, L. corniculatus, L. davidsonii, L.
nevadensis, L. scoparius, Lupinus breweri, Mimulus, Monardella villosa, Navarretia
heterodoxa, Nemophila, Penstemon heterodoxus, P. newberryi, P. spectabilis, Phacelia
davidsonii, P. distans, P. linearis, Potentilla glandulosa, Ranunculus californicus,
Rosa, Rubus leucodermis, Salix, Satureja douglasii, Sidalcea calycosa, Trifolium
gracilentum, T. variegatum, Verbena lasiostachys, Vicia americana. >

Osmia gracilis Michener, 1935. Pan-Pacifie Ent. 11: 183. °.
Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ., Pubs. Ent. 70: 51 (floral

relationships).

producta interior Michener. Alta., Wyo., Colo., Utah, N. Mex., Ariz. (west of plains). Pollen:

Apparently polylectic, visits flowers of Achillea millefolium, Astragalus, Helianthus
peliolaris, Monarda pectinata, Penstemon angustifolium, Phacelia linearis.

Hoplitis producta interior Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 286. 3, 9.

producta panamintana Michener. Calif. (Panamint and nearby mts.). Pollen: Unknown, but

visits flowers of Phacelia.

Hoplitis producta panamintana Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 287. 3.

producta producta (Cresson). Que. to Ga., west to Alta., Colo., and Tex. Ecology: Nests in

sumac borings. Parasite: Anthrax irroratus Say, Leucospis affinis Say, Nemognatha
nigripennis LeC., N. scutellaris LeC., Sapyga aculeata Cress., Stelis crassiceps Ckll., S.
labiata (Prov.), S. lateralis Cress., S. sexmaculata Ashm., Tricrania stansburyi Hald.
Pollen: Polylectic, visits a wide variety of flowers including Amorpha fruticosa,
Apocynum, Astragalus, Azalea, Baptisia, Barbarea, Chrysanthemum, Crataegus,
Erigeron philadelphicus, Geranium maculatum, Gilia, Lesquerella, Medicago sativa,
Melilotus officinalis, Mentha canadensis, Penstemon hirsutus, Phacelia dubia,
Potentilla, Rubus argutus, Salix, Solidago, Tephrosia virginiana, Trifolium. Predator:
Philanthus pulcher Dalla Torre.

Alcidamea producta Cresson, 1864. Ent. Soc. Phila., Proc. 2: 386. 3.
Autochelostoma canadensis Sladen, 1916. Canad. Ent. 48: 270. “3” = intersex.
Alcidamea helenae Cockerell, 1934. Amer. Mus. Novitates 732: 6. d.

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 153 (nest). —Comstock, 1924.

Introduction to entomology, p. 824 (nest). —Hicks, 1926. Colo. Univ. Studies 15: 217 (nest).
—Rau, 1928. Psyche 35: 100-107, 1 fig. (nest, life history). —Medler, 1961. Canad. Ent. 93:
571-573 (nest, life history).

producta subgracilis Michener. B. C., Wash., Oreg., Idaho.

Hoplitis producta subgracilis Michener, 1947. Amer. Mus. Nat. Hist., Bul. 89: 286. 6, 2.

sambuci Titus. B. C. to Calif., Nev. and Utah. Ecology: Nests in dry, pithy stems and

trap-nests of various sorts. Parasite: Gasteruption kirbii russewm Townes, Nemognatha
nigripennis LeC., N. scutellaris LeC., Sapyga aculeata Cress., Sphaeropthalma sp.,
Stelis sp. Pollen: Apparently polylectic, visits a wide variety of flowers including
Amorpha fruticosa, Asclepias eriocarpa, Astragalus antisellu, Cirsium, Clarkia
unguiculata, Cryptantha, Dicentra chrysantha, Eriodictyon californicum, E.
crassifolium, Eriogonum, Helianthus, Hesperochiron, Layia glandulosa, Lonicera
interrupta, Lotus glaber, L. scoparius, Lupinus formosus, L. hallii, L. paynei, Medicago
sativa, Mimulus guttatus, Nama parryi, Penstemon spectabilis, Phacelia, Potentilla
glandulosa, Rhamnus californica, Rubus leucodermis, Salvia mellifera, Sambucus
glauca, Stanleya pinnata, Trichostema ovatum, Vicia americana.

Hoplitis sambuci Titus, 1904. Ent. Soc. Wash., Proc. 6: 101. 2, d.

Biology: Clement and Rust, 1976. Pan-Pacific Ent. 52: 116-117, fig. 5, tables 1, 2 (nest

architecture, cell provisions, development, parasite).

Superfamily APOIDEA 2019

truncata mescalerium Cockerell. Colo., N. Mex., Ariz. Pollen: Unknown, but visits flowers of
Gilia.
Hoplitis mescalerium Cockerell, 1910. Entomologist 43: 90. °.
truncata truncata (Cresson). Que. and Maine to Fla., west to N. Dak., Wyo., Colo. and Miss.
Parasite: Sapyga centrata Say. Pollen: Apparently polylectic, visits a wide variety of
flowers including Baptisia, Berlandiera, Ceanothus, Cleome, Convolvulus, Erigeron,
Gillenia, Ilex, Melilotus, Oenothera, Penstemon, Pogonia graminifolia, Rubus,
Tephrosia virginiana, Trifolium, Vaccinium, Vicia.
Alcidamea truncata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 108. ¢.
uvulalis (Cockerell). Calif., Oreg., Idaho, Utah. Parasite: Nemognatha scutellaris LeC. Pollen:
Unknown, but visits flowers of Castilleia, Senecio.
Alcidamea uvulalis Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 139. ¢.

Genus HOPLITIS Subgenus FORMICAPIS Sladen

Formicapis Sladen, 1916. Canad. Ent. 48: 271.
Type-species: Formicapis clypeata Sladen. Monotypice.

Revision: Michener, 1938. Ent. News 49: 129 (as clypeata).
robusta (Nylander). Holarctic, Alaska, N. W. T., Que., B. C., Alta., Sask., Mont. Wyo., Colo.,

Oreg., Calif.; Europe and Asia. Pollen: Apparently polylectic, evidently stores pollen of
Astragalus, Silene ruprestris, Taraxacum, Trifolium. Predator: Philanthus pulcher
Dalla Torre.

Heriades robusta Nylander, 1848. Notis. Saellsk. Faun. Flor. Fenn. Forh. 1: 270. 2, d.

Osmia rhinoceros Giraud, 1861. Zool.-Bot. Gesell. Wien, Verh. 11: 464. 2.

Formicapis clypeata Sladen, 1916. Canad. Ent. 48: 271. °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 57-58, figs. 19-20
(redescription). —Peters, 1970. Senckenb. Biol. 51: 193-197 (synonymy, distribution, floral
relationships).

Biology: Clement and Rust, 1975. Ent. News 86: 115-120, 2 figs. (nest, cocoon, supersedure,
floral relationships).

Genus HOPLITIS Subgenus ROBERTSONELLA Titus

Robertsonella Titus, 1904. N. Y. Ent. Soc., Jour. 12: 22.
Type-species: Robertsonella gleasoni Titus Monotypie and orig. desig.

Revision: Michener, 1938. Ent. News 49: 130.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 64-68, fig. 25, table 3 (eastern
U.S. spp.).
gleasoni (Titus). Tex. to Ill. and N. J. Pollen: Unknown, but visits flowers of Geranium,
Phacelia, Rubus.
Robertsonella gleasoni Titus, 1904. N. Y. Ent. Soe., Jour. 12: 23. 2, d.
Robertsonella crataegina Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 28. d.
micheneri Mitchell. Kans., Ga. Pollen: Unknown, but visits flowers of Amorpha fruticosa.
Hoplitis (Robertsonella) micheneri Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 64,
fig. 25. 9, 6.
simplex (Cresson). Tex. to Conn. Parasite: Stelis labiata (Prov.). Pollen: Unknown, but visits

flowers of Phacelia, Salix.
Heriades simplex Cresson, 1864. Ent. Soe. Phila., Proc. 2: 384. °.

Genus HOPLITIS Subgenus Unassigned

alboscopata (Provancher). Que.
Heriades alboscopatum Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 425.

2
Taxonomy: Titus, 1906. Ent. Soc. Wash., Proce. 7: 160.

2020 Hymenoptera in America North of Mexico

imperfecta (Provancher). Calif.
Monumetha imperfecta Provancher, 1896. Nat. Canad. 23: 9. 2. Perhaps a Megachile.

Genus ANTHOCOPA Lepeletier

This rather large genus of chiefly Holarctie bees occurs in Eurasia, Africa and in western
North America, west of the Great Plains. Although there is some evidence which indicates that
Anthocopa should be merged with Hoplitis, these taxa are maintained as distinct in this catalog.

Revision: Michener, 1943. Ent. Soc. Amer., Ann. 36: 49-86 (Nearctic spp.).

Taxonomy: Hurd and Michener, 1955. Calif. Ins. Survey, Bul. 3: 94-126, pls. 8, 19-20, figs.
71-91, maps 40-61 (Calif. spp.). —Michener and Sokal, 1957. Evolution 11: 130-162 (phenetic
classification of the Hoplitis complex). —Sokal and Michener, 1958. Kans. Univ. Sci. Bul.
38: 1409-1438 (systematic relationships within the Hoplitis complex). —Michener, 1966.
Entomologist 99: 146 (status of Anthocopa and Hoplitis). —Rubin, 1966. Syst. Zool. 15:
176-182, tables 1-3 (phenetic classification of the Hoplitis complex). —Michener and Sokal,
1966. Ent. Soc. Amer., Ann. 59: 1211-1217, 5 figs. (phenetic similarities among the species
of the Hoplitis complex). —Michener, 1968. Ent. South Africa, Jour. 31: 337 (status of
Anthocopa and Hoplitis).

Genus ANTHOCOPA Subgenus ANTHOCOPA Lepeletier

Anthocopa Lepeletier, 1825. Encycl. Meth., Dict. Ins., v. 10, p. 314.
Type-species: Apis papaveris Latreille. Monotypic.
Phyllotoma Dumeril, 1860. Acad. Sci. Inst. Imp. France, Mem. 31: 842. Preocce.
Type-species: Apis papaveris Latreille. Desig. by Michener, 1941.
Pseudosmia Radoszkowski, 1872. Soc. Ent. Rossica, Horae 8: xviii. Republished by
Radoszkowski, 1874. Soc. Imp. Nat. Moscou, Bul. 48 (1): 152.
Type-species: Megachile cristata Fonscolombe. Desig. by Cockerell, 1922.
Pseudo-osmia Radoszkowski, 1873. Soc. Imp. Nat. Moscou, Bul. 46 (2): 187. Emend.
Peudocosmia Radoszkowski, 1886. Soc. Ent. Rossica, Horae 20: 14. Emend.

This subgenus does not occur in the New World.
Genus ANTHOCOPA Subgenus ATOPOSMIA Cockerell

Osmia subg. Atoposmia Cockerell, 1935. Pan-Pacific Ent. 11: 50.
Type-species: Osmia triodonta Cockerell. Orig. desig.

All species visit chiefly flowers of Penstemon in the mountains.

abjecta abjecta (Cresson). Calif. (White Mts.), Oreg., Utah, Wyo., Colo. Ecology: Nests under
stones. Pollen: Oligolege of Penstemon, including P. alpinus, P. heterodoxus, P. moffatti,
P. newberryi.
Osmia abjecta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 103. °.
Hoplitis mesae Cockerell, 1930. Amer. Mus. Novitates 397: 2. ¢.
Anthocopa (Atoposmia) nigrior Michener, 1943. Ent. Soc. Amer., Ann. 36: 54. 2 (6
misdet.).

Taxonomy: Michener, 1954. Pan-Pacific Ent. 30: 43 (synonymy).

Biology: Parker, 1975. Pan-Pacific Ent. 51: 119-121, table 1 (nest, cocoon, sex ratio).
abjecta alta (Michener). Calif. (Sierra Nevada Mts.).
Osmia alta Michener, 1936. Canad. Ent. 68: 41. °.
anthodyta anthodyta Michener. Calif., Nev., Oreg., Idaho; Mexico (Baja California). Pollen:
Oligolege of Penstemon including P. cinereus, P. grinnellii, P. newberryi, P. palmeri,
but also visits flowers of Collinsia parviflora.
Anthocopa (Atoposmia) anthodyta Michener, 1943. Ent. Soc. Amer., Ann. 36: 60. ?, d.
anthodyta bequaerti Michener. Ariz. (Santa Rita Mts.). Pollen: Penstemon.
Anthocopa anthodyta bequaerti Michener, 1954. Pan-Pacific Ent. 30: 42. d, 9.
arizonensis Michener. Ariz.
Anthocopa arizonensis Michener, 1954. Pan-Pacifie Ent. 30: 43. 9.

Superfamily APOIDEA 2021

elongata (Michener). Calif., Oreg., Wash., Mont., Colo. Pollen: Oligolege of Penstemon including
P. heterodoxus, P. newberryi, but also visits flowers of Phacelia, Potentilla presumably
for nectar.
Osmia elongata Michener, 1936. Canad. Ent. 68: 41. 9.
hebitis Michener. Calif. (Sierra Nevada Mts.). Pollen: Presumably an oligolege of Penstemon,
but also visits flowers of Mimulus.
Anthocopa hebitis Michener, 1954. Pan-Pacifie Ent. 30: 44. 2, d.

Taxonomy: Snelling, 1962. Pan-Pacifiec Ent. 38: 228 (geogr. and floral records).
oregona Michener. Calif. (Sierra Nevada Mts.), Oreg.
Anthocopa (Atoposmia) oregona Michener, 1943. Ent. Soc. Amer., Ann. 36: 53. 2, 6.
panamintensis Michener. Calif. (Panamint Mts.).
Anthocopa (Atoposmia) panamintensis Michener, 1943. Ent. Soc. Amer., Ann. 36: 65. 2, d.
pycnognatha pycnognatha Michener. East. Calif. Pollen: Oligolege of Penstemon including P.
breviflorus.
Anthocopa (Atoposmia) pycnognatha Michener, 1943. Ent. Soc. Amer., Ann. 36: 64. 2, ¢.
pycnognatha solata Michener. Calif. Pollen: Oligolege of Penstemon including P.
antirrhinoides, P. spectabilis, but also visits flowers of Euphorbia albomarginata,
Salvia columbariae presumably for nectar.
Anthocopa (Atoposmia) pycnognatha solatus Michener, 1949. Kans. Ent. Soc., Jour. 22: 48.
Taxonomy: Snelling, 1962. Pan-Pacific Ent. 38: 228 (geogr. and floral records).
triodonta shastensis (Cockerell). Calif. (Sierra Nevada Mts.), Nev., Oreg. Pollen: Oligolege of
Penstemon including P. cinereus.
Osmia shastensis Cockerell, 1935. Pan-Pacifie Ent. 11: 46. °.
triodonta triodonta (Cockerell). Calif. (Mt. Diablo, Sierra Nevada and White Mts.). Pollen:
Oligolege of Penstemon including P. heterophyllus.
Osmia (Atoposmia) triodonta Cockerell, 1935. Pan-Pacific Ent. 11: 50. d, °.
triodonta usingeri Michener. South. Calif. Pollen: Oligolege of Penstemon including P.
grinnellii, P. labrosus, P. spectabilis, but also visits other flowers presumably for nectar
including Nama parryi.
Anthocopa triodonta usingeri Michener, 1943. Ent. Soc. Amer., Ann. 36: 68. d, 2.

Genus ANTHOCOPA Subgenus HEXOSMIA Michener

Anthocopa subg. Hexosmia Michener, 1943. Ent. Soc. Amer., Ann. 36: 74.
Type-species: Osmia copelandica Cockerell. Orig. desig.
copelandica albomarginata (Cockerell). Calif. west. Oreg. Parasite: Chrysura sonorensis

Cam., Epistenia sp., Nemognatha scutellaris LeC., Sapyga pumila Cress. Pollen:
Apparently an oligolege of Phacelia including P. ciliata, P. davidsonii, P. distans, P.
heterophylla, P. hydrophylloides, P. ramosissima, but also visits other flowers including
Collinsia wrightii, Mimulus rubellus, Nemophila integrifolia, N. menziesii.

Osmia albomarginata Cockerell, 1935. Pan-Pacifie Ent. 11: 49. 2, 6.

Biology: Parker, 1975. Pan-Pacific Ent. 51: 116-119, figs. 7-8, table 1 (nest, cocoon, parasites).
copelandica arefacta (Cockerell). South. Calif., deserts. Parasite: Leucospis affinis Say. Pollen:
Apparently an oligolege of Phacelia including P. distans, but also visits other flowers
presumably for nectar including Cryptantha intermedia, Malacothrix glabrata.

Predator: Cymatodera sp.
Osmia arefacta Cockerell, 1935. Pan-Pacifie Ent. 11: 42. 2, d.

Biology: Parker, 1975. Pan-Pacific Ent. 51: 116, 117, 119, fig. 9, table 1 (nest, cocoon, nest
associates).
copelandica copelandica (Cockerell). Colo., Utah, Wyo., Mont., Idaho, east. Oreg., Wash., B. C.
Pollen: Apparently an oligolege of Phacelia.
Osmia copelandica Cockerell, 1908. Entomologist 41: 59. °.
Osmia besseyae Cockerell, 1910. Entomologist 43: 92. 2, d.

Biology: Parker, 1975. Pan-Pacific Ent. 51: 116 (Idaho nests only).

2022 Hymenoptera in America North of Mexico

phaceliarum (Cockerell). South. Calif. Pollen: Apparently an oligolege of Phacelia including P.
distans.
Osmia phaceliarum Cockerell, 1935. Pan-Pacifie Ent. 11: 45. 9.

Genus ANTHOCOPA Subgenus EREMOSMIA Michener

Anthocopa subg. Eremosmia Michener, 1943. Ent. Soc. Amer., Ann. 36: 66.
Type-species: Osmia robustula Cockerell. Orig. desig.

Anthocopa subg. Phaeosmia Michener, 1943. Ent. Soc. Amer., Ann. 36: 77.
Type-species: Osmia enceliae Cockerell. Orig. desig.

Revision: Michener, 1949. Kans. Ent. Soc., Jour 22: 53 (spp. formerly placed in Phaeosmia).
beameri Michener. Tex. (Big Bend Natl. Park, Marathon). Pollen: Unknown, but visits flowers
of Nama, Phacelia.
Anthocopa (Eremosmia) beameri Michener, 1951. Pan-Pacific Ent. 27: 64. ?, 6.
daleae Michener. Tex. (Big Bend Natl. Park, Sanderson). Pollen: Unknown, but visits flowers
of Dalea, Phacelia.
Anthocopa (Phaeosmia) daleae Michener, 1951. Pan-Pacifie Ent. 27: 62. 2, 6.
enceliae enceliae (Cockerell). South. Calif., desert border. Pollen: Unknown, but visits flowers
of Baileya, Encelia farinosa.
Osmia enceliae Cockerell, 1935. Pan-Pacifie Ent. 11: 43. 2.
enceliae mortua (Cockerell). Ariz., Calif., Nev. Pollen: Unknown, but visits flowers of Encelia
actoni, E. farinosa.
Osmia viguierae var. mortua Cockerell, 1935. Pan-Pacifie Ent. 11: 44. 9, 6.
Anthocopa (Eremosmia?) mallognatha Michener, 1943. Ent. Soc. Amer., Ann. 36: 67. 9.
hemizoniae (Cockerell). South. Calif. Pollen: Unknown, but visits flowers of Helianthus
gracilentus, Hemizonia paniculata.
Osmia hemizoniae Cockerell, 1935. Pan-Pacifie Ent. 11: 47. 9.

Taxonomy: Michener, 1954. Pan-Pacific Ent. 30: 50. 6.
hypostomalis Michener. Calif. Ecology: Nests in preexisting burrows in wood including
prebored elderberry trap stems. Parasite: Leucospis affinis Say, Sphaeropthalma
amphion (Fox), Stelis sp. Pollen: Apparently an oligolege of Dalea including D.
fremontii, D. schottii, but also visits other flowers including Cryptantha barbigera,
Palafoxia linearis. Predator: Cymatodera sp.
Anthocopa (Phaeosmia) hypostomalis Michener, 1949. Kans. Ent. Soc., Jour. 22: 50. °.

Taxonomy: Michener, 1954. Pan-Pacific Ent. 30: 49. 6.

Biology: Parker, 1975. Pan-Pacific Ent. 51: 113-116, figs. 1-6, table 1 (nest, cocoon, sex ratio,
nest associates).
maryae Michener. Tex. Pollen: Unknown, but visits flowers of Dalea.
Anthocopa (Phaeosmia) maryae Michener, 1949. Kans. Ent. Soc., Jour. 22: 50. 2.
mirifica Michener. Calif. (Inyo and Riverside Cos.). Pollen: Unknown, but visits flowers
Chaenactis brachypappa, C. carphoclinia, Cryptantha, Encelia farinosa.
Anthocopa mirifica Michener, 1954. Pan-Pacific Ent. 30: 51. 2, 6.
namatophila Michener. Calif. (Twentynine Palms). Pollen: Unknown, but visits flowers Nama
demissum.
Anthocopa namatophila Michener, 1954. Pan-Pacifie Ent. 30: 47. °.
nitidivitta Michener. South. and east. Calif., desert. Pollen: Unknown, but visits flowers of
Astragalus fremontii, Dalea fremontii, Mentzelia.albicaulis, Nama demissum, Phacelia.
Anthocopa (Eremosmia) nitidivitta Michener, 1943. Ent. Soc. Amer., Ann. 36: 70. 2, 6.
robustula (Cockerell). South. and east. Calif., Nev. Pollen: Unknown, but visits flowers of
Astragalus coulteri, Cercidium floridum, Cryptantha barbigera, Dalea californica, D.
fremontu, D. polyadenia, D. saundersii, D. schottii, Geraea canescens, Lotus scoparius,
Lupinus odoratus, Palafoxia linearis, Senecio.
Osmia robustula Cockerell, 1935. Pan-Pacifie Ent. 11: 44. 9.

Superfamily APOIDEA 2023

rupestris (Cockerell). South. and east. Calif., desert. Pollen: Unknown, but visits flowers of
Chaenactis stevioides, Eriodictyon trichocalyx, Larrea tridentata, Nama demissum,
Phacelia distans.
Osmia rupestris Cockerell, 1935. Pan-Pacifie Ent. 11: 47. 9.
segregata Michener. Calif. (Inyo Co.). Pollen: Unknown, but visits flowers of Dalea fremontii.
Anthocopa segregata Michener, 1954. Pan-Pacifie Ent. 30: 48. 9.
timberlakei (Cockerell). South. Calif., desert. Pollen: Unknown, but visits flowers of
Astragalus fremontii, Palafoxia linearis.
Osmia timberlakei Cockerell, 1935. Pan-Pacifie Ent. 11: 41. 2.
viguierae (Cockerell). South. Calif., desert border. Pollen: Apparently an oligolege of vernal
flowering Compositae including Encelia farinosa, Enceliopsis argophylla var.
grandiflora, Viguiera parishii, but also visits flowers of Hyptis emoryi.
Osmia viguierae Cockerell, 1935. Pan-Pacifiec Ent. 11: 44. 2 (6 misdet.).

Genus ANTHOCOPA Subgenus ISOSMIA Michener and Sokal

Anthocopa subg. Isosmia Michener and Sokal, 1957. Evolution 11: 159.
Type-species: Anthocopa (Phaeosmia) rubrella Michener. Orig. desig.
hurdiana Michener. Calif. (Panamint Mts.). Pollen: Unknown, but visits flowers of Dalea
fremontii.
Anthocopa hurdiana Michener, 1954. Pan-Pacifie Ent. 30: 47. 2, d.
rubrella macswaini Michener. Calif., Nev. Pollen: Unknown, but visits flowers of Dalea
mollis.
Anthocopa rubrella macswaini Michener, 1954. Pan-Pacific Ent. 30: 45. 2, ¢.
rubrella ere Michener. Tex.; Mexico (Sonora). Pollen: Unknown, but visits flowers of
Dalea.
Anthocopa (Phaeosmia) rubrella Michener, 1949. Kans. Ent. Soc., Jour. 22: 51. 2, ¢.
rubrella rubrior Michener. South. Calif. Pollen: Unknown, but visits flowers of Dalea mollis.
Anthocopa rubrella rubrior Michener, 1954. Pan-Pacific Ent. 30: 46. 2, d.

Genus ASHMEADIELLA Cockerell

Revision: Michener, 1936. Amer. Mus. Novitates 875: 1-16. —Michener, 1939. Amer. Midland
Nat. 22: 1-84 (Nearctic spp.).

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1051, figs. 134, 138, 139, 141 (larva).
—Hurd and Michener, 1955. Calif. Ins.,Survey, Bul. 3: 153-215, pls. 10, 23-24, figs. 2,
112-141, maps 77-111 (Calif. spp.). —Michener and Sokal, 1957. Evolution 11: 130-162
(phenetic classification of the Hoplitis complex). —Sokal and Michener, 1958. Kans. Univ.
Sci. Bul. 38: 1409-1438 (systematic relationships within the Hoplitis complex). —Mitchell,
1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 53-56, figs. 1, 17-18, table 3 (eastern U. S. spp.).
—Rubin, 1966. Syst. Zool. 15: 176-182, tables 1-3 (phenetic classification of the Hoplitis
complex). —Michener and Sokal, 1966. Ent. Soc. Amer., Ann. 59: 1211-1217, 5 figs.
(phenetic similarities among species of the Hoplitis complex).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 36 (Larrea visiting spp.).
Morphology: Michener, 1943. Pan-Pacifiec Ent. 19: 96-100 (sex anomalies).

Genus ASHMEADIELLA Subgenus CHILOSIMA Michener

Ashmeadiella subg. Chilosima Michener, 1939. Amer. Midland Nat. 22: 78.
Type-species: Ashmeadiella rhodognatha Cockerell. Orig. desig.
holtii Cockerell. N. Mex., Ariz., deserts.
Ashmeadiella holtii Cockerell, 1898. Canad. Ent. 30: 51. d.
rhodognatha Cockerell. Tex., Nev. South. and east. Calif.; north. Mexico (Baja California and
Sonora). Pollen: Unknown, but visits flowers of Cercidium torreyanum, Cryptantha
angustifolia, Dalea californica, D. emoryi, D. fremontii, D. neomexicana, D. polyadenia,

2024 Hymenoptera in America North of Mexico

D. schottii, Heliotropium curassavicum, Larrea tridentata, Melilotus, Nama hispidum,
Olneya tesota, Pluchea sericea, Prosopis glandulosa var. torreyanum.
Ashmeadiella rhodognatha Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 557. °.

Taxonomy: Michener, 1942. Ent. News 53: 51. —Michener, 1943. Pan-Pacifie Ent. 19: 96.

Genus ASHMEADIELLA Subgenus ASHMEADIELLA Cockerell

Ashmeadiella Cockerell, 1897. Ent. News 8: 197.
Type-species: Heriades opuntiae Cockerell. Orig. desig.

Titusella Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 445.
Type-species: Titusella pronitens Cockerell. Monotypic.

Taxonomy: Michener and Sokal, 1957. Evolution 11: 159 (synonymy).
altadenae Michener. South. and cent. Calif. Pollen: Unknown, but visits flowers of
Adenostoma fasciculatum, Lotus scoparius.
Ashmeadiella altadenae Michener, 1936. Pan-Pacifie Ent. 12: 63. 6.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 75. @.

aridula aridula Cockerell. Wash., Oreg., Calif., Nev., Utah, Idaho, Wyo., Colo. Parasite:
Anthrax irroratus Say, Chrysura sonorensis (Cam.), Epistenia sp., Leucospis affinis
Say, Sapyga pumila Cress. Pollen: Presumably polylectic, visits a wide variety of
flowers including Chaetopappa aurea, Cleomella obtusifolia, Cryptantha intermedia, C.
Micrantha, Eriogonum gracile, Eriodictyon, Gutierrezia lucida, G. sarothrae,
Heliotropium curassavicum var. oculatum, Lotus americanus, L. argyrophyllus, L.
canadensis, L. davidsonii, L. glaber, L. hamatus, L. nevadensis, Nama parryi, Phacelia
imbricata, P. ramosissima, Prosopis, Sidalcea malvaeflora, Solidago californica,
Sphaeralcea ambigua, Viguiera nevadensis.

Ashmeadiella aridula Cockerell 1910. Entomoligist 43: 91. 6d.

aridula astragali Michener. Wash., Wyo., Oreg., Calif., Nev. Pollen: Presumably polylectic,

visits a wide variety of flowers including Astragalus bolanderi, Calycadenia
multiglandulosa, Centromadia pungens, Cressa cretica, Dalea polyadenia, Euphorbia,
Grindelia camporum, Heliotropium curassavicum var. obovatum, H. c. var. oculatum,
Hemizonia pungens, Lotus americanus, L. glaber, Marrubium vulgare, Melilotus alba,
Phacelia distans, Solidago californica, Trichostema laxuwm, Trifolium.

Ashmeadiella (Ashmeadiella) cactorum astragali Michener, 1939. Amer. Midland Nat. 22:
44.9, 6.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 74 (geogr. records and variation).

Morphology: Snelling, 1962. Pan-Pacifie Ent. 38: 229-230, fig. 1 (intersex).

bigeloviae (Cockerell). Tex., to south. Calif., Nev.; north. Mexico. Ecology: Nests readily in
trap nests. Parasite: Anthrax irroratus Say, Chryswra sonorensis (Cam.), Leucospis
affinis Say, Monodontomerus anthidii (Ashm.), Nemognatha nigripennis LeC.,
Pyemotes ventricosus (Newport). Pollen: Polylectic, visits a wide variety of flowers
including Acacia greggii, Asclepias galioides, Baileya pleniradiata, Bebbia juncea,
Cercidium floridum, C. torreyanum, Cleomella obtusifolia, Coldenia palmeri,
Cryptantha barbigera, C. intermedia, Dalea fremontii, D. mollis, D. spinosa,
Echinocereus engelmannii, Eriastrum virgatum, Eriogonum deflexans, E. deserticola,
E. fasciculatum, E. f. var. polifolium, E. inflatum, E. trichopes, Eucnide urens, Geraea
canescens, Gutierrezia californica, G. lucida, G. sarothrae, Haplopappus acradenius, H.
palmeri, Helenium laciniatum, Heliotropium curassavicum, Hymenopappus flavescens,
Isomeris arborea, Larrea tridentata, Lepidium alyssoides, Lotus davidsonii, Medicago
sativa, Melilotus alba, Monardella exilis, Pectis papposa, Petalonyx thurberi, Phacelia
crenulata, Pyrrhopappus multicaulis, Prosopis, Rhus trilobata, Salvia pilosa, Senecio
douglasii, Sesuvium sessile, Sphaeralcea ambigua, S. orcutti, Stephanomeria pauciflora,
Tamariz gallica, Thelypodium cooperi, Tidestromia oblongiflora, Verbesina auriculata,
Wislizenia refracta. Predator: Trichodes ornatus Say.

Heriades bigeloviae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 136. d, °.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 282-284 (nest architecture,
supersedure, parasite).

Superfamily APOIDEA 2025

biscopula Michener. Ariz., N. Mex. Pollen: Unknown, but visits flowers of Prosopis.
Ashmeadiella (Titusella) biscopula Michener, 1935. Amer. Midland Nat. 22: 54. 9.
bucconis bucconis (Say). N. Dak., south to N. Mex. and Tex., east to Wis., Ill., Ind., Mo., Ark.,
La. and Ga. Pollen: Appears to collect pollen principally from the Compositae, visits
flowers of Ambrosia psilostachya, Anthemis cotula, Brauneria pallida, Callirrhoe
imvolucrata, Coreopsis, Gaillardia pulchella, Grindelia squarrosa, Helenium
laciniatum, Helianthus petiolaris, Heliopsis helianthoides, Lactuca pulchella, Monarda,
Petalostemon, Sideranthus, Solidago, Tetragonotheca ludoviciana, Verbena, Verbesina.
Osmia bucconis Say, 1837. Boston Jour. Nat. Hist. 1: 400. 2, 3.
Megachile osmioides Cresson, 1872. Amer. Ent. Soc., Trans. 4: 269. 2, 6.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 54-55, fig. 17 (redescription).
bucconis denticulata (Cresson). B. C. to Calif., east to Mont., Wyo., Colo., N. Mex. and Tex.;

Mexico. Ecology: Nests readily in trap-nests. Parasite: Anthrax irroratus Say, Chrysura
sonorensis (Cam.), Leucospis affinis Say, Nemognatha nigripennis LeC., Pyemotes
ventricosus (Newport). Pollen: Appears to collect pollen principally from the
Compositae, visits flowers of Acamptopappus sphaerocephalus, Adenostoma
sparsifolium, Aster tephrodes, Baileya multiradiata, B. pleniradiata, Chaenactis
glabriuscula, Chrysothamnus nauseosus, Coreopsis grandiflora, C. lanceolata,
Cryptantha intermedia, Encelia californica, E. farinosa, Eremocarpus setigerus,
Erigeron divergens, E. foliosus var. stenophyllus, Eriogonum gracile, Eriophyllum
confertiflorum, Gnaphalium californicum, Grindelia camporum, G. hallii, G. nana,
Gutierrezia californica, G. lucida, G. sarothrae, Haplopappus acradenius, H.
squarrosus, H. teretifolius, H. venetus, H. vernonioides, Helianthus, Heterotheca
grandiflora, Navarretia viscidula, Pectis papposa, Penstemon, Senecio Hemota ste
Solidago californica, Stephanomeria exigua.

Heriades? denticulatum Cresson, 1878. Amer. Ent. Soc., Trans. 7: 108. 6.

Heriades? rotundiceps Cresson, 1879. Amer. Ent. Soc., Trans. 7: 205. 2.

Ashmeadiella wislizeni Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 545. 6.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 285-286 (nest architecture, life
history, parasites).
cactorum basalis Michener. South. and cent. Calif., west of deserts, also isolated localities in

east Calif. Pollen: Apparently polylectic, visits flowers of Baileya multiradiata,
Castilleia, Chilopsis linearis, Chrysopsis villosa, Chrysothamnus, Cordylanthus nevinii,
C. pilosus, Cryptantha inaequata, C. intermedia, C. micrantha var. lepida, Dalea
fremontii, Encelia farinosa, Erigeron divergens, Eriastrum virgatum, Eriogonum
fasciculatum, E. wrightii subscaposum, Eryngium aristatulum, Grindelia camporum,
Gutierrezia californica, G. sarothrae, Lotus argophyllus, L. davidsonii, L. glaber, L.
nevadensis, L. rosea, L. scoparius, Mimulus, Monardella linoides var. stricta,
Penstemon bridgesii, P. grinnellii, P. ternatus, Phacelia heterophylla, P. imbricata, P.
ramosissima, Salvia pachyphylla, Trichostema lanatum, T. parishii, Verbena
lasiostachys.

Ashmeadiella basalis basalis Michener, 1936. Amer. Mus. Novitates 875: 6. d, @.

cactorum cactorum (Cockerell). B. C., Wash., Oreg., north. and east. Calif., Mont., Wyo., Utah,

Colo., Ariz., N. Mex., Tex.; north. Mexico. Pollen: Stores pollen of Prosopis and small
amounts of Anisacanthus, but is most probably quite polylectic; visits a wide variety of
flowers including Allium parvum, Asclepias tuberosa, Chilopsis linearis, Clarkia
pulchella, C. rhomboidea, Cordylanthus nevini, Cryptantha, Dalea argyraea, D.
pogonothera, Echinocereus, Encelia farinosa, Eriastrum virgatum, Erigeron divergens,
E. stenophyllus, Eriogonum fasciculatum, Lasthenia chrysostoma, Lotus argyrophyllus,
L. scopariws, Monardella stricta, Nama rothrockii, Opuntia, Penstemon ternatus,
Phacelia ramosissima, Trichostema lanatum.

Heriades cactoruwm Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 140. @.

Ashmeadiella curriei Titus, 1904. Ent. Soc. Wash., Proc. 6: 100. 2.

Ashmeadiella echinocerei Cockerell, 1911. Canad. Ent. 43: 132. °.

Ashmeadiella echinocacti(!) Cockerell, 1931. Ann. and Mag. Nat. Hist. (8) 10: 5438.

Ashmeadiella basalis nigra Michener, 1936. Amer. Mus. Novitates 875: 7. 2, d.

2026 Hymenoptera in America North of Mexico

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 286-288 (nest architecture, larval
food, life history, cocoon).
californica californica (Ashmead). B. C. to Calif. (except deserts and Sierra Nevada Mts.),

Idaho, Nev., Utah; Mexico (Baja California). Parasite: Anthrax irroratus Say, Chrysura
sp., Leucospis affinis Say, Sphaeropthalma wnicolor (Cress.), Stelis ashmeadiellae
Timberlake. Pollen: Apparently polylectic, visits a wide variety of flowers including
Asclepias, Aster adscendens var. yosemitanus, Calycadenia multiglandulosa,
Camissonia campestris, Centaurea melitensis, Chaenactis glabriuscula, Chorizanthe
staticoides, Chrysothamnus, Corethrogyne, Cressa cretica, Cryptantha intermedia,
Encelia californica, E. farinosa, Eriastrum virgatum, Erigeron divergens, E. foliosus
var. stenophyllus, Eriogonum fasciculatum, E. subscaposum, Eriophyllum
confertiflorum, Eupatorium occidentale, Frankenia grandiflora, Gilia multicaulis,
Grindelia camporum, G. elata, Gutierrezia californica, G. sarothrae, Haplopappus
linearifolius, H. parishii, H. squarrosus, H. vernonioides, Helianthus gracilentus,
Hemizonia fasciculata, H. luzulaefolia, H. paniculata, H. wrightii, Heterotheca
grandiflora, Lessingia germanorum, L. grandiflora, L. leptoclada, L. tenuis, Lotus
americanus, L. glaber, L. hamatus, L. scoparius, Medicago sativa, Melilotus,
Monardella douglasii, M. stricta, Navarretia heterodoxa, N. viscidula, Pentachaeta
aurea, Phacelia ciliata, P. davidsonii, Potentilla glandulosa, Senecio douglasii,
Sesuvium sessile, Solidago lasiostachys, Stephanomeria exigua, Verbena prostrata,
Viguiera.

Chalicodoma californica Ashmead, 1897. South. Calif. Acad. Sci., Proc. 1 (3): 1. d, 2.

Ashmeadiella coquilletti Titus, 1904. Ent. Soc. Wash., Proc. 6: 99. 2, d.

Biology: Timberlake, 1941. N. Y. Ent. Soc., Jour. 49: 134 (parasite).
californica florissantensis Michener. Idaho. Wyo., Colo., N. Mex. Pollen: Unknown, but visits
flowers of Aster, Chrysolepis.
Ashmeadiella florissantensis Michener, 1936. Pan-Pacifie Ent. 12: 62. d, °.
californica sierraensis Michener. Calif. (Sierra Nevada Mts.). Pollen: Apparently polylectic,
principally flowers of Compositae, including Aster adscendens, A. foliaceus, Erigeron,
Helianthus, Senecio canus, Solidago multiradiata.
Ashmeadiella (Ashmeadiella) californica sierraensis Michener, 1939. Amer. Midland Nat.
Zo Olea enor
cockerelli Michener. South. Calif.
Ashmeadiella cockerelli Michener, 1936. Pan-Pacific Ent. 12: 62. d.
cubiceps clypeata (Michener). Calif., low and middle altitudes; Mexico (Baja California).
Parasite: Chrysura sp., Leucospis affinis Say, Monodontomerus anthidii (Ashm.).
Pollen: Apparently polylectic, principally Compositae, visits flowers of Baileya,
Chaenactis, Cryptantha, Encelia, Eriophyllum confertiflorum, Haplopappus
linearifolius, Hemizonia, Lessingia germanorum, Melilotus, Sphaeralcea, Viguiera.
Titusella clypeata Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 93. °.
cubiceps cubiceps (Cresson). Nev., Calif., Oreg., high mts.
Heriades? cubiceps Cresson, 1879. Amer. Ent. Soc., Trans. 7: 205. °.
difugita difugita Michener. South. Calif. Pollen: Unknown, but visits flowers of Chrysopsis
fastigiata, Eriodictyon trichocalyx, Eriogonum fasciculatum, E. inflatum, Gilia,
Phacelia heterophylla, Verbena lasiostachys.
Ashmeadiella (Ashmeadiella) difugita Michener, 1939. Amer. Midland Nat. 22: 41. 9, d.
difugita emarginatula Michener. Calif. (Sierra Nevada Mts.), Oreg., Idaho, Nev. Pollen:
Unknown, but visits flowers of Camissonia clavaeformis citrina, C. tanacetifolia,
Chrysothamnus nauseosus, Cirsium tioganum, Clarkia, Eriogonum, Grindelia,
Haplopappus apargoides.
Ashmeadiella difugita emarginatula Michener, 1951. Kans. Ent. Soc., Jour. 24: 53. 2.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 74 (geogr. and floral records).

dimalla Michener. Ariz.
Ashmeadiella (Ashmeadiella) dimalla Michener, 1939. Amer. Midland Nat. 22: 45. 9, d.

Superfamily APOIDEA 2027

femorata (Michener). South. and east. Calif., Nev., Ariz., deserts. Pollen: Unknown, but visits
flowers of Baileya, Cercidium floridum, C. torreyanum, Croton californicus, Dalea
fremontu, Haplopappus interior, Hyptis emoryi, Larrea tridentata, Prosopis pubescens.
Predator: Cymatodera sp.
Osmia femorata Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 91. 9.
floridana (Robertson). N. C. to Fla. Pollen: Unknown, but visits flowers of Lupinus.
Heriades floridanus Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 348. 9.

Taxonomy: Mitchell, 1962. N. C. Agr. Exp. Sta. Tech. Bul. 152: 55-56, fig. 18, table 3
(redescription).
foveata Michener. Cent. and east. Calif., Ariz., Nev., Utah. Ecology: Nests under stones.

Pollen: Unknown, but visits flowers of Collinsia torreyi, Cryptantha intermedia, Dalea,
Diplacus auriantiacus, Encelia, Erigeron miser, Haplopappus interior, Helianthus
gracilentus, Heliotropium curassavicum var. oculatum, Lotus scoparius, Monardella
lanceolata, Phacelia platyloba, Prosopis, Salvia pilosa.

Ashmeadiella (Ashmeadiella) foveata Michener, 1939. Amer. Midland Nat. 22: 39. 2.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 74. 3.
gillettei cismontanica Michener. Calif. (Riverside, Fresno, Contra Costa Cos.). Pollen:
Unknown, but visits flowers of Cryptantha intermedia, Eriastrum virgatum,
Eriogonum, Heliotropium curassavicum var. oculatum, Melilotus.
Ashmeadiella (Ashmeadiella) gillettei cismontanica Michener, 1951. Pan-Pacifie Ent. 27:
Gis

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 24: 52. d.
gillettei gillettei Titus. Colo., N. Dak., S. Dak., Nebr. Pollen: Unknown, but visits flowers of
Sedum stenopetalum.
Ashmeadiella gillettei Titus, 1904. Ent. Soc. Wash., Proce. 6: 100. 9.
Ashmeadiella coloradensis Cockerell, 1934. Amer. Mus. Novitates 732: 4. 3.
gillettei rubra Michener. Tex., N. Mex. Pollen: Unknown, but visits flowers of Dalea formosa.
Ashmeadiella (Ashmeadiella) gillettei rubra Michener, 1951. Pan-Pacifie Ent. 27: 67. 2, 3.
gillettei rufiventris Michener. South. and east. Calif.; Mexico (Baja California). Parasite:
Anthrax irroratus Say, Chrysura sp., Leucospis affinis Say, Stelis sp. Pollen: Unknown,
but visits flowers of Aster abatus, Astragalus fremontii, Baileya pleniradiata,
Cercidium floridum, C. torreyanum, Chaenactis carphoclinia, Cryptantha, Dalea
mollis, Geraea canescens, Heliotropium curassavicum, Lupinus, Malacothrix, Palafoxia
linearis, Phacelia distans, Prosopis glandulosa var. torreyanum, Stephanomeria.
Predator: Cymatodera sp.
Ashmeadiella (Ashmeadiella) rufiventris Michener, 1939. Amer. Midland Nat. 22: 32. 2, 6.
leucozona Cockerell. South. Calif., Ariz.; north. Mexico, deserts. Pollen: Unknown, but visits
flowers of Cercidium torreyanum, Heliotropium curassavicum var. oculatum, Nama
hispidum, Teucrium depressum.
Ashmeadiella lewcozona Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 556. d, 2.
maxima Michener. Tex.
Ashmeadiella maxima Michener, 1936. Pan-Pacific Ent. 12: 61. 2, ¢.
meliloti meliloti (Cockerell). South. Calif., Ariz., N. Mex., Tex.; north. Mexico; deserts.
Parasite: Anthrax irroratus Say, Chrysura sp., Epistenia sp., Leucospis affinis Say,
Nemognatha nigripennis LeC., Sapyga pumila Cress., Stelis sp. Pollen: Apparently
polylectic, analyzed pollen masses (2) ranged from 100 per cent Prosopis to a mixture of
several pollens consisting of 74 per cent Stemodia and 26 per cent Chenopodiaceae with
traces of pollen from Acacia, Krameria, entomophilous Compositae and Pinus; visits a
wide variety of flowers including Acacia greggii, Acamptopappus sphaerocephalus,
Baileya multiradiata, Chilopsis linearis, cleome, Cryptantha, Crusea subulata, Dalea
argyraea, D. formosa, D. pogonathera, Eriogonum gracile, E. inflatum, Eriastrum
virgatum, Gutierrezia californica, G. lucida, Haplopappus interior, Heliotropium
curassavicum var. oculatum, Lepidium alyssoides, L. montanum, Lotus davidsonii,
Melilotus alba, Opuntia megacarpa, Penstemon, Phacelia cicutaria, P. ramosissima,
Prosopis glandulosa var. torreyanum, Pyrrhopappus multicaulis, Rhus trilobata,

2028 Hymenoptera in America North of Mexico

Salvia pilosa, Solidago californica, Sphaeralcea ambigua, Viguiera nevadensis.
Predator: Cymatodera sp., Trichodes horni Wolcott and Chapin. Another subspecies,
crassa Ckll., occurs in Mexico (Baja California).

Heriades meliloti Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 141. d, 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 288-290 (nest architecture, larval
food, life history, supersedure, parasites and predators).
occipitalis Michener. South. Ariz., west. Tex.; Mexico (Sonora, Baja California). Ecology:
Readily nests in trap-nests. Parasite: Chrysura sonorensis (Cam.), Nemognatha
nigripennis LeC. Predator: Pyemotes ventricosus (Newport), Trichodes horni Wolcott
and Chapin.
Ashmeadiella (Ashmeadiella) occipitalis Michener, 1939. Amer. Midland Nat. 22: 22. 2, 6.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 76 (geogr. records, tax. characters of
female).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 290-294, pl. 17, fig. 78 (nest
architecture, larval food, life history, male production, supersedure, parasites, predators).
opuntiae (Cockerell). Tex. to south. Calif., Utah and Colo.; north. Mexico (Sonora). Parasite:
Anthrax irroratus Say, Chrysura sp., Leucospis affinis Say, Melittobia chalybii Ashm.
Pollen: Oligolege of Cactaceae, analyzed pollen masses (4) contained 95.2 per cent
Opuntia pollen of the subgenus Cylindropuntia and 4.8 per cent pollen of the subgenus
Platyopuntia; obtains pollen and nectar from flowers of Cactaceae including
Echinocactus cylindraceus, Echinocereus engelmannii, Ferrocactus acanthodes,
Opuntia echinocarpa, O. megacarpa, O. occidentalis, O. parryi, O. vaseyi, but also visits
other flowers for nectar including Chrysothamnus, Penstemon, Viguiera nevadensis.
Predator: Cymatodera sp.
Heriades opuntiae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 139. 2.
Ashmeadiella submaxima Michener, 1936. Amer. Mus. Novitates 875: 3. 6.
Ashmeadiella arizonensis Michener, 1936. Amer. Mus. Novitates 875: 3. 6.

Taxonomy: Michener, 1943. Pan-Pacifie Ent. 19: 97. —Hurd and Michener, 1955. Calif. Ins.
Survey, Bul. 3: 192 (synonymy).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 294-295 (nest architecture, larval
food, life history, parasite).
pronitens (Cockerell). Colo., Wyo.
Titusella pronitens Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 446. 9.
prosopidis (Cockerell). Tex. to south. Calif.; north. Mexico, deserts. Pollen: Apparently an
oligolege of Prosopsis including P. glandulosa var. torreyanum, P. pubescens, but visits
other flowers presumably for nectar including Acacia greggii, Cryptantha barbigera,
Heliotropium curassavicum, Larrea tridentata, Melilotus, Pluchea sericea, Salix nigra,
Sphaeralcea ambigua.
Heriades prosopidis Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 20: 140. 2 (6 misdet.).
Ashmeadiella schwarzi Titus, 1904. Ent. Soc. Wash., Proc. 6: 98. 2, ¢.
Ashmeadiella subangusta Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 588. °.

Taxonomy: Hurd and Michener, 1955. Calif. Ins. Survey, Bul. 3: 161 (synonymy).
rufipes Titus. South. and east. Calif., north. Mexico, deserts. Parasite: Monodontomerus

anthidii (Ashm.), Stelis sp. Pollen: Unknown, but visits flowers of Abronia, Cryptantha
barbigera, C. intermedia, Dalea emoryi, D. mollis, Heliotropium curassavicum, H. c.
var. oculatum, Palafoxia linearis, Pectis papposa, Petalonyx thurberi, Tidestromia
oblongiflora.

Ashmeadiella rufipes Titus, 1904. Ent. Soc. Wash., Proc. 6: 99. °.

Ashmeadiella haematopoda Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 555. 9.

Ashmeadiella rhodopus Michener, 1936. Pan-Pacific Ent. 12: 59. 6, 9.

Taxonomy: Michener, 1936. Pan-Pacific Ent. 12: 58. 2, 6. —Hurd and Michener, 1955. Calif.
Ins. Survey Bul. 3: 175-177 (synonymy).

Superfamily APOIDEA 2029

rufitarsis Michener. Calif. Pollen: Apparently an oligolege of Eriogonum, including LE.
fasciculatum, E. gracile, E. vimineum, but also visits other flowers presumably for
nectar including Euphorbia.
Ashmeadiella (Ashmeadiella) rufitarsis Michener, 1939. Amer. Midland Nat. 22: 37. 2, d.
sonora Michener. Calif., Ariz., Utah; north. Mexico (Baja California). Pollen: Unknown, but
visits flowers of Asclepias, Baileya multiradiata, Dalea, Eriastrum virgatum,
Eriogonum trichopes, Euphorbia polycarpa var. hirtella, Fouquieria splendens,
Gutierrezia sarothrae, Lepidium alyssoides, Olneya tesota, Pectis papposa.
Ashmeadiella (Ashmeadiella) sonora Michener, 1939. Amer. Midland Nat. 22: 35. d, @.
stevensi Michener. N. Dak.
Ashmeadiella stevensi Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 405. °.
titusi Michener. Calif. Pollen: Possibly an oligolege of Lotus including L. scoparius, but visits
other flowers presumably for nectar including Helianthus gracilentus, Phacelia
ramosissima.
Ashmeadiella (Ashmeadiella) tituwsi Michener, 1939. Amer. Midland Nat. 22: 25. 2, d.

Taxonomy: Snelling, 1962. Pan-Pacific Ent. 38: 230 (geogr. and floral records).
truncativentris Michener. Tex. (Sanderson). Pollen: Unknown, but visits flowers of Dalea.
Ashmeadiella (Ashmeadiella) truncativentris Michener, 1951. Pan-Pacific Ent. 27: 68. 2,
é.
vandykiella Michener. Tex.
Ashmeadiella (Ashmeadiella) vandykiella Michener, 1949. Kans. Ent. Soc., Jour. 22: 46. 2,
é.

Genus ASHMEADIELLA Subgenus AROGOCHILA Michener

Ashmeadiella subg. Arogochila Michener, 1939. Amer. Midland Nat. 22: 58.
Type-species: Ashmeadiella timberlakei Michener. Orig. desig.
Ashmeadiella subg. Corythochila Michener, 1939. Amer. Midland Nat. 22: 74.
Type-species: Ashmeadiella inyoensis Michener. Orig. desig.
Rhamphorhina Michener, 1939. Amer. Midland Nat. 22: 8. Republished by Michener and
Sokal, 1957. Evolution 11: 185 and 159, lapsus for Corythochila Michener.
australis (Cockerell). Calif., Nev., Utah. Pollen: Apparently an oligolege of Penstemon
including P. breviflorus, P. grinnellii, P. palmeri, but visits other flowers presumably
for nectar including Geranium, Monardella linoides var. stricta, Solidago californica,
Streptanthus tortuosus.
Chelostoma australis Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 139. 9.
Chelostomopsis australis nanus Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 206. °.
Taxonomy: Snelling, 1962. Pan-Pacifie Ent. 38: 234 (geogr. and floral records).
barberi Michener. Ariz.
Ashmeadiella (Arogochila) barberi Michener, 1939. Amer. Midland Nat. 22: 61. °.
breviceps Michener. South. Calif., Ariz., Nev.; Mexico (Baja Calif., Sonora). Parasite: Chryswra
sp., Stelis sp. Pollen: Apparently polylectic, visits a wide variety of flowers including
Cercidium floridum, Dalea mollis, Dipetalia linifolia, Hyptis emoryi, Larrea tridentata,
Prosopis glandulosa var. torreyanum, Stephanomeria. Predator: Cymatodera sp.
Ashmeadiella (Corythochila) breviceps Michener, 1939. Amer. Midland Nat. 22: 77. @, 6.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 36 (floral relationships).
cazieri Michener. South. and east. Calif., Ariz., deserts. Parasite: Nemognatha nigripennis
LeC., Stelis sp. Pollen: Apparently polylectic, visits flowers of Cercidium torreyanum,
Dalea californica, D. fremontii, D. johnsonti, Larrea tridentata.
Ashmeadiella (Arogochila) cazieri Michener, 1939. Amer. Midland Nat. 22: 72. 9, 6.
clypeodentata clypeodentata Michener. Ariz., south. Calif., north. Mexico, deserts. Pollen:
Apparently polylectic, visits flowers of Cercidium floridum, C. torreyanum, Dalea
emoryi, Heliotropium curassavicum var. oculatum, Phacelia hispida, Prosopis

glandulosa var. torreyanum.
Ashmeadiella clypeodentata Michener, 1936. Pan-Pacifie Ent. 12: 57. 2.

2030 Hymenoptera in America North of Mexico

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 295 (nest architecture, life
history).
clypeodentata simplicior Michener. Tex., Ariz. Pollen: Unknown, but visits flowers of
Prosopis.
Ashmeadiella (Arogochila) clypeodentata simplicior Michener, 1951. Pan-Pacific Ent. 27:
70. @.
erema Michener. Tex., Ariz., south Calif., deserts. Pollen: Unknown, but visits flowers of Dalea
including D. fremontii.
Ashmeadiella (Arogochila) erema Michener, 1939. Amer. Midland Nat. 22: 65. 9.
eurynorhyncha Michener. East. Calif., desert. Pollen: Unknown, but visits flowers of Dalea
fremontii.
Ashmeadiella (Arogochila) eurynorhyncha Michener, 1939. Amer. Midland Nat. 22: 62. ?,
3.
foxiella Michener. Wash., Oreg., cent. Calif., Idaho. Pollen: Unknown, but visits flowers of
Penstemon, Phacelia.
Ashmeadiella (Arogochila) foxiella Michener, 1939. Amer. Midland Nat. 22: 73. 3.
Ashmeadiella (Chilosima) Washingtonensis Michener, 1939. Amer. Midland Nat. 22: 80. °.
inyoensis Michener. East. Calif., deserts. Pollen: Unknown, but visits flowers of Dalea
polyadenia, D. schottii.
Ashmeadiella (Corythochila) inyoensis Michener, 1939. Amer. Midland Nat. 22: 75. 2, d.
lateralis Michener. South. Calif.
Ashmeadiella lateralis Michener, 1936. Pan-Pacifie Ent. 12: 60. 6.
leachi Michener. Calif.
Ashmeadiella (Arogochila) leachi Michener, 1949. Kans. Ent. Soc., Jour. 22: 43. 2.
lutzi (Cockerell). Utah, Colo. Pollen: Unknown, but visits flowers of Phacelia leucophylla.
Chelostomopsis lutzi Cockerell, 1930. Amer. Mus. Novitates 397: 2. 9°.
micheneri Snelling. Calif. (Stanislaus and Mariposa counties). Pollen: Unknown, but visits
flowers of Phacelia.
Ashmeadiella (Arogochila) micheneri Snelling, 1962. Pan-Pacific Ent. 38: 231, fig. 2. d, ?.
neomexicana (Cockerell). N. Mex.
Chelostoma Neomexicanum Cockerell, 1904. Canad. Ent. 36: 13. 9.
salviae Michener. South. and cent. Calif. Pollen: Apparently polylectic, visits flowers of
Brassica, Calochortus luteus, Dalea, Lepechinia calycina, Rhamnus crocea, Salvia
mellifera, Trichostema parishii.
Ashmeadiella (Arogochila) salviae Michener, 1939. Amer. Midland Nat. 22: 69. 2, 3.
sculleni Michener. Oreg.
Ashmeadiella (Arogochila) sculleni Michener, 1939. Amer. Midland Nat. 22: 60. °.
stenognatha Michener. Calif. Pollen: Unknown, but visits flowers of Phacelia platyloba.
Ashmeadiella (Arogochila) stenognatha Michener, 1939. Amer. Midland Nat. 22: 63. °.

Taxonomy: Michener, 1954. Kans. Ent. Soc., Jour. 27: 77. 3.
timberlakei solida Michener. Cent. and north. Calif., Oreg., Nev. Pollen: Apparently polylectic,
visits flowers of Astragalus bolanderi, Eriodictyon, Grindelia camporum, Kelloggia
galioides, Lotus argophyllus, L. glaber, L. nevadensis, L. scoparius, Nama rothrockii,
Nemophila, Phacelia distans, Solidago californica, Trifolium variegata.
Ashmeadiella (Arogochila) timberlakei solida Michener, 1939. Amer., Midland Nat. 22: 68.
Gy Oe
timberlakei timberlakei Michener. South. Calif. Parasite: Anthrax irroratus Say, Leucospis
affinis Say, Nemognatha nigripennis LeC., Rhydinofoenus sp., Stelis sp. Pollen:
Apparently polylectic, visits flowers of Chaenactis, Chorizanthe staticoides, Cryptantha
intermedia, C. lepida, Dalea californica, Eriodictyon crassifolium, E. trichocalyx, Lotus
argophyllus, L. davidsonii, L. glaber, L. nevadensis, L. scoparius, Lupinus arizonicus, L.
concinnus, Mimulus fremontii, Oenothera dentata, Penstemon spectabilis, Phacelia
cicutaria, P. davidsonii, P. heterophylla, P. ramosissima, Trichostema lanata, T.
parishi, Trifolium variegata.
Ashmeadiella timberlakei Michener, 1936. Pan-Pacifie Ent. 12: 56. 2, do.

Superfamily APOIDEA 2031

Genus ASHMEADIELLA Subgenus CUBITOGNATHA Michener

Ashmeadiella subg. Cubitognatha Michener, 1939. Amer. Midland Nat. 22: 81.
Type-species: Ashmeadiella xenomastax Michener. Orig. desig. and monotypic.
xenomastax Michener. South. and east. Calif., Nev.; Mexico (Sonora); deserts. Pollen:

Apparently polylectic, visits flowers of Aster abatus, Chaenactis xantiana, Dalea
californica, D. mollis, D. polyadenia, D. saundersii, D. schottii, Eriogonum
fasciculatum, Lotus, Lupinus odoratus, Phacelia distans, Pholisma arenarium, Salvia
columbariae.

Ashmeadiella (Cubitognatha) xenomastax Michener, 1939. Amer. Midland Nat. 22: 81. 2.

Taxonomy: Michener, 1942. Ent. News 53: 51. ¢.

Genus OSMIA Panzer

Revision: Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 1-167 (Nearctic spp.). —Rust, 1974.
Wasmann Jour. Biol. 32: 1-93, 45 figs., 9 tables (treats Nearctic spp. of the subgenera
Cephalosmia, Chalcosmia and Osmia and includes information on their biologies).

Taxonomy: Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 357-364 (key to western spp.).
—Sandhouse, 1925. Canad. Ent. 57: 35-41 (key to Canad. spp.). —Michener, 1941. Amer.
Midland Nat. 26: 147-167 (classification). —Sinha, 1958 (1956). Xth Internatl. Congress
Ent., Proc. 1: 243-251, 1 fig., 4 tables (phylogenetic relationships). —Sinha, 1958. Kans.
Univ. Sci. Bul. 39: 211-261, 151 figs. (reclassification of New World subgenera). —Mitchell,
1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 69-107, figs. 1, 25-35, table 4 (eastern WES
spp.). —Snelling, 1967. South. Calif. Acad. Sci., Bul. 66: 103-108, 1 fig. (treats subgenera
Euthosmia and Mystacosmia).

Biology: Rau, 1937. Ent. Soc. Amer., Ann. 30: 324-343, 6 figs., 2 tables (nesting habits).
—Bohart, 1955. Ent. Soe. Wash., Proc. 57: 203-204 (gradual nest supersedure).
—Crosswhite and Crosswhite, 1966. Amer. Midland Nat. 76: 450-467 (Penstemon visiting
spp.). —Krombein, 1967. Trap-nesting wasps and bees, pp. 295-320, pl. 16, figs. 70-77, pl. 17,
figs. 82-85, 87, pl. 18, figs. 88-91, pl. 23, figs. 111-114, pl. 27, figs. 128-130, tables 29-33 (life
histories, nests, assa¢iates). —Medler, 1967. Ent. Soc. Amer., Ann. 60: 338-344, figs. 1-2, 2
tables (life histories, nests and associates of Wis. spp.). —Maeta, 1969. The life study,
Fukui 13: 41-43 (supersedure). —Rust, 1974. Wasmann Jour. Biol. 32: 1-93, 45 figs., 9 tables
(life histories, nests and associates of the subgenera Cephalosmia, Chalcosmia and
Osmia).

Genus OSMIA Subgenus OSMIA Panzer

Osmia Panzer, 1806. Krit. Rev. Insektenf. Deutschlands, v. 2, p. 230.

Type-species: Apis rufa Linnaeus. Desig. by Latreille, 1810. (=Apis bicornis
Linnaeus).

Amblys Klug, 1807. Mag. Insektenk. 6: 198.
Type-species: Apis rufa Linnaeus. Desig. by Latreille, 1810.

Osmia subg. Ceratosmia Thomson, 1872. Hym. Seand., v. 2, p. 2382.
Type-species: Apis rufa Linnaeus. Monotypic.

Osmia subg. Aceratosmic Schmiedeknecht, [1885]. Apidae Europaeae, v. 2, p. 19.
Type-species: Osmia emarginata Lepeletier. Desig. by Sandhouse, 1939.

Osmia subg. Pachyosmia Ducke, 1900. Ber. naturw.-med. Ver. Innsbruck 25: 18.
Type-species: Osmia rufa (Linnaeus). Desig. by Sandhouse, 1939.

Revision: Rust, 1974. Wasmann Jour. Biol. 32: 8-82, figs. 1-2, 7-15, tables 1-3 (Nearctic spp.).

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 9-11 (summary of information on life histories).
cornifrons (Radoszkowski). Utah (Garden City and Providence Canyon); China, Korea, Japan.
Ecology: Nests in preexisting burrows, holes or cavities in wood, stems or culms

including trap-nests. Purposely introduced into Utah from Morioka, Japan in 1965 and
possibly is an established species. Parasite: Tricran ia stansbury? (Hald.). Pollen:
Apparently polylectic, visits flowers of Malus, Prunus, Pyrus.

Chalicodoma cornifrons Radoszkowski, 1887. Soc. Ent. Ross., Horae Zl F430nne-

2032 Hymenoptera in America North of Mexico

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 14-15 (nest architecture, cocoon, life history,
parasite).
lignaria lignaria Say. N. S. to Ga., west to Mich., Iowa, Nebr., Kans., Okla., and Tex. Ecology:

Nests in a wide variety of preexisting borings, burrows, cavities, crevices, and mud
nests of wasps as well as attaching their cells in clusters on exposed surfaces; uses mud
for nest construction. Parasite: Chaetodactylus krombeini Baker, Chrysura kyrae
Krombein, Leucospis affinis Say, Melittobia chalybii Ashm., Monodontomerus obscurus
Westw., Sapyga angustata Cress. Pollen: Polylectic, visits a wide variety of flowers
including Camassia, Cardamine, Cercis canadensis, Chaerophyllum, Claytonia,
Collinsia, Dentaria laciniata, Dicentra, Diervilla japonica, Dirca, Erigenia,
Erythronium albidum, E. americanum, Fragaria, Geranium, Glechoma hederacea,
Hydrophyllum, Lupinus, Malus, Mertensia, Nepeta hederacea, Osmorrhiza, Oxalis,
Polemonium, Prunus, Pyrus, Ranunculus, Ribes, Rosa, Rubus, Salix, Senecio,
Stellaria, Taenidia, Taraxacum, Viburnum, Vicia, Viola, Zanthoxylum.

Osmia lignaria Say, 1837. Boston Jour. Nat. Hist. 1: 399. 2, 3.

Osmia purpurascens Smith, 1849. Zoologist 7 (app.): lviii. ¢.

Biology: Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 203 (nest). —Rau, 1937. Ent. Soc. Amer.,
Ann. 30: 324-342, 1 pl. (nest architecture, life history, effect of cell size and placement on
sex ratio, foraging behavior, parasite). —Bohart, 1955. Ent. Soc. Wash., Proce. 57: 203-204
(nest architecture, life history, supersedure). —Chandler, 1959 (1958). Ind. Acad. Sci., Proc.
68: 199-204 (nest). —Balduf, 1961. Brooklyn Ent. Soc., Bul. 56: 83-84 (nest, foraging
behavior). —Krombein, 1962. Biol. Soc. Wash., Proc. 75: 239-242, pl. 1 (nest architecture,
parasite). —Chandler, 1962. Ent. Soc. Amer., North Central Branch, Proe. 16: 18-19 (nest
architecture, life history, parasite). —Krombein, 1967. Trap-nesting wasps and bees, pp.
296-306; pl. 16, figs. 70-77; pl. 17, figs. 82-85, 87; pl. 18, fig. 88; pl. 23, figs. 111-114; pl. 27,
figs. 128-130 (nest architecture, life history, supersedure, effect of cell size and placement
on sex ratio, foraging behavior, cocoon, parasites). —Medler, 1967. Ent. Soc. Amer., Ann.
60: 340-341, fig. 2, table 1 (nest architecture, life history). —Byers, 1972. Kans. Ent. Soc.,
Jour. 45: 235-238 (supersedure by Monobia quadridens). —Matthews and Kislow, 1973.
Environ. Ent. 2: 157-158, 1 fig. (cocoon orientation). —Rust, 1974. Wasmann Jour. Biol. 32:
21-27, table 3 (nest architecture, life history, parasites). —Torchio, 1976. Kans. Ent. Soc.,
Jour. 49: 475-482, 4 figs., 4 tables (use as pollinator in apple and prfine orchards).

lignaria propinqua Cresson. Sask., to west. Tex., west to B. C., Wash., Oreg. and Calif.

Ecology: Nests in a wide variety of preexisting borings, burrows, cavities, crevices and
mud nests of wasps as well as attaching their cells in clusters on exposed surfaces; uses
mud for nest construction. Parasite: Lewcospis affinis Say, Sapyga emarginata Cress.,
Stelis sp., Tricrania stansburyi Hald. Pollen: Polylectic,visits a wide variety of flowers
including Acer, Amsinckia intermedia, Arbutus menziesii, Arctostaphylos drupacea, A.
glauca, A. patula, Astragalus, Berberis californica, Blennosperma nanum, Brassica,
Ceanothus integerrimus, Cercis occidentalis, Cercocarpus betulifolius, Cirsiwm,
Clarkia, Collinsia heterophylla, Cryptantha, Cytisus scoparius, Eriodictyon,
Erythronium, Geraea canescens, Gilia tricolor, Hydrophyllum, Isomeris arborea,
Limnanthes douglasii, Lonicera, Lupinus albifrons, L. bicolor, Lycium, Mimulus,
Nemophila exilis, N. menziesti, Penstemon, Phacelia cicutaria, P. distans, P.
tanacetifolia, Prunus subcordata, Ranunculus californicus, Raphanus sativus,
Rhamnus californica, R. crocea, Ribes cereum, R. menziesii, R. roezlii, R. velutina, R.
viscosissimum, Rubus ursinus, Salix laevigata, S. lasiolepis, Salvia carduacea, Senecio,
Sisymbrium irio, Tamarix, Taraxacum vulgare, Trifolium repens, Vicia californicum,
Viola, Wyethia. Predator: Ptinus californicus Pic., Trichodes ornatus Say.

Osmia propinqua Cresson, 1864. Ent. Soc. Phila., Proc. 3: 23. 2.

Osmia lignaria lignariella Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 227. 2.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1054, figs. 135-137, 140 (larva).

Biology: Hicks, 1934. Colo. Univ., Studies 21: 265 (nest). —Linsley and MacSwain, 1941.
South. Calif. Acad. Sci., Bul. 40: 129 (nest, parasites). —Levin, 1957. Econ. Ent., Jour. 50:
506-507 (nest architecture, life history). —Levin and Haydak, 1957. Bee World 38: 221-226
(comparative nutritional values of different pollens). —Levin, 1966. Kans. Ent. Soc., Jour.

Superfamily APOIDEA 2033

39: 524-535, figs 2-6 (nest architecture, life history, cocoon, effect of cell size and

placement on sex ratio, mud collecting, foraging behavior, parasites, predators). —Rust,

1974. Wasmann Jour. Biol. 32: 21-27, table 3 (nest architecture, life history, mating

behavior, parasites, predators).

ribifloris biedermannii Michener. Oreg., Calif., Nev., Utah, Ariz., west. N. Mex.; Mexico (Baja

California). Ecology: Nests in borings, abandoned Sceliphron nests and between cracks
in shingles. Pollen: Polylectic, visits flowers of Arctostaphylos drupacea, A. glandulosa,
A. glauca, A. patula, Asclepias, Astragalus, Berberis, Eriodictyon crassifolium,
Fendlera, Phlox, Prunus, Rhus, Ribes, Salix, Salvia sonomensis, Wyethia.

Osmia ribifloris biedermannii Michener, 1936. Amer. Mus. Novitates 875: 19. 9, 6.

Biology: Leech, 1959. Pan-Pacifie Ent. 35: 53 (nest). —Krombein, 1967. Trap-nesting wasps
and bees, p. 306 (nest architecture, life history, as ribifloris). —Rust, 1974. Wasmann
Jour. Biol. 32: 32 (nest architecture, foraging behavior).

ribifloris ribifloris Cockerell. Tex., N. Mex., Colo., Utah, Nev. Pollen: Unknown, but visits
flowers of Berberis, Sophora.
Osmia (Melanosmia) ribifloris Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 410. 2.

Genus OSMIA Subgenus CHALCOSMIA Schmiedeknecht

Osmia Subg. Chalcosmia Schmiedeknecht, [1885]. Apidae Europaeae, v. 2, p. 886.
Type-species: Osmia fulviventris Latreille. Desig. by Sandhouse, 1939.
Gnathosmia Robertson, 1903. Amer. Ent. Soc., Trans. 29: 165.
Type-species: Osmia georgica Cresson. Orig. desig. and monotypic.

Revision: Rust, 1974. Wasmann Jour. Biol. 32: 32-56, figs. 3-4, 15-26, tables 4-6.

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 34 (summary of information on life histories).
chalybea Smith. N. J., N. C., Ga., Fla., La., Tex. Pollen: Unknown, but visits flowers of
Ceanothus, Cirsium, Vaccinium.
Osmia chalybea Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 148. 2, 6.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 33. d.
coerulescens (Linnaeus); Holarctic. N. S. to Minn. south to N. C., Ind., Il., Mo., and Neb.;
Idaho (Twin Falls), where probably adventive. Ecology: Nests in preexisting burrows in
wood or in open cavities as in old mud dauber nests. Parasite: Melittobia chalybii
Ashm., Monodontomerus sp. Pollen: Collects pollen from Leonurus cardinea, Medicago
sativa, Nepeta cataria, but visits other flowers including Agastache, Cercis, Cirsium,
Lotus, Philadelphus, Salvia, Trifolium, Vicia. Predator: Attagenus piceus Oliv.,
Chaetodactylus sp., Trogoderma inclusum LeC., T. teukton Beal.
Apis coerulescens Linnaeus, 1758. Syst. Nat., ed. 10, p. 576. °.
Apis aenea Linnaeus, 1761. Fauna Suecica, ed. 2, p. 421. d.
Osmia cyanea Giraud, 1866. Soc. Ent. France, Ann. 6: 451. 9, d.
Osmia purpurea Cresson, 1864. Ent. Soc. Phila., Proc. 3: 27. @.
Osmia rustica Cresson, 1864. Ent. Soc. Phila., Proc. 3: 27. d.
Osmia caerulescens Dalla Torre, 1896. Cat. Hym., v. 10, p. 388. Emend.

Biology: Chandler, 1963. North Central Branch, Ent. Soc. Amer., Proc. 18: 30 (nest,
predators, parasites). —Krombein, 1967. Trap-nesting wasps and bees, pp. 310-311 (nest
architecture, life history, predator). —Medler, 1967. Ent. Soe. Amer., Ann. 60: 341, table 1
(nest architecture). —Tasei, 1972. Apidologie 3: 149-165 (nest architecture, life history,
parasites). —Rust, 1974. Wasmann Jour. Biol. 32: 42 (life history, predators, parasites).

coloradensis Cresson. Nebr. to Tex., west to B. C., Wash., Oreg. and Calif.; Mex. (Baja
California). Ecology: Nests in burrow in Pinus jeffrey? cone as well as trap-nests.
Parasite: Chrysura pacifica (Say). Pollen: Apparently prefers pollen from the flowers of
Compositae, but also visits other flowers possibly for nesting material; visitation records
include Amsinckia intermedia, Barbarea orthoceras, Calandrinia caulescens var.
menziesii, Calendula, Carduus tenuiflorus, Centawrea, Chaenactis fremontii, C.
glabriuscula, Chrysothamnus, Cirsium californieum, Coreopsis lanceolata, Cryptantha
intermedia, Encelia farinosa, Erigeron, Eriodictyon, Eriogonum nudum, Eriophyllum

2034 Hymenoptera in America North of Mexico

confertiflorum, E. lanatum, Eucnide, Gilia multicaulis, Grindelia camporum,
Gutierrezia, Helenium bigelovii, Helianthella californica, Helianthus gracilentus,
Hyptis emoryji, Iris, Lasthenia chrysostoma, Layia platyglossa var. breviseta,
Malacothrix, Mimulus guttatus, Penstemon, Phacelia minor, Phalacoseris bolanderi,
Ranunculus californicus, Rhus trilobata, Senecio integerrimus, S. lugens, Silybum
marianum, Sisymbrium irio, Sonchus oleraceus, Sphaeralcea ambigua, Stenotopsis
linearifolius, Taraxacum officinale, Verbena, Viguiera laciniata, Wyethia.

Osmia coloradensis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 105. 2.

Osmia hypochrysea Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 449. 9.

Osmia hypochrysea Rohweri Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 366. 2.

Osmia pasadenae Cockerell, 1910. Ent. News 21: 122. 6.

Biology: Ruckes, 1956. Pan-Pacifie Ent. 32: 122, 1 fig. —Rust, 1974. Wasmann Jour. Biol. 32:
47-48 (nest architecture, cocoon, parasite).
georgica Cresson. Mass. to Mich., south to Fla. and Tex. Ecology: Nests in borings and glass

tubes. Parasite: Chrysura pacifica (Say). Pollen: Unknown, but visits flowers of Aronia,
Brassica, Cardamine, Chrysanthemum leucanthemum, Claytonia, Coreopsis, Erigeron,
Erythronium, Fragaria, Geranium, Hydrangea, Krigia Linaria, Lupinus, Malus,
Pyrus, Ranunculus, Rubus, Salix, Senecio, Taraxacum, Verbena, Vicia. Predator:
Camponotus sp., Crematogaster sp.

Osmia georgica Cresson, 1878. Amer. Ent. Soc., Trans. 7: 105. 2.

Osmia (Gnathosmia) louisianae Cockerell, 1901. Canad. Ent. 42: 171. 9.

Biology: Hartman, 1944. Psyche, 51: 162-165 (nest architecture, life history). —Krombein,
1967. Trap-nesting wasps and bees. p. 311 (nest architecture, life history). —Hawkins,
1975. Kans. Ent. Soc., Jour. 48: 493-499, 3 figs. (nest architecture, life history, parasite,
predator).

texana Cresson. N. Y., Mich., Ohio, Nebr., Tex., and B. C., Alta. and N. Dak., south to N. Mex.,

Ariz. and Calif.; Mexico (Baja California and Tamaulipas). Ecology: Nests in abandoned
cells of Anthophora occidentalis Cr. in clay bank and in elderberry trap-nests; uses
masticated leaf material from Malva neglecta and Sphaeralcea coccinea for cell
partitions and cell plug. Parasite: Anthrax irroratus Say, Chrysis densa Cress., Hornia
minutipennis Riley, Monodontomerus montivagus Ashm., Nemognatha sp., Stelis
montana Cress. Pollen: Apparently prefers pollen from Compositae, visits a wide
variety of flowers including Asclepias, Campanula, Carduus tenuiflorus, Calochortus
splendens, Centaurea, Chaenactis glabriuscula, Cirsium californicum, C. vulgare,
Coreopsis lanceolata, Cryptantha intermedia, Diplacus aurantiacus, Erigeron,
Eriodictyon californicum, Eriophyllum lanatum, Grindelia camporum, Frasera parryi,
Helianthus, Lotus, Marrubium vulgare, Monarda, Monardella villosa, Parkinsonia
aculeata, Penstemon, Phalacoseris bolanderi, Rubus leucodermis, Senecio, Stenotopsis
linearifolius, Sidalcea malvaeflora, Viguiera laciniata, V. multiflora, Wyethia
helenioides. Predator: Trichodes simulator (Horn).

Osmia texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 261. ¢.

Osmia mandibularis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 102. 2.

Osmia faceta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 103. 6.

Osmia davidsoniella Cockerell, 1905. Canad. Ent. 37: 370. ¢.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 217 (nest). —Mickel, 1928. Ent. News 39: 69
(nest). —Hobbs, Nummi and Virostek, 1961. Canad. Ent. 93: 143 (nest, parasite). —Rust,
1974. Wasmann Jour. Biol. 32: 54-56 (nest architecture, life history, parasites).

Genus OSMIA Subgenus CEPHALOSMIA Sladen

Cephalosmia Sladen, 1916. Canad. Ent. 48: 270.
Type-species: Osmia montana Cresson. Monotypic. (=Osmia armaticeps Cresson).

Revision: Rust, 1974. Wasmann Jour. Biol. 32: 56-85, figs. 5-6, 27-45, tables 7-9.

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 58 (summary of information on life histories).

Superfamily APOIDEA 2035

californica Cresson. B. C. to Calif., east to Mont., Wyo. and Colo. Ecology: Nests in borings

and preexisting burrows in various wood substrates. Parasite: Anthrax sp., Chrysis sp.,
Leucospis affinis Say, Nemognatha sp., Stelis sp., Tricrania stansburyi Hald. Pollen:
Apparently mainly dependent upon the pollens of Compositae, especially Cirsium, but
visits many flowers including Amsinckia, Arnica, Balsamorrhiza, Brodiaea lutea,
Calochortus, Carduus tenuiflorus, Ceanothus, Chaenactis glabriuscula, Cirsium
californicum, C. proteanum, Clarkia, Cryptantha intermedia, Erigeron pygmaeus,
Eriodictyon californicum, Eriogonum, Eriophyllum lanatum, Gilia capitata, Grindelia
camporum, Helianthus nuttallu, Layia platyglossa, Lotus strigosus var. hirtellus,
Lupinus, Mimulus guttatus, Penstemon, Phacelia heterophylla, P. humilis,
Physocarpus, Ranunculus, Salix, Senecio integerrimus, Sidalcea, Stenotopsis
linearifolius, Taraxacum officinale, Viguiera multiflora, Wyethia angustifolia, W. elata,
W. glabra, W. helenioides, W. longicaulis, W. mollis, W. ovata, W. reticulata. Predator:
Ptinis californicus Pic, Trichodes ornatus Say.

Osmia californica Cresson, 1864. Ent. Soc. Phila., Proc. 3: 24. 2.

Osmia pascoensis Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 342. 2.

Osmia nassa Cockerell, 1910. Ent. News 21: 272. @.

Osmia occidentalis Michener, 1936. Canad. Ent. 68: 42. 9.

Biology: Bohart, 1955. Ent. Soc. Wash., Proc. 57: 203-204 (nest, supersedure). —Levin, 1966.
Kans. Ent. Soc., Jour. 39: 524-535, 6 figs., 3 tables (nest architecture, life history,
supersedure, sex relationships, pollen collection, parasites). —Rust, 1974. Wasmann Jour.
Biol. 32: 66-69 (nest architecture, life history, parasites).

grinnelli Cockerell. Calif., Nev., Utah, Idaho. Pollen: Unknown, but visits flowers of
Arctostaphylos drupacea, A. patula, Aster abatus, Astragalus pachypus, Chaenactis
glabriuscula, Coreopsis lanceolata, Cryptantha intermedia, Descurainia sophia, Encelia
actoni, E. frutescens, Erysimum, Helianthus gracilentus, Hesperochiron californicum,
Hulsea callicarpha, Lupinus, Nepeta, Penstemon, Prunus subcordata, Sisymbrium,
Stenotopsis linearifolius, Taraxacum, Wyethia.
Osmia grinnelli Cockerell,-1910. Ent. News 21: 120. 2.
marginipennis Cresson. Calif., Nev., Oreg., Idaho, Utah, Colo., Wyo.; Mexico (Baja California).
Pollen: Unknown, but visits flowers of Chaenactis.
Osmia marginipennis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 106. d.
Osmia gaillardiae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 227. °.
Osmia viridior Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 261. 6.
Osmia leonis Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 128. °.
montana montana Cresson. B. C. to Calif. (White Mts.), east to Idaho and Mont., south to
Ariz. and N. Mex. Ecology: Nests in trap-nests. Parasite: Stelis montanu Cress. Pollen:
Analyzed pollen stores indicates reliance on Compositae including Balsamorrhiza
sagittata, Cosmos, Helianthus annuus, Wyethia amplexicaulis, but also visits other
flowers for nectar or pollen including Arnica, Aster, Chaenactis, Gutierrezia, Senecio,
Silybum, Taraxacum.
Osmia montana Cresson, 1864. Ent. Soe. Phila., Proc. 3: 24. d.
Osmia armaticeps Cresson, 1878. Amer. Ent. Soc., Trans. 7: 104. 2.
Osmia armaticeps var. sapellonis Cockerell, 1901. Ann. and Mag. Nat. Hist. (7) 7: 336. 2.

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 76-80 (nest architecture, life history,
supersedure, parasite).

montana quadriceps Cresson. Wash. to Calif., Nev., Ariz. (Santa Catalina Mts.). Ecology:
Nests in abandoned beetle burrows in Pinus contorta. Pollen: Apparently polylectic,
although most floral records from noncomposite genera refer to visits for leaf or dried
flower material; visitation records include Agoseris heterophylla, Arctostaphylos patula,
Aster, Astragalus, Balsamorhiza deltoides, Ceanothus integirrimus, Chaenactis
glabriuscula, Cirsium, Clarkia biloba, Coreopsis, Eriodictyon, Eriophyllum
confertiflorum, E. lanatum, Gilia capitata, Gormania obtusata, Haplopappus
linearifolius, Helianthella californica, Helianthus gracilentus, Hesperochiron
californicus, Lasthenia chrysostoma, Layia platyglossa, Lotus scoparius, Lupinus,
Mimulus suksdorfii, Nemophila integrifolia, Penstemon, Phacelia humilis, Potentilla,

2036 Hymenoptera in America North of Mexico

Raillardella scaposa, Ribes cereum, Salvia, Senecio, Sidalcea, Sisymbrium irio,
Tamarix gallica, Taraxacum officinale, Wyethia angustifolia, W. helenioides, W. mollis.

Osmia quadriceps Cresson, 1878. Amer. Ent. Soc., Trans. 7: 104. 2.

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 81 (nest).

subaustralis Cockerell. N. W. T. east to Ont. and Mich., south to Calif., Nev., Ariz. and N. Mex.

Ecology: Nests in abandoned beetle burrows in fir log, probably Abies lasiocarpa.
Pollen: Analyzed pollen stores revealed 99 per cent from Compositae, probably Senecio;
visits flowers of Aster, Centaurea, Encelia farinosa, Erigeron, Gutierrezia, Lupinus,
Penstemon, Phacelia humilis, Raillardella scaposa, Senecio, Solidago, Streptanthus,
Taraxacum, Wyethia, Yucca.

Osmia nigrifrons var. subaustralis Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 410. 2.

Osmia wardiana Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 368. 9.

Osmia seneciophila Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 446. 3.

Osmia nelsoni Cockerell, 1930. Ann. and Mag. Nat. Hist. (10) 5: 411. °.

Osmia wardiana austromontana Michener, 1936. Canad. Ent. 68: 42. 2.

Osmia lyncis Cockerell, 1937. South. Calif. Acad. Sci., Bul. 36: 110. 9.

Osmia (Cephalosmia) mendocinensis Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 27. 9.

Biology: Rust, 1974. Wasmann Jour. Biol. 32: 85 (nest architecture, pollen).
Genus OSMIA Subgenus CENTROSMIA Robertson

Centrosmia Robertson, 1903. Amer. Ent. Soc., Trans. 29: 165.
Type-species: Osmia bucephala Cresson. Orig. desig. and monotypic.

Revision: Sinha and Michener, 1958. Kans. Univ. Sci. Bul. 39: 275-303 (Nearctic spp.).

austromaritima Michener. Calif., Colo., Utah.
Osmia austromaritima Michener, 1936. Canad. Ent. 68: 43. 6.
bakeri Sandhouse. Oreg., Calif. Pollen: Unknown, but visits flowers of Clarkia, Collinsia
childii, C. heterophylla, C. wrightii, Cryptantha lepida, Gilia tricolor, Nemophila
integrifolia.
Osmia baker Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 345. ¢.
bucephala Cresson. Holarctic; Alaska, Yukon and N. W. T. to Que., south to Calif., Utah, N.
Mex., Ill, Tenn., and N. C.; limited to mountains in southern part of range. Ecology:
Nests in borings and in a tunnel in a maple tree. Parasite: Chaetodactylus krombeini
Baker, Melittobia chalybii Ashm. Pollen: Unknown, but visits flowers of Aesculus,
Azalea, Baptisia, Cercis canadensis, Dentaria, Dicentra, Erythronium, Mertensia
virginica, Nepeta hederacea, Penstemon, Taraxacum, Tephrosia virginiana, Viola.
Osmia bucephala Cresson, 1864. Ent. Soe. Phila., Proc. 3: 17. 9°.
Osmia megacephala Cresson, 1864. Ent. Soc. Phila., Proc. 3: 18. °.
Osmia latitarsis Cresson, 1864. Ent. Soc. Phila., Proc. 3: 20. ¢.
Osmia lignivora Packard, 1867. Amer. Nat. 1: 375. @.
Osmia lignicola Provancher, 1882. Nat. Canad. 13: 208. °.
Osmia subornata Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 342. 2.

Biology: Packard, 1867. Amer. nat. 1: 375 (nest, as lignivora). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 307-310, pl. 18, figs. 89-91 (nest architecture, life history,
supersedure, parasites).

nigriventris (Zetterstedt). Holarctic; Alaska to Hudson Bay and Ont., south to Oreg., Colo. and
Minn.; Eurasia.
Anthophora nigriventris Zetterstedt, 1838. Ins. Lapponica, v. 1, p. 465. 9.
Osmia frigida Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 142. 2, ¢.
Osmia hudsonica Cresson, 1864. Ent. Soc. Phila., Proc. 3: 21. d.
Osmia corticalis Gerstaecker, 1869. Stettin. Ent. Ztg. 30: 331. 2, d.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 331. 2, d. —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 152: 77-78, fig. 33, table 4 (redescription, synonymy).

Superfamily APOIDEA 2037

pikei Cockerell. B. C., Wash., Oreg., Nev., Calif., Colo., Wyo. Parasite: Nemognatha scutellaris
LeC., Sapyga fulvicornis Cress. Pollen: Unknown, but visits flowers of Ribes cereum.
Osmia pikei Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 367. 9.
Osmia universitatis Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 538. 6.
Osmia integrella Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 124. 6.
Osmia vallicola Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 449. ¢.
Osmia amala Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 447. 6.
Osmia metitia Cockerell, 1909. Canad. Ent. 41: 130. ¢.
raritatis Michener. Colo., Wash., Calif.
Osmia raritatis Michener, 1957. Kans. Ent. Soc., Jour. 30: 40, figs. 1, 3, 5. ¢.
tanneri Sandhouse. Utah. Ecology: Nests made entirely of mud on concave underside of stone.
Parasite: Chrysura pacifica (Say). Pollen: Traces of pollen attached to cocoon and exines
in feces in cell indicate an unknown species of Umbelliferae.
Osmia (Nothosmia) tanneri Sandhouse, 1939. Ent. Soe. Wash., Mem. 1: 87. ¢.

Taxonomy: Parker, 1975. Pan-Pacific Ent. 51: 179-180, figs. 1-2. 9°.

Biology: Parker, 1975. Pan-Pacifie Ent. 51: 180-182, figs. 3-4 (nest architecture, larval habits,
life history, parasite).
tarsata Provancher. Que.
Osmia tarsata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 328. d.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 78-79 (redescription, tax.
status).
thysanisca Michener. Calif., Oreg., Wyo. Pollen: Unknown, but visits flowers of Hackelia
jessicae, Trifolium.
Osmia thysanisca Michener, 1957. Kans. Ent. Soc., Jour. 30: 39, fig. 2. ¢.

vandykei Sandhouse. Oreg.
Osmia vandykei Sandhouse, 1924. Calif. Acad. Sci., Proe. (4) 18: 344. 6.

Genus OSMIA Subgenus ACANTHOSMIOIDES Ashmead

Acanthosmioides Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 76.
Type-species: Osmia odontogaster Cockerell. Orig. desig. and monotypic.

Revision: White, 1952. Kans. Univ. Sci. Bul. 35: 219-307, pls. 39-44 (Nearctie spp.).

Taxonomy: Michener, 1936. Amer. Mus. Novitates 875: 27-28 (Key).
ashmeadii (Titus). Oreg. (Dalles).
Acanthosmioides ashmeadii Titus, 1904. Ent. Soe. Wash., Proc. 6: 101. d.
calearata White. Calif. (Sierra Nevada Mts.). Pollen: Unknown, but visits flowers of
Astragalus, Lotus.
Osmia calcarata White, 1952. Kans. Univ. Sci. Bul. 35: 245, pl. 39, fig. 15; pl. 40, fig. 11; pl.
41, fig. 9; pl. 42, fig. 9; pl. 43, fig. 11; pl. 44, fig. 15. d, °.
dakotensis Michener. N. Dak., Mont., east. Colo., Wyo.
Osmia dakotensis Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 407. °.
Osmia (?Acanthosmioides) dacotensis Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 62.
Emend.
francisconis White. Calif. (San Francisco). Ecology: Collected in sand dunes.
Osmia francisconis White, 1952. Kans. Univ. Sci. Bul. 35: 269, pl. 40, fig. 2. °.
giffardi Sandhouse. Calif. (Sierra Nevada Mts.).
Osmia (Acanthosmioides) giffardi Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 50. d.
giliarum Cockerell. Idaho, Mont., Wyo., Colo. Pollen: Unknown, but visits flowers of Gilia,
Penstemon.
Osmia giliarum Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 451. 2.
hurdi White. Calif., B. C. Pollen: Unknown, but visits flowers of Allium.
Osmia hurdi White, 1952. Kans. Univ. Sci. Bul. 35: 278, pl. 39, fig. 7. 2.

2038 Hymenoptera in America North of Mexico

integra Cresson. Calif. to B. C., east to Man., south to Ariz., N. Mex., Tex., Mich. Pollen:
Unknown, but visits flowers of Astragalus pomonensis, Lotus scoparius, Phacelia
distans.
Osmia integra Cresson, 1878. Amer. Ent. Soc., Trans. 7: 106. 6.
Osmia novomexicana Cockerell, 1903. Ent. News 14: 331. @.
Osmia florissanticola Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 450. 9.
Osmia integra nigrigena Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 90. d, °.

Taxonomy: Cockerell, 1910. Ent. News 21: 270. 6.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 246 (nesting habits, as novomexicana).

kenoyeri Cockerell. Yukon, Alta., Calif., Colo.
Osmia kenoyeri Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 16: 483. d, 2.
lanei Sandhouse. Calif.
Osmia (Acanthosmioides) lanei Sandhouse, 1939. Ent. Soe. Wash., Mem. 1: 51. do.
longula Cresson. B. C. to Calif., east to Mont., Wyo., Colo. and Utah. Ecology: Nests on side of
stone beneath a slight overhang. Pollen: Stores pollen of an unidentified Leguminosae;
visits flowers of Astragalus, Erigeron compositus, Penstemon heterodoxus, Wyethia
mollis.
Osmia longula Cresson, 1864. Ent. Soc. Phila., Proc. 3: 19. 2.
Osmia abnormis Cresson, 1878. Amer. Ent. Soce., Trans. 7: 105. 6.
Osmia grandior Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 343. °.
Osmia permorata Cockerell, 1910. Canad. Ent. 42: 310. 9.

Biology: Parker, 1975. Pan-Pacific Ent. 51: 182-183, figs. 5-6 (nest architecture, larval habits,
sex ratio, nest provisions).
nifoata Cockerell. Wash. to Calif., east to Idaho, Utah, Colo. and Wyo. Parasite: Dioxys

pomonae Ckll. Pollen: Unknown, but visits flowers of Amsinckia, Astragalus
pomonensis, Chaenactis, Lotus argophyllus, L. crassifolius, L. davidsonii, L. glaber, L.
scoparius, L. strigosus, Lupinus confertus, L. formosus, L. lobbii var. lyalli, L.
odoratus, Potentilla glandulosa, Rhus trilobata, Streptanthus bernardinus, Trifolium
involucratum, T. variegatum.

Osmia (Acanthosmioides) nifoata Cockerell, 1909. Entomologist 42: 95. d.

Osmia pellax Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 356. °.

Osmia (Acanthosmioides) wyomingensis Michener, 1936. Amer. Mus. Novitates 875: 27. ¢.

Biology: Fulton and Bergen, 1935. Econ. Ent., Jour. 28: 729 (nest). —Hurd, 1958. Calif. Univ.
Pubs. Ent. 14: 285 (parasite).
nigrifrons Cresson. N. Dak. and Alta., west to B. C., south to Calif., Utah, Colo. and Wyo.

Ecology: Nest in borings, bamboo culms, and in corrugated steel drain pipes. Parasite:
Chrysura pacifica (Say), Dibrachys sp., Monodontomerus montivagus Ashm., Stelis
chlorocyanea Ckll. Pollen: Polylectic, apparently prefers flowers of Leguminosae for
pollen and nectar; visits other flowers for pollen including Astragalus, Lotus,
Onobrychis, Trifolium, Vicia; visitation records include Amsinckia douglasiana,
Astragalus douglasiana, A. parishii, A. utahensis, Brassica nigra, Lotus crassifolius, L.
davidsonii, L. humistratus, L. scoparius, Medicago sativa, Melilotus alba, M. officinalis,
Onobrychis sativa, Phacelia, Trifolium repens, Vicia americana.

Osmia nigrifrons Cresson, 1878. Amer. Ent. Soc., Trans. 7: 108. 9.

Osmia casta Cockerell, 1910. Ent. News 21: 272. 2.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 246 (nest). —Rust and Thorp, 1973. Kans. Ent.
Soe., Jour. 46: 548-562, 26 figs., 2 tables (parasite). —Rust, Thorp and Torchio, 1974. Nat.
Hist., Jour. 8: 29-47, 14 figs., 2 tables (nest architecture, life history, parasites).

nigrobarbata Cockerell. Wash., Calif., Ariz., Oreg.?, Idaho?, Utah? and Sask.? Ecology: Nests
in burrows in the ground. Parasite: Diorys pomonae Ckll. Pollen: Stores pollen of
Astragalus including A. antisellii, A. parishii, A. pomonensis, but visits other flowers
including Amsinckia, Brodiaea, Castilleia, Cryptantha, Lotus, Lupinus, Mimulus,
Penstemon, Salvia, Vicia.
Osmia nigrobarbata Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 52. °.
Osmia vanduzeei Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 348. ¢.

Superfamily APOIDEA 2039

Biology: Rozen and Favreau, 1967. N. Y. Ent. Soe., Jour. 75: 197-203, 31 figs. (nest
architecture, life history, parasite).
obliqua White. Calif., Oreg. Pollen: Unknown, but visits flowers of Astragalus.
Osmia obliqua White, 1952. Kans. Univ. Sci. Bul. 35: 238, pl. 40, fig. 7; pl. 41, fig. 5; pl. 42,
fig. 2; pl. 43, fig. 9; pl. 44, figs. 1, 5. d.
odontogaster Cockerell. B. C., Wash., Oreg., Calif.
Osmia odontogaster Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 338. ¢.
physariae Cockerell. B. C., Alta. and Sask., south to Calif., Nev., Utah, Colo. and Wyo. Pollen:
Unknown, but visits flowers of Allium, Iris missouriensis, Lappula, Lotus, Trifolium
hybridum.
Osmia physariae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 446. 3.
Osmia pinorum Cockerell, 1935. Amer. Mus. Novitates 766: 4. 2.
Osmia (Acanthosmioides) erecta Michener, 1936. Canad. Ent. 68: 40. 2.
Osmia crenulaticornis Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 84. ¢.

Taxonomy: Michener, 1949. Kans. Ent. Soe., Jour. 22: 56 (crenulaticornis).
sedula Sandhouse. B. C., Wash., Wyo., Calif. Pollen: Unknown, but visits flowers of Layia
platyglossa, Lotus, Phacelia distans.
Osmia sedula Sandhouse, 1924. Calif. Acad. Sei., Proc. (4) 13: 353. 9.
Osmia lutzi Michener, 1936. Amer. Mus. Novitates 875: 25. 9.
sladeni Sandhouse. B. C., Alta., Wyo., Utah.
Osmia sladeni Sandhouse, 1925. Canad. Ent. 57: 33. 6.
trifoliama Sandhouse. Oreg. Pollen: Unknown, but visits flowers of Trifolium hybridum, T.
pratense.
Osmia (Acanthosmioides) trifoliama Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 44. 9.
unca Michener. Oreg., Wash., B. C., Idaho, Mont., Colo., Utah. Ecology: Cells are constructed
with a leaf mastic lining surrounded by mud.
Osmia (Acanthosmioides) uncinata Michener, 1936. Canad. Ent. 68: 39. 3. Preoce.
Osmia (Acanthosmioides) unca Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 406. N.
name.
Osmia (Acanthosmioides) hicksi Sandhouse, 1939. Ent. Soc. Amer., Mem. 1: 53. 3.
Osmia (Acanthosmioides) depressa Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 406.
3.

Biology: Stephen, Bohart and Torchio, 1969. The biology and external morphology of bees,
pp. 76, 100, 104 (nest, cell construction).

watsoni Cockerell. N. Mex. (Albuquerque).
Osmia watsoni Cockerell, 1911. Amer. Ent. Soe., Trans. 37: 235. 6.

Genus OSMIA Subgenus NOTHOSMIA Ashmead

Nothosmia Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 75.
Type-species: Osmia distincta Cresson. Orig. desig. and monotypic.
Leucosmia Robertson, 1903. Amer. Ent. Soc., Trans. 29: 166.
Type-species: Osmia albiventris Cresson. Orig. desig. and monotypic.
Xanthosmia Robertson, 1903. Amer. Ent. Soc., Trans. 29: 166.
Type-species: Osmia cordata Robertson. Orig. desig. and monotypic.
albiventris Cresson. Que. to Ga., west to Ill. and Minn. Pollen: Unknown, but visits flowers of
Brassica, Geranium, Gillenia, Malus, Penstemon, Rubus, Trifolium, Vicia.
Osmia albiventris Cresson, 1864. Ent. Soc. Phila., Proc. 3: 31. 2, 6.

Biology: Medler, 1967. Ent. Soc. Amer., Ann. 60: 341-342, fig. 1, table 1 (nest architecture,
supersedure, life history).
cordata Robertson. Colo., N. Mex., N. Dak., IIl., Mo., Ohio, Ind. Ecology: Nests in abandoned
mud-dauber cells, and in borings in wood. Parasite: Monodontomerus mandibularis
Gahan, M. montivagus Ashm., M. obscurus Westw., Sapyga conflwenta Cress. Pollen:
Unknown, but visits flowers of Aesculus, Cardamine, Lithospermum, Penstemon,
Phlox, Rubus, Salvia, Trifolium, Verbena.
Osmia cordata Robertson, 1902. Ent. News 13: 79. d.

2040 Hymenoptera in America North of Mexico

Osmia hesperella Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 227. 9.
Osmia coloradella Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 229. °.
Osmia Ramaleyi Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 126. 6, °.
Osmia figginsi Cockerell, 1935. Amer. Mus. Novitates 766: 2. 3.

Taxonomy: Cockerell, 1911. Canad. Ent. 43: 389 (tax. characters, synonymy).

Biology: Turner, 1911. Jour. Anim. Behavior 1: 374 (as “Stelidae”). —Rau, 1916. Jour. Anim.
Behavior 6: 39. —Rau, 1923. Ent. News 34: 308. —Rau, 1928. Acad. Sci. St. Louis, Trans.
25: 363. —Hicks, 1934. Colo. Univ., Studies 21: 265. —Rau, 1937. Ent. Soc. Amer., Ann. 30:
324 (nest architecture, life history, parasite). —Chandler, 1959 (1958). Indiana Acad. Sci.,
Proc. 68: 199-204 (life history). —Chandler, 1962. Ent. Soc. Amer., North Central Branch,
Proc. 16: 18-19 (interspecific competition). —Chandler, 1963. Ent. Soc. Amer., North
Central Branch, Proc. 18: 30 (interspecific competition).
distincta Cresson. Ont. and Que., Maine to N. C., west to N. Dak., and Colo. Pollen: Unknown,
but visits flowers of Blephilia, Dianthera, Gillenia, Hypoxis, Lithospermum,
Penstemon, Rubus, Trifoliwm.
Osmia distincta Cresson, 1864. Ent. Soc. Phila., Proc. 3: 30. 2.
enixa Sandhouse. Colo., Calif.
Osmia enixa Sandhouse, 1924. Calif. Acad. Sci., Proe. (4) 13: 347. 3.
grindeliae Cockerell. Alta., Colo., Wash., Oreg., Calif. Pollen: Unknown, but visits flowers of
Grindelia.
Osmia grindeliae Cockerell, 1910. Psyche 17: 246. °.
inspergens Lovell and Cockerell. Que. and Maine, south to Ga. Pollen: Unknown, but visits
flowers of Baptisia, Polycodium, Trifolium, Vaccinium.
Osmia inspergens Lovell and Cockerell, 1907. Psyche 14: 17. °.
liogastra Cockerell. Calif. Pollen: Unknown, but visits flowers of Lotus, Lwpinus, Phacelia
distans.
Osmia liogastra Cockerell, 1933. Pan-Pacifie Ent. 9: 26. 2.
lupinicola Cockerell. Calif. Pollen: Unknown, but visits flowers of Astragalus douglasii,
Chaetopappa aurea, Cryptantha, Lotus davidsonii, Lupinus, Nolina.
Osmia lupinicola Cockerell, 1937. Amer. Mus. Novitates 948: 12. 9.
marginata Michener. South. Calif., desert. Parasite: Nemognatha macswaini Enns. Pollen:
Polylectic, visits flowers of Amsinckia douglasiana, Antirrhinum filipes, Astragalus
coulteri, Beleperone californica, Chaenactis glabriuscula, Dalea mollis, D. saundersii,
Helianthus annuus, H. gracilentus, Hyptis emoryi, Isomeris arborea, Larrea tridentata,
Lotus scoparius, Lupinus sparsifolius, Mirabilis glutinosa, Prosopis, Salvia carduacea.
Osmia marginata Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 86. 9.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 37 (floral relationships).

melanopleura Cockerell. Calif.
Osmia melanopleura Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 52. °.
michiganensis Mitchell. Mich. (Grand Traverse Co.).
Osmia (Nothosmia) michiganensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 92.
Gir
mixta Michener. Calif.
Osmia mixta Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 88. 9.

Taxonomy: Michener, 1949. Kans. Ent. Soc., Jour. 22: 56.
morongana Cockerell. Calif. Pollen: Unknown, but visits flowers of Amsinckia, Lupinus
odoratus, Orthocarpus ornatus, Hyptis emoryi.

Osmia morongana Cockerell, 1937. Amer. Mus. Novitates 948: 11. °.
nigritula Friese. No locality known.

Osmia parva Provancher, 1883. Nat. Canad. 14: 37. ¢. Preoce.

Osmia parvula Dalla Torre, 1896. Cat. Hym., v. 10, p. 405. N. name. Preoce.

Osmia nigritula Friese, 1902. Ztschr. System. Hym. Dipt. 2: 109. N. name.
novaescotiae Cockerell. N. S. Probably a synonym of simillima or proxima.

Osmia novaescotiae Cockerell, 1912. Canad. Ent. 44: 355. 2.

Superfamily APOIDEA 2041

phenax Cockerell. N. Mex.
Osmia phenax Cockerell, 1897. Canad. Ent. 29: 66. 2.
prunorum Cockerell. N. Mex.
Osmia prunorum Cockerell, 1897. Canad. Ent. 29: 65. 2 (¢ misdet.).
pumila Cresson. Que. to Ga., west to IIl., Minn., Kans. Ecology: Readily nests in borings.
Parasite: Chaetodactylus? sp., Chrysura pacifica (Say), Leucospis affinis Say,
Melittobia chalybui Ashm., Pyemotes ventricosus (Newport), Sapypa centrata Say.
Pollen: Apparently polylectic, visits a wide variety of flowers including Anemonella,
Antennaria, Arabis, Astragalus, Barbarea, Camassia, Cardamine, Cercis,
Chaerophyllum, Claytonia, Collinsia, Comandra, Cornus, Crataegus, Dentaria,
Diospyros, Ellisia, Erigenia, Erythronium, Fragaria, Geranium, Houstonia, Isopyrum,
Krigia, Malus, Malva, Melilotus, Mertensia, Nepeta, Oxalis, Pedicularis, Penstemon,
Polemonium, Potentilla, Ranunculus, Ribes, Rosa, Rubus, Salix, Sassafras, Scutellaria,
Sisyrinchium, Smilax, Taraxacum, Trifolium, Vaccinium, Viburnum, Vicia, Viola,
Zanthoxylum, Zizia.
Osmia pumila Cresson, 1864. Ent. Soc. Phila., Proc. 3: 35. 2.
Osmia vicina Cresson, 1864. Ent. Soe. Phila., Proce. 3: @8. dc.
Osmia pumilia(!) Pearson, 1933. Ecol. Monogr. 3: 381.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 312-318, tables 32-33 (nest
architecture, life history, supersedure, parasites). —Medler, 1967. Ent. Soc. Amer., Ann.
60: 342, table 1 (nest architecture, life history, parasite).
sandhouseae Mitchell. New England to Fla., west to Tex. Pollen: Unknown, but visits flowers

of Astragalus, Claytonia, Crataegus, Fragaria, Gaylussacia, Geranium, Ilex, Iris,
Linaria, Lupinus, Oenothera, Pedicularis, Penstemon, Polycodium, Rubus, Tephrosia,
Toxicodendron, Trifolium, Vaccinium, Vicia.

Osmia albohirta Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 164. 2. Preoce.

Osmia sandhouseae Mitchell, 1927. Psyche 34: 178. N. name.

solitaria Sandhouse. Calif.
Osmia solitaria Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 346. 3.

Taxonomy: Michener, 1949. Kans. Ent. Soc., Jour. 22: 55.
titusi Cockerell. South. Calif., N. Mex.? Pollen: Unknown, but visits flowers of Astragalus
fremontii, Lotus, Larrea tridentata, Lupinus sparsifolius, Phacelia fremontii.
Osmia titusi Cockerell, 1905. Canad. Ent. 37: 370. 9.

Genus OSMIA Subgenus CHENOSMIA Sinha

Osmia Subg. Chenosmia Sinha, 1958. Kans. Univ. Sci. Bul. 39: 233.
Type-species: Osmia pentstemonis Cockerell. Orig. desig.
atriventris Cresson. N. S. to Alta., south to Ia., Ill, Tenn. and Ga. Ecology: Readily nests in
artificial nesting devices. Parasite: Chrysis coerulans Fabr., Dibrachys maculipennis
Szelenyi?, Leucospis affinis Say, Sapyga sp., Stelis foederalis Sm. Pollen: Unknown, but
visits flowers of Arabis, Astragalus, Barbarea, Cardamine, Cercis canadensis,
Chrysanthemum leucanthemum, Claytonia, Collinsia, Ellisia, Erigenia, Fragaria,
Geranium, Gillenia, Hydrangea, Hypoxis, Lithospermum, Mertensia, Nothoscordum,
Penstemon, Polemonium, Psoralea, Rubus, Trifolium, Uvularia, Vaccinium, Vicia,
Viola, Zizia. J
Osmia atriventris Cresson, 1864. Ent. Soc. Phila., Proc. 3: 29. 2.

Biology: Fye, 1965. Canad. Ent. 97: 872-874, fig. 4, tables 3-4 (nest architecture, life history,
parasites).

bruneri Cockerell. Mont. to N. Mex., west to B. C. and Calif. Pollen: Unknown, but visits
flowers of Chamaebatia foliolosa, Clarkia rhomboidea, Collinsia heterophylla,
Eriodictyon californicum, Erysimum perenne, Geranium molle, Gilia capitata,
Hesperochiron, Lepechinia calycina, Lotus argophyllus, L. scoparius, Limnan thes
douglasii, Lupinus lobbii var. lyallii, Malacothamnus arcuatus, Medicago sativa,
Mimulus guttatus, Monardella lanceolata, Nemophila spatulata, Penstemon
heterophyllus, P. spectabilis, Phacelia distans, P. frigida, P. heterophylla, P. imbricata,

2042 Hymenoptera in America North of Mexico

Sidalcea malvaeflora, Solidago multiradiata, Stellaria crispa, Streptanthus tortuosus,
Taraxacum, Trifolium microcephalum, T. repens, Verbena californica, V. lasiostachys,
Vicia americana, V. californica, V. cracca.

Osmia bruneri Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 337. 9.

Osmia Bennettae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 122. d.

Osmia holochlora Cockerell, 1923. Canad. Ent. 55: 205. 6.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 242, 243 (nesting habits).
calla Cockerell. Idaho to Colo., west to Calif. and B. C. Pollen: Unknown, but visits flowers of
Astragalus, Brassica geniculata, Brodiaea lutea, Cirsium californicum, Clarkia
rhomboidea, Cryptantha intermedia, C. lepida, Eriodictyon trichocalyx, Lepechenia
calycina, Lotus corniculatus, Lotus davidsonii, L. scoparius, Lupinus albifrons,
Malacothamnus arcuatus, Marrubium vulgare, Nama parryi, Nemophila integrifolia,
Penstemon grinnellii, P. labrosus, Phacelia distans, P. heterophylla, P. imbricata, P.
ramosissima, Phalacoseris bolanderi, Potentilla glandulosa, Streptanthus bernardinus,
Trichostema lanatum, Trifolium involucratum, T. microcephalum, T. repens, T.
variegatum, Verbena lasiostachys.
Osmia calla Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 337. d.
Osmia Ednae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 123. “2” = 6.

caulicola Cockerell. Wyo., Colo.
Osmia caulicola Cockerell, 1934. Brooklyn Ent. Soc., Bul. 29: 17. 2, 6.

cerasi Cockerell. N. Mex.
Osmia cerasi Cockerell, 1897. Canad. Ent. 29: 66. °.

clarescens Cockerell. South. Calif., N. Mex., Sask.? Ecology: Nest in abandoned cells of
Sceliphron. Pollen: Polylectic, visits flowers of Amsinckia douglasii, Arctostaphylos
patula, Astragalus parishii, A. pomonensis, Beleperone californica, Chaenactis
glabriuscula, Cirsium californicum, Cryptantha intermedia, Dalea saundersii,
Eriodictyon crassifolium, Helianthus gracilentus, Heliotropium curassavicum, Hyptis
emoryi, Isomeris arborea, Larrea tridentata, Koeberlinia, Lotus davidsonii, L. glaber, L.
scoparius, Marrubium vulgare, Penstemon antirrhinoides, P. spectabilis, Phacelia
davidsonii, P. distans, P. fremontii, Rhus trilobata, Ribes indecorum, Salvia carnosa, S.
columbariae, S. mellifera, Sphaeralcea ambigua, Trifolium repens, T. variegatum.

Osmia clarescens Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 764. 2, d.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 36-37 (floral relationships).

cobaltina Cresson. Idaho to Colo., west to B. C. and Calif., Ariz.? Pollen: Unknown, but visits a
wide variety of flowers including Ambrosia, Arctostaphylos drupacea, A. patula,
Chamaebatia foliolosa, Clarkia pulchella, C. rhomboidea, Eriogonum inflatum,
Horkelia bolanderi, H. tillingi, Lotus argophyllus, L. davidsonii, L. glaber, Lupinus
austromontanus, Mimulus primuloides, Monardella stricta, M. linoides, M. stricta,
Orthocarpus purpurascens, Penstemon bridgesii, P. grinnellii, P. labrosus, P. palmeri,
Phacelia heterophylla, P. imbricata, P. ramosissima, Phalacoseris bolanderi, Potentilla
glandulosa, Ribes, Sambucus nevadensis, Senecio lugens, Streptanthus bernardinus, S.
major, Vicia americana.

Osmia cobaltina Cresson, 1878. Amer. Ent. Soc., Trans. 7: 104. 2.

Osmia bella Cresson, 1878. Amer. Ent. Soc., Trans. 7: 107. 3.

Osmia basilissa Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 764. °.
Osmia kerminesina Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 349. 9.

cockerelli Sandhouse. Colo., N. Mex.

Osmia (Nothosmia) cockerelli Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 104. d.
collinsiae Robertson. Maine to N. C., west to Minn. and III. Pollen: Unknown, but visits
flowers of Astragalus, Cardamine, Cercis, Claytonia, Collinsia, Dicentra, Erythronium,
Geranium, Lupinus, Nepeta, Pedicularis, Penstemon, Rubus, Trifolium, Vicia, Viola.
Osmia collinsiae Robertson, 1905. Canad. Ent. 37: 236. d.

cyanopoda Cockerell. Idaho, Nev., Wash., Oreg., Calif. Pollen: Unknown, but visits flowers of
Amsinckia intermedia, Castilleia, Cryptantha intermedia, Erysimum asperum, Lotus
scoparius, Salvia columbariae, S. mellifera.
Osmia cyanopoda Cockerell, 1916. Pomona Col. Jour. Zool. 8: 52. 2.

Superfamily APOIDEA 2043

Osmia parallela Michener, 1936. Amer. Mus. Novitates 875: 19. ¢.

dolerosa Sandhouse. B. C., Wash., Oreg., Calif. Pollen: Unknown, but visits flowers of Clarkia
rhomboidea, Collinsia tinctoria, Horkelia fusca, Limnanthes douglasti, Lotus
argophyllus, Rosa, Stachys, Trifolium microcephalum, T. gracilentwm.
Osmia dolerosa Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 128. 2.
exigua Cresson. South. Colo., N. Mex., Calif. Pollen: Unknown, but visits flowers of Allinm
parvum, Anagallis arvensis, Brodiaea elegans, B. laxa, B. lutea, Centaurea melitensis,
Cirsium vulgare, Clarkia unguiculata, Diplacus aurantiacus, Epilobium minutum,
Eriodictyon californicum, Eriophyllum confertiflorum, Geranium molle, Helianthella
californica, Hesperochiron, Lepechinia calycina, Lotus scoparius, L. subpinnatus,
Lupinus densiflorus, Mimulus guttatus, Monardella villosa, Navarretia heterodoxa,
Penstemon speciosus, Phacelia cicutaria, P. distans, Plectritis macrocera, Satureja
douglasii, Streptanthus tortuosus, Trifolium microcephalum, T. variegatum, Verbena
lasiostachys, Viola purpurea.
Osmia exigua Cresson, 1878. Amer. Ent. Soc., Trans. 7: 107. d.
Osmia granulosa Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 767. 2, d.
Osmia exiguua(!) Cresson, 1916. Amer. Ent. Soc., Mem. 1: 118.
Osmia vana Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 349. °.

Taxonomy: Michener, 1962. Kans. Ent. Soc., Jour. 35: 253-254 (tax. characters, geogr.
records).
gaudiosa Cockerell. N. Mex., Colo., Ariz., Calif. Pollen: Analyzed pollen masses from two cells

revealed 96 per cent Papillionoideae (cf. Astragalus, Vicia, or Lathyrus) and 4 per cent
probably Scrophulariaceae (cf. Stemodia); visitation records include Arctostaphylos
patula, Astragalus, Dalea californica, Fragaria californica, Gilia multicaulis, Lathyrus,
Lotus davidsonii, L. scoparius, Mirabilis laevis, Nemophila integrifolia, Phacelia
davidsonii, P. distans, Rhus trilobata, Ribes indecorum, Salvia columbariae, Sambucus
caerulea. Predator: Metaponium sp., Telabis sp.

Osmia gaudiosa Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 588. d.

Taxonomy: Michener, 1962. Kans. Ent. Soc., Jour. 35: 253-254 (tax. characters, geogr.
records).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 318-320 (nest architecture, life
history, larval food, supersedure, predators).
hemera Sandhouse. Calif.
Osmia (Nothosmia) hemera Sandhouse, 1939. Ent. Soe. Wash., Mem. 1: 114. d.
illinoensis Robertson. South. IIl., Tex. Pollen: Unknown, but visits flowers of Astragalus,
Fragaria, Hypoxis, Lithospermum, Ranunculus.
Osmia illinoensis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 347. ¢.
indeprensa Sandhouse. Idaho, Nev., Wash., Oreg., Calif. Pollen: Unknown, but visits flowers of
Calyptridium wmbellatum, Clarkia pulchella, Collinsia torreyi, Limnanthes douglasii,
Lotus nevadensis, Phacelia hydrophylloides.
Osmia (Nothosmia) indeprensa Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 115. 2, d.
inermis (Zetterstedt). Holarctic; Labrador, Que., N. B., Ont., Maine, Mass., Wis., Colo., N. W.
T., Alta., Mont., B. C., Wash., Oreg., Calif.; Eurasia. Pollen: Unknown, but visits flowers
of Kalmia polifolia, Ledum glandulosum, Phyllodoce breweri, Rubus, Salix, Vaccinium.
Anthophora (Osmia) inermis Zetterstedt, 1838. Ins. Lapponica, v. 1, p. 466. °.
Osmia parietina Smith, 1844. Zoologist 2: 743. 2.
Osmia globosa Cresson, 1864. Ent. Soc. Phila., Proc. 3: 36. 2.
Osmia vulpecula Gerstaecker, 1869. Stettin. Ent. Ztg. 30: 335. °.
Osmia globosiformis Cockerell, 1910. Canad. Ent. 42: 311. d.
inurbana Cresson. Wyo., Colo., Idaho, Wash., Oreg., Calif., Que.?, Man., Alta.?, B. C.? Pollen:
Unknown, but visits flowers of Gilia multicaulis, Lotus davidsonii, Nemophila
integrifolia, Potentilla glandulosa, Senecio lugens, Streptanthus tortuosus.
Osmia inurbana Cresson, 1878. Amer. Ent. Soc., Trans. 7: 107. 6.
Osmia eutrichosa Cockerell, 1910. Canad. Ent. 42: 312. ¢.

2044 Hymenoptera in America North of Mexico

kineaidii Cockerell. Nebr., N. Mex., Idaho, B. C., Wash., Oreg., Calif. Parasite: Chrysura
sonorensis (Cam.), Epistenia sp., Leucospis affinis Say, Nemognatha scutellaris LeC.
Pollen: Unknown, but visits flowers of Allium parvum, Amsinckia douglasiana,
Brodiaea, Calochortus nudus, Chamaebatia foliolosa, Clarkia rhomboidea, C.
unguiculata, Fragaria californica, Geranium molle, Heterogaura heterandra, Linanthus
montanus, Lotus scoparius, L. subpinnatus, Lupinus micranthus, Monardella villosa,
Nemophila spatulata, Phacelia distans, Rhamnus californica, Sida hederacea,
Trifolium microcephalum, Vicia.
Osmia kincaidii Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 334. 2, 6.
lacus Sandhouse. Oreg., Calif.
Osmia (Nothosmia) lacus Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 125. 6.
laeta Sandhouse. Calif., N. Mex.? Parasite: Nemognatha scutellaris LeC. Pollen: Unknown, but
visits flowers of Arctostaphylos drupacea, A. patula, Astragalus, Brassica geniculata,
Cryptantha intermedia, Eriodictyon trichocalyx, Gilia exilis, Lotus argophyllus, L.
davidsonii, L. glaber, L. nevadensis, L. scoparius, L. strigosus, Nama parryi, Nemophila
integrifolia, Penstemon centranthifolius, P. labrosus, P. spectabilis, Phacelia imbricata,
P. heterophylla, P. ramosissima, Rubus leucodermis, Trifolium variegatum, Vicia
villosa.
Osmia laeta Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 18: 350. °.
Osmia aglaia Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 112. @.
malina Cockerell. Wash., Calif., Colo. Pollen: Unknown, but visits flowers of Arctostaphylos
drupacea, Ceanothus cordulatus, C. lewcodermis, C. integerrimus, Chamaebatia
foliolosa, Comandra umbellata, Lotus argophyllus, L. crassifolius, L. davidsonii, L.
nevadensis, Lupinus austromontanus, Mimulus guttatus, Phacelia ramosissima,
Streptanthus tortwosus, Trifolium gracilentwm.
Osmia malina Cockerell, 1909. Entomologist 42: 94. °.
mertensiae Cockerell. Colo., Calif.?, B. C.? Pollen: Unknown, but visits flowers of Lappula
floribunda, Mertensia.
Osmia mertensiae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 488. ¢.

Taxonomy: Cockerell, 1934. Amer. Mus. Novitates 732: 5. °.
nanula Cockerell. Idaho, Utah, Colo., B. C., Wash., Oreg. Pollen: Unknown, but visits flowers
of Phacelia.
Osmia nanula Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 339. 2.
Osmia phaceliae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 364. ?.
pagosa Sandhouse. Oreg., Colo., Calif. Pollen: Unknown, but visits flowers of Vicia californica.
Osmia (Nothosmia) pagosa Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 89. °.
pentstemonis Cockerell. Alta., Idaho, Utah, Colo., B. C., Wash., Oreg., Calif. Pollen: Unknown,
but visits flowers of Fragaria californica, Gormania obtusata, Penstemon heterodoxus,
P. newberryi. Predator: Philanthus pulcher Dalla Torre.
Osmia pentstemonis Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 451. 9.
pingreeana Michener. Colo.
Osmia pingreeana Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 408. 2.
potentillae Michener. Calif. Pollen: Unknown, but visits flowers of Calyptridium umbellatum,
Fragaria californica, Lotus nevadensis, L. scoparius, Navarretia divaricata, Nemophila
integrifolia, Phacelia ramosissima, Potentilla glandulosa.
Osmia potentillae Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 89. 2.

Taxonomy: Michener, 1949. Kans. Ent. Soc., Jour. 22: 56.
proxima Cresson. Alaska to N. S., south to Oreg., Idaho, Wyo., N. Dak., Ill. and Ga. Parasite:
Stelis subemarginata (Cress.). Pollen: Unknown, but visits flowers of Houstonia,
Penstemon, Rubus, Trifolium.
Osmia proxima Cresson, 1864. Ent. Soc. Phila., Proe. 3: 32. 6.
Osmia sericea Cresson, 1864. Ent. Soc. Phila., Proc. 3: 27. 6.
Osmia melanotricha Lovell and Cockerell, 1907. Psyche 14: 16. °.

Superfamily APOIDEA 2045

Biology: Fye, 1965. Canad. Ent. 97: 874, tables 2-4 (nest, parasite). —Medler, 1967. Ent. Soc.
Amer., Ann. 60: 342 (nest, parasite).
pulsatillae Cockerell. Idaho, Wyo., Colo., Wash., Oreg., Calif. Pollen: Unknown, but visits
flowers of Epilobium glaberrimum.
Osmia pulsatillae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 363. 6.
pusilla Cresson. Wash., Oreg., Calif., Colo., N. Mex.? Pollen: Unknown, but visits flowers of
Gayophytum diffusum, Nemophila integrifolia, Phacelia davidsonii, Phalacoseris
bolanderi, Potentilla glandulosa, Sanicula nevadensis.
Osmia pusilla Cresson, 1864. Ent. Soc. Phila., Proc. 3: 35. 3.
regulina Cockerell. Oreg., Calif. Pollen: Unknown, but visits flowers of Astragalus,
Calandrinia caulescens var. menziesii, Chaenactis glabriuscula, Cryptantha
intermedia, Gilia multicaulis, Lotus argophyllus, L. scoparius, Salvia mellifera,
Trifolium involucratum, T. repens, T. variegatum.
Osmia regulina Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 766. 2 (¢ misdet.).
sanctaerosae Cockerell. Calif. Pollen: Unknown, but visits flowers of Eriodictyon
californicum, Lotus argophyllus, L. davidsonii, Lupinus confertus.
Osmia sanctae-rosae Cockerell, 1910. Ent. News 21: 121. 2.
sequoiae Michener. Calif.
Osmia sequoiae Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 89. °.

Taxonomy: Michener, 1949. Kans. Ent. Soc., Jour. 22: 57.
tersula Cockerell. Alaska to Hudson Bay and N. B., south to Oreg., Colo, Wis. and Mich.
Parasite: Sapyga martinii Sm.? Pollen: Unknown, but visits flowers of Rubus.
Osmia tersula Cockerell, 1912. Canad. Ent. 44: 358. 6.
Osmia subarctica Cockerell, 1912. Canad. Ent. 44: 357. 2.

Biology: Medler, 1967. Ent. Soc. Amer., Ann. 60: 342-343, figs. 1-2, tables 1-2 (nest
architecture, life history, parasite).

tokopahensis Michener. Calif.
Osmia tokopahensis Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 87. °.

Taxonomy: Michener, 1949. Kans. Ent. Soc., Jour. 22: 58.
trevoris Cockerell. B. C. and Alta., south to Calif., Nev., Colo., and Wyo.
Osmia trevoris Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 341. °.
Osmia subtrevoris Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 451. 2.
Osmia corkinsi Sandhouse, 1924. Ent. News 35: 351. °.
tristella cyanosoma Cockerell. Calif. Pollen: Unknown, but visits flowers of Ceanothus, Dalea
saundersii, Diplacus aurantiacus, Lotus davidsonii, Lupinus longipes, Monardella,
Nemophila integrifolia, Penstemon grinnellii, Phacelia hydrophylloides, Prunus
emarginata, Rubus vitifolius, Senecio lugens.
Osmia cyanosoma Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 52. °.
tristella tristella Cockerell. B. C., Wash., Oreg., Calif., Idaho, Colo., Que.? Pollen: Unknown,
but visits flowers of Diplacus aurantiacus, Lepechinia calycina, Lotus scoparius.
Osmia tristella Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 340. °.
Osmia hypoleuca Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 366. 2.
virga Sandhouse. Mass. to Wis., south to Va. Pollen: Unknown, but visits flowers of Salix.
Osmia virga Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 130. 6.
zephyros Sandhouse. Calif. Ecology: Nests in oak gall. Pollen: Unknown, but visits flowers of
Collinsia parviflora.
Osmia zephyros Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 132. d, ?.

Genus OSMIA Subgenus EUTHOSMIA Sinha

Osmia subg. Euthosmia Sinha, 1958. Kans. Univ. Sci. Bul. 39: 235.
Type-species: Heriades glaucum Fowler. Orig. desig.
glauca (Fowler). Calif. Ecology: Nests in abandoned burrows and also trap-nests; uses mud for
cell construction. Parasite: Stelis sermaculata Ashm. Pollen: Oligolege of Collinsia

2046 Hymenoptera in America North of Mexico

including C. bicolor, C. heterophylla, C. sparsifolia, but visits other flowers presumably
for nectar including Alyssum maritimum, Convolvulus, Cryptantha, Geranium
dissectum, G. molle, Gilia tricolor, Lasthenia chrysostoma, Lupinus bicolor, L.
micranthus, Melilotus indica, Mimulus, Nemophila integrifolia, N. pulchella, Phacelia
distans, P. douglasii, P. rattani, Platystemon californicus, Plectritis macrocera, Rhus
trilobata, Trifolium microcephalum.

Heriades glaucum Fowler, 1899. Psyche 8: 405. 6.

Osmia exilis Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 351. 2.

Taxonomy: Snelling, 1967. South. Calif. Acad. Sci., Bul. 66: 103 (tax. status).

Biology: Linsley and MacSwain, 1941. South. Calif. Acad. Sci., Bul. 40: 129 (nest). —Rust and
Clement, 1972. Kans. Ent. Soc., Jour. 45: 523-528, 1 fig., 1 table (nest architecture, life
history, floral relationships, parasite, biological characteristics).

Genus OSMIA Subgenus MYSTACOSMIA Snelling

Osmia subg. Mystacosmia Snelling, 1967. South. Calif. Acad. Sci., Bul. 66: 104.
Type-species: Osmia nemoris Sandhouse. Orig. desig.

nemoris Sandhouse. B. C., Idaho and Mont., south to Calif. and Utah. Ecology: Nests in

abandoned burrows of Diadasia diminuta Cress. as well as trap-nests; uses masticated
leaf material from Malva rotundifolia and Sphaeralcea coccinea cemented with resin
for cell construction. Pollen: Collects pollen primarily from flowers of Penstemon and
various legumes; visitation records include Astragalus, Brassica, Brodiaea congesta, B.
elegans, B. laxa, B. lutea, B. pulchella, Calochortus luteus, C. venustus, Chamaebatia
foliolosa, Chaenactis glabriuscula, Cirsium, Clarkia biloba, C. cylindrica, C. dudleyana,
C. gracilis, C. pulchella, C. purpurea viminea, C. speciosa, C. unguiculata, Collinsia
callosa, Convolvulus malacophyllus, C. subacaulis, Cryptantha, Dodecatheon
hendersoni, Eriogonum fasciculatum, Geranium molle, Grindelia, Lasthenia
chrysostoma, Layia platyglossa, Linanthus parviflorus, Lotus scoparius, L.
subpinnatus, Lupinus, Medicago sativa, Mimulus, Monardella villosa, Orthocarpus
purpurascens, Penstemon, Phacelia ciliata, P. distans, Plectritis macrocera, Prunus
ilicifolia, Raphanus sativus, Rhus ovata, Rosa californica, Rubus, Salvia, Satureja,
Sisymbrium, Trifolium involucratum, T. microcephalum, T. repens, T. tridentatum,
Vicia americana, Wyethia.

Osmia nemoris Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 345. 6.

Osmia seclusa Sandhouse, 1924. Calif. Acad. Sci., Proce. (4) 13: 352. 2.

Osmia abdominalis Michener, 1935. Pan-Pacific Ent. 11: 184. ¢.

Taxonomy: Snelling, 1967. South. Calif. Acad. Sci., Bul. 66: 103 (synonymy).

Biology: Bohart, 1955. Ent. Soc. Wash., Proc. 57: 235-236 (nest architecture, life history).
—Rust and Clement, 1972. Kans. Ent. Soce., Jour. 45: 523-528, table 1 (nest architecture,
life history, biological characteristics). —MacSwain, Raven and Thorp, 1973. Calif. Univ.
Pubs. Ent. 70: 52 (floral relationships, as seclusa).

Genus OSMIA Subgenus MONILOSMIA Robertson

Monilosmia Robertson, 1903. Amer. Ent. Soc., Trans. 29: 166.
Type-species: Osmia simillima Smith. Orig. desig. and monotypic. (~Osmia
canadensis Cresson).
albolateralis albolateralis Cockerell. Alta. to N. Mex., west to B. C. and Wash.
Osmia dubia Cresson, 1864. Ent. Soe. Phila., Proc. 3: 29. 2. Preoce.
Osmia albolateralis Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 450. 9.
2Osmia enena Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 448. d.
albolateralis visenda Sandhouse. Oreg., Calif. Pollen: Unknown, but visits flowers of
Astragalus antisellii, A. douglasii, Clarkia cylindrica, C. rhomboidea, Collinsia
tinctoria, Erysimum asperum, Horkelia tillingi, Iris hartwegii, Lathyrus alfeldvi, Lotus
argophyllus, L. davidsonii, Montia perfoliata, Nemophila integrifolia, Penstemon

Superfamily APOIDEA 2047

spectabilis, Phacelia, Senecio lugens, Sidalcea malvaeflora, Vicia californica, V.
truncata, Viola purpurea.

Osmia visenda Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 355. @.

atrocyanea atrocyanea Cockerell. Wash., Oreg., Calif., Colo., Utah, Nev. Parasite: Sapyga

angustata Cress. Pollen: Unknown, but visits flowers of Clarkia rhomboidea,
Cryptantha intermedia, Cirsium, Eriodictyon californicum, Lotus glaber, L. scoparius,
Lupinus, Penstemon laetus, Phacelia, Ribes, Rubus ursinus, Sidalcea malvaeflora,
aD involucratum, T. variegatum, Verbena lasiostachys, Vicia californica, V.
villosa.

Osmia atrocyanea Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 344. 9.

Osmia senior Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 368. °.

Osmia pseudamala Cockerell, 1910. Canad. Ent. 42: 312. d.

Taxonomy: Cockerell, 1930. Ann. and Mag. Nat. Hist. (10) 5: 163. d, 9.

atrocyanea putata Cockerell. South. Calif. Pollen: Unknown, but visits flowers of
Arctostaphylos drupacea, A. patula, Cryptantha intermedia, Iris hartwegii, Lathyrus
californicus, Lupinus confertus, L. latifolius, Nemophila integrifolia, Penstemon
spectabilis, Ribes roezlii, Vicia californica.
Osmia putata Cockerell, 1910. Ent. News 21: 272. 2.
brevis brevis Cresson. Wyo., Colo., Utah, Ariz., B. C., Oreg., north. and cent. Calif.
Osmia brevis Cresson, 1864. Ent. Soe. Phila., Proc. 3: 36. °.
Osmia wilmattae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 228. 2.
2Osmia wheeleri Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 449. ¢.
brevis peridonea Sandhouse. South. Calif. Pollen: Unknown, but visits flowers of Astragalus
antisellu, Arctostaphylos drupacea, Castilleia, Clarkia, Collinsia heterophylla, C.
tinctoria, Cryptantha intermedia, C. lepida, Eriodictyon californicum, E. crassifolium,
Fragaria californica, Gilia exilis, Horkelia tillingii, Lotus davidsonii, L. scoparius,
Lupinus bicolor, Nama parryi, Penstemon antirrhinoides, P. grinnellii, P.
heterophyllus, P. spectabilis, Phacelia andersonii, P. davidsonii, Phalacoseris bolanderi,
Psoralea californica, Rhus trilobata, Salvia carnosa, Streptanthus bernardinus, S.
tortuosus, Taraxacum.
Osmia peridonea Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 348. d.
bridwelli Sandhouse. Nev., Calif. Pollen: Unknown, but visits flowers of Agastache urticifolia,
Brodiaea coronaria, B. lutea, Chamaebatia foliolosa, Glycyrrhiza lepidota, Iris
hartwegii, Lathyrus sulphureus, Lotus nevadensis, Lupinus albifrons, L. densiflorus, L.
succulentus, Mimulus guttatus, Penstemon spectabilis, Psoralea physodes, Sedum,
Streptanthus tortuosus, Vicia californica, Wyethia helenioides.
Osmia (Nothosmia) bridwelli Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 100. 6, 2.
cara Cockerell. South. Calif., Nev. Pollen: Unknown, but visits flowers of Cirsium
californicum, Clarkia, Dicentra chrysantha, Eriodictyon californicum, E. trichocalyx,
Lotus crassifolius, L. davidsonii, L. glaber, L. scoparius, Lupinus densiflorus, L. nanus,
Mimulus, Nama parryi, Penstemon heterophyllus, P. laetus, P. spectabilis, Trichostema
parishii.
Osmia cara Cockerell, 1910. Ent. News 21: 271. °.
cyanella Cockerell. Wash., Oreg., Calif., Colo., Utah, Nev. Pollen: Unknown, but visits flowers
of Chamaebatia foliolosa, Cirsium californicum, Collinsia heterophylla, C. tinctoria,
Cryptantha intermedia, Eriodictyon californicum, Eriophyllum confertiflorum,
Fragaria californica, Heterogaura californica, Iris hartwegii, Lepechinia calycina,
Linanthus parviflorus, Lotus davidsonii, L. nevadensis, L. scoparius, Lupinus,
Mimulus guttatus, Penstemon grinnellii, P. heterodoxus, P. speciosus, Phacelia
californica, P. heterophylla, P. imbricata, P. ramosissima, Potentilla glandulosa,
Ranunculus californicus, Streptanthus tortuosus.
Osmia cyanella Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 340. °.
Osmia aprilina Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 364. 6.
Osmia aprilina atrovirens Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 347. d.
Osmia brevior Michener, 1936. Amer. Mus. Novitates 875: 20. 2.
cyaneonitens Cockerell. S. Dak., Colo.
Osmia cyaneonitens Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 448. 3.

2048 Hymenoptera in America North of Mexico

Osmia brevihirta Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 365. @.
densa densa Cresson. Mont. to N. Mex., west to B. C., cent. Calif., and Ariz. Pollen: Polylectic,
visits a wide variety of flowers including Arctostaphylos crustacea, A. mariposa,
Brodiaea lutea, B. pulchella, Calochortus nudus, Ceanothus integerrimus, Chamaebatia
foliolosa, Cirsium californicum, Clarkia breweri, C. dudleyana, C. rhomboidea, C.
unguiculata, Convolvulus malacophyllus, Eriodictyon californicum, Eriogonum
imflatum, Gilia capitata, Hackelia jessicae, Horkelia fusca, Iris hartwegii, Lathyrus
graminifolius, L. sulphureus, Limnanthes douglasii, Lotus argophyllus, Lupinus
albifrons, L. bicolor, L. latifolius, L. micranthus, L. stiversi, Mimulus guttatus, M.
primuloides, M. suksdorfii, Monardella lanceolata, Penstemon spectabilis, Phacelia
heterophylla, Ranunculus californicus, Ribes nevadensis, Spraguea umbellata,
Streptanthus tortwosus, Thermopsis malacophyllus, Verbena lasiostachys, Vicia
californica.
Osmia densa Cresson, 1864. Ent. Soe. Phila., Proc. 3: 25. 2.
Osmia olivacea Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 362. d.
Osmia proposita Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 354. 2.
densa pogonigera Cockerell. Calif. Parasite: Tricrania stansburyi Hald. Pollen: Polylectic,
visits a wide variety of flowers including Arctostaphylos drupacea, A. patula,
Astragalus antisellii, A. parishii, Barbarea orthoceras, Ceanothus cordulatus, Clarkia
breweri, C. unguiculata, Convolvulus, Cryptantha intermedia, Dicentra chrysantha,
Epilobium angustifolium, Horkelia bernardina, Lathyrus, Lotus crassifolius, L.
davidsonii, L. scoparius, Lupinus austromontanus, L. confertus, L. formosus, L. gray,
Malus, Marrubium vulgare, Nemophila integrifolia, Penstemon grinnellii, P.
spectabilis, Phacelia davidsonii, P. distans, P. heterophylla, Rhamnus californica,
Rhododendron occidentale, Ribes cereum, R. roezlii, Vicia arvensis, V. californica.
Osmia pogonigera Cockerell, 1910. Ent. News 21: 121. 9.
Osmia celsa Sandhouse, 1924. Calif. Acad. Sci., Proe. (4) 13: 354. @.
felti Cockerell. New England to Minn., south to West Va.
Osmia felti Cockerell, 1911. Ent. News 22: 18. 9.
gabrielis Cockerell. Oreg., Calif. Pollen: Unknown, but visits flowers of Brodiaea lutea,
Castilleia, Clarkia cylindrica, Diplacus aurantiacus, Glycyrrhiza lepidota, Iris
hartwegii, Lathyrus sulphureus, Lotus glaber, L. scoparius, Lupinus albifrons, L.
formosus, L. subspicata, Marrubium vulgare, Orthocarpus lithospermoides, Penstemon
speciosus, Psoralea physodes, P. rigida, Salvia mellifera, Streptanthus bernardinus,
Trifolium variegatum, Vicia californica.
Osmia gabrielis Cockerell, 1910. Ent. News 21: 120. °.
hendersoni Cockerell. Mont., Wyo., Colo., Utah, Oreg. Calif. (Sierra Nevada Mts.).
Osmia Hendersoni Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 365. °.
hesperos Sandhouse. Calif., Oreg.
Osmia hesperos Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 131. 6.
iridis Cockerell and Titus. Colo., N. Mex., Calif. Pollen: Unknown, but visits flowers Collinsia
callosa, Malus.
Osmia iridis Cockerell and Titus, 1902. Amer. Nat. 36: 816. ¢.
juxta juxta Cresson. Alta. to N. Mex., west to Idaho and Ariz.
Osmia juxta Cresson, 1864. Ent. Soc. Phila., Proc. 3: 19. 9.
Osmia theta Sandhouse, 1925. Canad. Ent. 57: 34. ¢.
Osmia subpurpurea jamesi Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 409. °.
juxta subpurpurea Cockerell. B. C. to Calif. Pollen: Unknown, but visits flowers of Brodiaea
congesta, Geranium, Lotus scoparius, Lupinus lobbii var. lyallii, L. longipes, Mertensia,
Penstemon speciosus, Phalacoseris bolanderi, Plectritis macrocera, Salvia columbariae,
Trifolium microcephalum, Wyethia angustifolia.
Osmia subpurpurea Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 344. 9.
paradisica Sandhouse. Wash., Oreg., Calif. (Sierra Nevada Mts.). Pollen: Unknown, but visits
flowers of Eriogonum.
Osmia paradisica Sandhouse, 1924. Calif. Acad. Sci., Proce. (4) 13: 348. ¢.
rawlinsi Sandhouse. Wyo., Calif.
Osmia (Nothosmia) rawlinsi Sandhouse, 1939. Ent. Soe. Wash., Mem. 1: 115. 2.

Superfamily APOIDEA 2049

rostrata Sandhouse. Calif. Pollen: Unknown, but visits flowers of Amsinckia douglasiana,
Grossularia, Phacelia tanacetifolia.
Osmia rostrata Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 351. 9.
sculleni Sandhouse. Oreg., Calif., Utah, Colo. Pollen: Unknown, but visits flowers of Hackelia
jessicae.
Osmia (Nothosmia) sculleni Sandhouse, 1939. Ent. Soc. Wash., Mem. 1: 82. @.

simillima Smith. N. S., west to N. W. T. and B. C., south to Oreg., Ariz., and N. C. Parasite:

Leucospis affinis Say, Stelis suwhemarginata (Cress.). Pollen: Unknown, but visits
flowers of Althaea, Barbarea, Oxalis, Penstemon, Ranunculus, Rubus, Vicia, Viola.

Osmia simillima Smith, 1853. Cat. Hym. Brit. Mus., v.11, p. 142. 2, 6.

Osmia canadensis Cresson, 1864. Ent. Soc. Phila., Proc. 3: 33. 3.

Osmia cognata Cresson, 1864. Ent. Soc. Phila., Proc. 3: 33. 3.

Osmia major Robertson, 1902. Ent. News 13: 79. 2 (36 misdet.).

Osmia chlorops Cockerell and Titus, 1902. Amer. Nat. 36: 816. 6.

Osmia stasima Lovell, 1909. Ent. News 20: 125. °.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 332. —Cockerell, 1906. Amer. Mus.
Nat. Hist., Bul. 22: 448. ¢. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 80-81,
figs. 28, 32, 35 (redescription, synonymy).

Biology: Packard, 1867. Amer. Nat. 1: 377 (nest). —Saunders, 1872. Canad. Ent. 4: 237 (nest,
as canadensis). —Graenicher, 1935. Ent. Soc. Amer., Ann. 28: 285 (nest, parasite).

Genus OSMIA Subgenus TRICHINOSMIA Sinha

Osmia subg. Trichinosmia Sinha, 1958. Kans. Univ. Sci. Bul. 39: 244.

Type-species: Osmia latisulcata Michener. Monotypic and orig. desig.
latisulcata Michener. Ariz., Nev., Calif. Pollen: Unknown, but visits flowers of Amsinckia,
Arctostaphylos glandulosa, Astragalus douglasii, Chaetopappa aurea, Dalea,

Cryptantha intermedia, Eriodictyon californicum, Lotus scoparius, L. strigosus,
Lupinus, Salix, Salvia carnosa, S. columbariae.
Osmia latisulcata Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 86. 9.

Genus OSMIA Subgenus DICERATOSMIA Robertson

Diceratosmia Robertson, 1904. Amer. Ent. Soc., Trans. 29: 166.
Type-species: Osmia conjuncta Cresson. Monotypic and orig. desig. (~Osmia
quadridentata Cresson).

Revision: Michener, 1949. Ent. Soc. Amer., Ann. 42: 258-264 (Nearctic spp.).

Taxonomy: Hurd and Michener, 1955. Calif. Ins. Surv., Bul. 3: 215-217, fig. 4, pl. 11, map 112
(Calif. spp.). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 81-85, figs. 29, 31-82,
table 4 (eastern U. S. spp.).

botitena (Cockerell). Tex.
Osmia botitena Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 30. °.

Osmia conjuncta marilaunidii Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 363. 2.
conjuncta (Cresson). Ont. to Minn., south to N. C. and Tex. Ecology: Nests in snail shells.
Pollen: Polylectic, visits flowers of many families, especially Leguminosae and

Scrophulariaceae, including Anemonella, Blephilia, Cardamine, Cercis, Collinsia,
Dentaria, Fragaria, Geranium, Hydrophyllum, Lupinus, Osmorrhiza, Polemonium,
Psoralea, Ranunculus, Rubus, Scutellaria, Stellaria, Trifoliwm, Viola.

Osmia conjuncta Cresson, 1864. Ent. Soc. Phila., Proc. 3: 31. 2.

Osmia 4-dentata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 107. 3. Preoce.

Osmia cressoni Dalla Torre, 1896. Cat. Hym., v. 10, p. 392. N. name.

Osmia quadridentata Dalla Torre, 1896. Cat. Hym., v. 10, p. 392. Emend.

Biology: Rau, 1937. Ent. Soc. Amer., Ann. 30: 330 (nest).

2050 Hymenoptera in America North of Mexico

subfasciata miamiensis Mitchell. Southern Fla. Pollen: Unknown, but visits flowers of
Crotalaria.
Osmia (Diceratosmia) subfasciata miamiensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech.
Bul. 152: 84. 2, ¢.
subfasciata subfasciata Cresson. N. J. to Fla., west to south. Calif.; northern Mexico. Ecology:
Nests in abandoned beetle burrows, also accepts artificial nesting devices. Parasite:
Melittobia chalybii Ashm. Pollen: Polylectic, visits a wide variety of flowers including
Achillea, Acacia greggii, Amorpha fruticosa, Astragalus, Cercis canadensis, Coreopsis,
Gaillardia pulchella, Helenium tenuifolium, Helianthus, Heliotropium curassavicum
var. oculatum, Larrea tridentata, Lesquerella, Lotus scoparius, Machaeranthera
tenacetifolium, Marilaunidium origanifolium, Marrubium vulgare, Medicago sativa,
Melilotus officinalis, Monarda citriodora, Oxalis, Palafoxia linearis, Parkinsonia
aculeata, Phacelia, Pluchea sericea, Prosopis juliflora, Prunus, Rhus, Rubus, Salix,
Sphaeralcea, Vicia.
Osmia subfasciata Cresson, 1872. Amer. Ent. Soc., Trans. 4: 261. 2, 6.
Osmia conjunctoides Robertson, 1893. Amer. Ent. Soc., Trans. 20: 276. d.
Osmia punctata Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 85. °.

Biology: Linsley, 1946. Econ. Ent., Jour. 39: 24 (nest sites, pollination of alfalfa). —Mitchell,
1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 84 (nest). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 311-312 (nest architecture, life history, parasite).

Genus OSMIA Subgenus Unassigned

angustipes Cockerell. Colo.
Osmia angustipes Cockerell, 1933. Pan-Pacific Ent. 9: 158. ¢.
claremontensis Michener. Mont., Wash., Oreg., Calif. Pollen: Unknown, but visits flowers of
Arctostaphylos patula, Collinsia heterophylla, Cryptantha intermedia, Phacelia distans,
Trifolium repens.
Osmia claremontensis Michener, 1936. South. Calif. Acad. Sci., Bul. 35: 84. 6.

Taxonomy: Snelling, 1967. South. Calif. Acad. Sci., Bul. 66: 106-107 (tax. status).

foxi Cameron. N. Mex.
Osmia Foxi Cameron, 1901. Amer. Ent. Soc., Trans. 27: 316. 6.

NOMEN NUDUM IN OSMIA
Osmia pacifica Packard, 1892. Psyche 6: 340.

Genus MEGACHILE Latreille

This genus, which is nearly cosmopolitan in distribution, has until recently included all of the
so-called leaf-cutter bees regardless of whether the various species cut leaves with which to
make their nests or not. As currently restricted, those species which make their nests with
pieces of leaves or petals belong to the genus Megachile while those which use resin, mud, or
other such material have been assigned to Chalicodoma Lepeletier and Creightonella Cockerell.
The latter genus does not occur in the Western Hemisphere. Some authors have concluded that
Chelostomoides Robertson warrants generic rank rather than subgeneric status within the
genus Chalicodoma as it is treated here.

Revision: Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 295-361, pls. XX-XXI (Part I,
classification of Nearctic subgenera including key; n. spp.). —Mitchell, 1935. Amer. Ent.
Soe., Trans. 61: 1-44, pl. I (Part II, treats Nearctic spp. of subgenera Cressoniella,
Litomegachile, Neomegachile). —Mitchell, 1935. Ent. Soc. Amer., Trans. 61: 155-205, pls.
VIII-IX (Part III, treats Nearctie spp. of the subgenera Anthemois and Delomegachile).
—Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 117-166, pls. VIII-XI (Part IV, treats
Nearctic spp. of the subgenera Derotropis, Megachiloides, Phaenosarus, and
Xanthosarus). —Mitchell, 1937. Amer. Ent. Soc., Trans. 62: 323-382, pls. XXII-XXVI
(Part V, treats Nearctic spp. of the subgenus Xeromegachile). —Mitchell, 1937. Amer.
Ent. Soc., Trans. 63: 45-83, pls. V-VI (Part VI, treats Nearctic spp. of the subgenera
Acentron, Argyropile, Leptorachis, Melanosarus and Pseudocentron). —Mitchell, 1937.

Superfamily APOIDEA 2051

Amer. Ent. Soc., Trans. 63: 175-206, pls. XII-XIII (Part VII, treats Nearctic spp. of the
subgenus Sayapis). —Mitchell, 1937. Amer. Ent. Soc., Trans. 63: 381-426, pls. XX VI-XXIX
(Part VIII, treats Nearctic spp. of the subgenus Chelostomoides; addenda and index to
Parts I-VIII).

Taxonomy: Michener, 1962. N. Y. Ent. Soc., Jour. 70: 17-29 (classification). — Mitchell, 1962.
N. C. Agr. Expt. Sta. Tech. Bul. 152: 108-191, figs. 1, 36-62, tables 5-6 (eastern U. S. spp.).
—Pasteels, 1965. Mus. Royal I’ Afrique Centr., Ann. Sci. Zool. 137: ix, 579 pp.
(classification). —Butler, 1965. Ariz. Agr. Expt. Sta. Tech. Bul. 187: 1-19 (Ariz. spp.).
—Michener, 1965. Amer. Mus. Nat. Hist., Bul. 130: 185-186 (classification). —Medler and
Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 1-80 (Wis. spp.). —Stephen, Bohart and Torchio,
1969. The biology and external morphology of bees, pp. 53-54 (classification).

Biology: Hobbs and Lilly, 1954. Ecology 35: 453-462 (alfalfa pollination in southern Alberta).
—Krombein, 1967. Trap-nesting wasps and bees, pp. 320-337, pl. 19, figs. 92-97; pl. 20, figs.
98-100 (life histories, nest associates). —Holm and Skou, 1972. Ent. Scandinavica 3:
169-180, 6 figs., 8 tables (nesting habits, parasites). —Hurd and Linsley, 1975. Smithsn.
Contrib. Zool. 193: 37-38 (Larrea visiting spp.).

Morphology: Mitchell, 1935. Amer. Ent. Soc., Trans. 61: 3-9 (male sternites and genital
armature). —Pasteels and Pasteels, 1971. Acad. Sci. Paris, Compt. Rend. 273: 1481-1483
(tergal glands).

Genus MEGACHILE Subgenus LITOMEGACHILE Mitchell

Megachile subg. Litomegachile Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 301, 308.
Type-species: Megachile brevis Cresson. Orig. desig.
brevis brevis Say. U. S. and south. Canada. Ecology: Nests in a wide variety of situations

including within dead plant stalks of Ambrosia trifida, Cirsium, Helianthus annuus, H.
tuberosa, Vernonia interior, Zea mays, in a rolled leaf of Eupatoriwm perfoliatum,
among green leaves of Erigeron canadensis, among rocks and under dried cow chips, in
a termite tunnel, and in various burrows and holes in the soil. Parasite: Anthrax
irroratus Say, Coelioxys novomexicana Ckll., C. octodentata Say, C. salinaria CkIl., C.
sayi Robt., Leucospis affinis Say, Nemognatha nigripennis Lec. Pollen: Polylectic, visits
a wide variety of flowers especially Compositae, Leguminosae and Labiatae; visitation
records include Abutilon, Acerates, Althaea rosea, Amorpha, Apios, Asclepias eriocarpa,
Aster canescens, Astragalus, Baptisia, Bidens laevis, Blephilia, Boltonia, Brauneria,
Campanula, Cassia, Ceanothus, Cephalanthus, Chaenactis glabriuscula, Chamacrista
fasciculata, Chrysanthemum, Chrysothamnus, Cicuta, Clarkia bottae, Convolvulus
occidentalis, Coreopsis lanceolata, Cornus, Crotalaria, Croton californicus, Cryptantha
intermedia, Cuscuta, Cynoglossum, Desmodium, Dianthera, Encelia farinosa,
Engelmania, Erechthites, Erigeron stenophyllus, Eryngium, Eupatorium, Fayopyrum,
Frankenia grandiflora, Gaillardia pulchella, Galactia, Geranium maculatum, Gerardia,
Gilia, Gossypium, Grindelia elata, Gutierrezia californica, G. sarothrae, Haplopappus
palmeri, H. vernonioides, Helenium, Helianthus annuus, Heliopsis, Heliotropium
curassavicum, Hemizonia fasciculata, Hieracium, Heterotheca grandiflora, Hibiscus,
Hydrophyllum, Hypericum perfoliatum, Impatiens, Koellia, Krigia, Kuhnistera,
Lactuca, Lepidospartum squamatum, Linaria, Lippia, Lobelia, Lotus scoparius,
Lupinus parryi, Lycopus, Lythrum, Marrubium vulgare, Medicago sativa, Melilotus
alba, Mentha piperita, Nepeta, Nutilla stricta, Oenothera, Onobrychis, Penstemon,
Petalostemon compactus, Phacelia distans, P. heterophylla, P. ramosissima, Phaseolus,
Phyla lanceolata, P. nodiflora rosea, Physostegia, Pluchea camphorata, Polygala,
Polygonum auberti, Psoralea, Psoralioides, Pycnanthemum, Rhus, Rosa, Rubus,
Rudbeckia hirta, Ruellia, Sabatia, Saggittaria, Salix lasiolepis, Salvia, Schinus molle,
Scutellaria, Senecio, Sidalcea reticulata, Silphium, Solidago canadensis, Sphaeralcea
fasciculata, Stachys, Stephanomeria exigua, S. virgata, Strophostyles, Suriana,
Tephrosia, Tecrium, Tradescantia, Trifolium pratense, T. repens, Verbena, Verbesina,
Vicia.

Megachile brevis Say, 1837. Boston Jour. Nat. Hist. 1: 407. d, °.

Megachile lanuginosa Smith, 1853. Cat. Hym. Ins. Coll. Brit. Mus., v. 1, p. 190. 2, ¢.

?Megachile nupta Cresson, 1872. Amer. Ent. Soc., Trans. 4: 268. °.

2052 Hymenoptera in America North of Mexico

Megachile perbrevis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 127. ¢.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1057, figs. 151-153, 157 (larva).
—Michener, 1953. Kans. Univ. Sci. Bul. 35: 1663 (tax. status).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 39 (nesting habits). —Rau, 1934. Acad.
Sei. St. Louis, Trans. 28: 220 (nesting habits). —Rockwood, 1951. Pan-Pacifie Ent. 27: 155
(nest, parasite). —Michener, 1953. Kans. Univ. Sci. Bul. 35: 1659-1748, 31 figs. (life history,
nest and associates). —Donahue, 1954. Nature Mag. 47: 300-302 (life history). —Medler and
Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 19-20 (life history, parasites).

brevis onobrychidis Cockerell. Southwest. U. S., intergrading with typical brevis Say in a wide
area from Nebr. and Tex. to Pacific Coast from Calif. to B. C. and Idaho. Ecology: Uses
grape leaves for nesting material. Parasite: Coelioxys octodentata Say. Pollen:
Polylectic, visits a wide variety of flowers especially Compositae, Leguminosae and
Labiatae, visitation records include Achillea lanulosa, A. millefolium, Agastache,
Asclepias eriocarpa, A. mexicana, Bigelovia, Brassica geniculata, Calochortus luteus, C.
venustus, Calycadenia multiglandulosa, Calyptridiwm umbellatum, Centaurea
solstitialis, Cercidium, Chamaebatia foliolosa, Chrysothamnus nauseosus speciosus, C.
viseidiflorus typicus, Cirsium californicum, Clarkia dudleyana, C. speciosa speciosa, C.
unguiculata, Cleomella obtusifolia, Convolvulus aridus, Cordylanthus filifolius, C.
pilosus, Coreopsis lanceolata, Corethrogyne bernardina, Crepis vescicaria, Croton
californicus, Cryptantha intermedia, Eriastrum virgatum, Evrigeron canadensis,
Eriodictyon, Eriogonum fasciculatum, E. gracile, E. latifolium var. nudum, Eryngium
aristatulum, Eschscholzia californica, Foeniculum vulgare, Frankenia grandiflora,
Grindelia camporum, G. hallii, Gutierrezia californica, G. lucida, G. sarothrae,
Haplopappus arborescens, H. bloomeri var. angustatus, H. pachylepis, H. palmeri, H.
venetus, Helenium, Helianthus, Heliotropium curassavicum, Hemizonia fasciculata, H.
luzulaefolia, H. pungens, H. wrightii, Heterotheca grandiflora, Hoffmanseggia
microphylla, Lotus americanus, L. glaber, L. hamatus, L. humistratus, L. purshianus,
L. scoparius, L. strigosus, Lupinus micranthus, Lythrum californicum, Malacothamnus
arcuatus, Marrubium vulgare, Medicago sativa, Melilotus alba, M. indica, Mimulus
brevipes, Onobrychis, Phacelia distans, P. ramosissima, Phyla lanceolata, P. nodiflora
var. rosea, Polygonum aubertii, Ribes, Rudbeckia serotina, Prosopis, Salix, Salvia
clevelandii, Sida hederacea, Solidago californica, S. occidentalis, Spiraea, Stanleya
pinnata, Stephanomeria exigua, S. virgata, Trichostema lanceolatum laxum, T. parishii,
T. repens, T. variegatum, Verbena lasiostachys, Wislizenia refracta.

Megachile onobrychidis Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 266. ¢.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1663 (tax. status).
brevis pseudobrevis Mitchell. N. C. to Fla. and Miss. Pollen: Presumably polylectic, visits
flowers of Afzelia, Chrysopsis, Crotalaria, Erigeron, Galactia, Gaylussacia, Helenium,
Helianthus, Lupinus, Melilotus, Opuntia, Polygonum, Rhus, Rubus, Solidago, Suriana,
Trilisa, Vaccinium.
Megachile (Litomegachile) brevis var. pseudobrevis Mitchell, 1936. Amer. Ent. Soc., Trans.
GL 20M ios

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 116 (tax. status).

coquilletti Cockerell. Idaho and Utah, west to B. C. to south. Calif. and Ariz. Pollen:

Apparently polylectic, visits a wide variety of flowers including Asclepias erosa, A.
mexicana, A. speciosa, Baccharis, Brassica geniculata, B. incana, Brodiaea crocea,
Centaurea solstitialis, Cercidium torreyanum, Cirsium vulgare, Clarkia unguiculata,
Cleome serrulata, Cordylanthus pilosus, C. rigidus, Croton californicus, Cryptantha
intermedia, Eriastrum virgatum, Erigeron stenophyllus, E. elatwm, E. fasciculatum
var. polifolium, E. latifolium var. nudum, E. plumatella, Eriophyllum confertiflorin,
Frasera parryi, Grindelia, Gutierrezia sarothrae, Helianthus gracilentus, Lotus
americanus, L. glaber, L. scoparius, Marrubium vulgare, Medicago sativa, Melilotus
alba, Phacelia ramosissima, Pluchea camphorata, Phyla filiformis, P. lanceolata,
Rhamnus californicus, Solidago, Swertia parryi.

Megachile mendica coquilletti Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 535. 6.

Superfamily APOIDEA 2053

gentilis Cresson. Tex. to Calif., Nev., Oreg. and Idaho, adventive and established in Hawaii.

Ecology: Nests in twigs of Sambucus and also in trap-nests. Parasite: Anthrax atriplex
Marston, A. daphne (O.S.), A. irroratus Say, A. melanopogon (Bigot), Aritranis notata
sierrae Townes, Coeliorys novomexicana Ckll., Leucospis affinis Say, Tetrastichus
megachilidis Burks. Pollen: Apparently polylectic, visits flowers of Acacia, Amorpha
fruticosa, Asclepias erosa, A. speciosa, Aster canescens, A. spinosus, Bigelovia,
Baccharis, Brassica geniculata, B. incana, Cercidium, Cereus engelmannii,
Chamaebatia foliolosa, Chrysopsis fastigiata, Chrysothamnus, Clarkia amoena, C.
dudleyana, C. unguiculata, Clematis, Cleomella obtusifolia, Collinsia tinctoria,
Condalia, Cordylanthus filifolius, C. nevinii, C. pilosus, C. rigidus, Croton californicus,
Cryptantha intermedia, Dalea, Echinocactus, Erigeron stenophyllus, Eriogonum
fasciculatum, E. gracile, E. latifolium var. nudum, Eriophyllum confertiflorum,
Grindelia camporum, Gutierrezia sarothrae, Haplopappus acradenius, H. arborescens,
H. bloomeri var. angustatus, H. vernonioides, Helenium bigelovii, Helianthus annuus,
Hemizonia pungens, Heteromeles arbutifolia, Heterotheca grandiflora, Hyptis emoryi,
Larrea tridentata, Lathyrus splendens, Lepidium, Lessingia glandulifera, Lippia, Lotus
argophyllus, L. glaber, L. heermannii, L. nevadensis, L. purshianus, Lupinus longipes,
Marrubium vulgare, Medicago sativa, Melilotus alba, M. indica, Monarda citriodora,
Nolina parryi, Opuntia, Phacelia distans, P. heterophylla, P. ramosissima, Phyla
nodiflora var. rosea, Pluchea camphorata, Polygonum, Prosopis, Ratibida columnaris,
Rhamnus californica, Sapindus, Senecio douglasii, Sisymbrium altissimum, Solidago
confinis, S. occidentalis, Spraguea umbellatum, Swertia neglecta, Trifolium repens, T.
variegatum, Vauquelinia, Verbena lasiostachys, Verbesina, Viguiera multiflora,
Wislizenia refracta. Predator: Trichodes horni Wolcott and Chapin.

Megachile gentilis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 267. ¢.

Megachile palmarum Perkins, 1899. Fauna Hawaii., v. 1, p. 114. 6, @.

Megachile murinella Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 263. 2.

Biology: Bechtel, 1958. Pan-Pacific Ent. 34: 12 (nest, parasite). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 320-321, pl. 20, figs. 99-100 (nest architecture, life
history, supersedure, parasites, predator).

lippiae Cockerell. Mont., Nebr., Tex., west to Utah and Calif. Ecology: Uses cuttings from rose
leaves for nest material. Pollen: Apparently polylectic, visits a wide variety of flowers
including Acacia, Agave, Althaea rosea, Amorpha, Asclepias, Aster, Baccharis,
Cercidium, Cirsium, Cucurbita, Erigeron, Eriogonum fasciculatum var. polifolium,
Haplopappus, Heleniwm, Helianthus, Hoffmanseggia, Hyptis emoryi, Larrea tridentata,
Lippia, Lotus, Marrubiwm vulgare, Medicago, Melilotus, Mortonia, Nama, Nolina,
Opuntia, Petalostemon, Prosopis, Sapindus, Sphaeralcea, Symphoricarpos, Trifolium,
Verbesina.

Megachile cleomis var. lippiae Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 15. 9, d.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 37 (floral relationship with
Larrea).

mendica mendica Cresson. Ont. and Maine to Fla., west to S. Dak., Colo., Ariz. and Calif.
Ecology: Nests in rose canes and trap nests. Parasite: Coeliorys octodentata Say, C.
sayi Robt., Melittobia chalybii Ashm., Physocephala marginata (Say). Pollen: Polylectie,
visits a wide variety of flowers, especially Compositae, Leguminosae, and Labiatae;
visitation records include Acerates, Agastache, Amorpha, Apios, Aralia, Asclepias
tuberosa, Aster, Baptisia, Bidens, Blephilia, Boltonia, Borrichia, Brassica nigra,
Brauneria, Buddleia campanula, Camassia, Cassia, Ceanothus, Cephalanthus,
Chaemaecrista, Chrysanthemum, Chrysopsis, Cicuta, Cirsium, Clematis, Cleomella
obtusifolia, Clethra, Conoclinum caelestinum, Coreopsis stellata, Cosmos, Crataegus,
Crotalaria, Cyrilla, Daucus, Desmodium, Diospyros, Elephantopus carolinianus,
Erigeron, Eriogonum, Eryngium, Eupatorium, Flaveria, Galactia, Gaura, Gaylussacia,
Geranium, Haplopappus, Helenium, Helianthus, Heliopsis helianthoides, Hypericum,
Ilex, Impatiens, Itea, Koellia mutica, Lepachys, Lespedeza, Linaria, Lupinus, Lycopus,
Lythrum, Marrubium vulgare, Medicago, Melilotus alba, Monarda, Nepeta, Nyssa,
Ocimum, Oenothera, Opuntia, Phacelia, Phaseolus, Polygala incarnata, Polygonum,

2054 Hymenoptera in America North of Mexico

Prunella, Psedera, Psoralea, Pycnanthemum, Pyrrhopappus, Rhus, Richardia, Robinia,

Rosa, Rubus, Rudbeckia, Salix, Senecio, Silphium, Solidago, Specularia, Spiraea,

Stachys, Strophostyles, Suriana, Symphoricarpos, Tephrosia, Teucrium, Trifolium,

Trilisia, Vaccinium, Verbena, Verbesina, Vernonia glauca, Veronica, Veronicastrum

virginicum, Vicia, Zinnia, Zizia. Predator: Pyemotes ventricosus (Newport), Vidia sp.
Megachile mendica Cresson, 1878. Amer. Ent. Soc., Trans. 7: 126. °.

Taxonomy: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 39 (life history).

Biology: Koerber and Medler, 1959. Wis. Acad. Sci., Arts and Letters 47: 56 (geogr. records).
—Medler, 1965. Ent. Soc. Wash., Proc. 67: 113-116, 1 table (life history, nest architecture,
cocoon, parasites). —Krombein, 1967. Trap-nesting wasps and bees, pp. 322-325 (life
history, nest architecture, supersedure, parasites, predators). —Medler and Lussenhop,
1968. Wis. Univ. Res. Bul. 274: 25-26 (nesting habits).

mendica snowi Mitchell. Colo., N. Mex., Ariz. Pollen: Apparently polylectic, visits flowers of

Asclepias, Ceanothus, Chilopsis, Cleome, Eriogonum, Geranium, Gilia, Helenium,
Heliopsis, Heterotheca, Ligustrum, Lupinus, Marrubium vulgare, Melilotus, Robinia,
Rudbeckia, Verbesina, Viguiera.

Megachile mendica snowi Mitchell, 1927. Psyche 34: 113. 9.

texana Cresson. Que. to Fla., west to B. C. and Calif. Ecology: Nests in preexisting holes in

the ground. Parasite: Coelioxys moesta Cress., C. octodentata Say, C. rufitarsis Sm., C.
sodalis Cress. Pollen: Polylectic, visits a wide variety of flowers especially Leguminosae,
Compositae, and Labiatae, but also commonly takes pollen from Larrea tridentata
(Zygophyllaceae) in the southwestern U. S.; visitation records include Acacia greggii,
Acerates, Achillea lanulosa, Afzelia, Agave deserti, Althaea rosea, Alyssum
maritimum, Apocynum androsaemifolium, Amorpha fruticosa, Asclepias galioides, A.
mexicana, A. speciosa, Aster, Baccharis emoryi, Baptisia, Berberis, Blephilia, Buddleia,
Calycadenia multiglandulosa, Chamaecrista fasciculata, Chrysothamnus, Cirsium,
Clematis ligusticifolia, Clethra, Convolvulus, Cordylanthus nevinti, Coreopsis,
Crotalaria, Croton californicus, Cryptantha, Dianthera, Elephantopus carolinianus,
Eriastrum virgatum, Eriogonum elongatum, E. fasciculatum, E. latifolium var.
nudum, E. subscaposum, E. wrightii, Erigeron coulteri, Eupatorium, Ewphorbia,
Frasera parryi, Galactia, Gilia inconspicua, Gutierrezia sarothrae, Haplopappus,
Helenium, bigelovii, Helianthus, Hypericum, Hyptis emoryi, Kuhnistera oligophylla,’
Larrea tridentata, Liatris, Lotus nevadensis, L. scoparius, Lupinus longipes,
Marrubium vulgare, Medicago sativa, Melilotus alba, Monardella lanceolata, Nepeta,
Nolina, Opuntia, Oxydendron, Penstemon, Petalostemon candidum, Phacelia
heterophylla, Phaseolus, Phyla nodiflora, Pluchea camphorata, Polygala, Polygonum
aubertii, Psoralea, Pycnanthemum, Pyrrhopappus, Rhamnus californica, Rhus, Rubus,
Sapindus, Schinus molle, Serinea, Silphium, Solidago californica, S. confinis, Stachys,
Stanleya pinnata, Strophostyles, Tamarix, Tephrosia, Trifolium, Verbena, Vicia,
Viguiera laciniata.

Megachile tecana Cresson, 1878. Amer. Ent. Soc., Trans. 7: 125. d (? misdet.).

Megachile generosa Cresson, 1878. Amer. Ent. Soc., Trans. 7: 125. °.

Megachile schismatura Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 267. d.

Megachile cleomis Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 18. @, d.

Megachile pruinosa Friese, 1903. Ztschr. System. Hym. Dipt. 3: 246. 2, ¢. Preoce.

Megachile vernonensis Cockerell, 1912. Canad. Ent. 44: 354. “9” = 6.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 38 (nest, as generosa). —Hicks, 1926.
Colo. Univ. Studies 15: 228 (nest, as cleomis). —Krombein, 1953. Ent. Soc. Wash., Proc.
55: 84-85 (nest). —Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 30 (nest).
—Krombein, 1970. Ent. Soc. Wash., Proc. 72: 415 (nest). —Hurd and Linsley, 1975.
Smithsn. Contrib. Zool. 193: 37-38 (floral relationships).

Genus MEGACHILE Subgenus NEOMEGACHILE Mitchell

Megachile subg. Neomegachile Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 302, 307.
Type-species: Megachile chichimeca Cresson. Orig. desig.

Superfamily APOIDEA 2055

aegra Mitchell. South. Tex., south to Brazil.

Megachile aegra Mitchell, 1930. Amer. Ent. Soc., Trans. 56: 283. 3.
chichimeca Cresson. South. Tex.; Mexico.

Megachile chichimeca Cresson, 1878. Amer. Ent. Soc., Trans. 7: 130. 2.

Megachile disparipennis Cockerell, 1917. In W. P. Cockerell, N. Y. Ent. Soc., Jour. 25: 192.
OF

Genus MEGACHILE Subgenus CRESSONIELLA Mitchell

Megachile subg. Cressoniella Mitchell, 1943. Amer. Ent. Soc., Trans. 59: 302, 307.
Type-species: Megachile zapoteca Cresson. Orig. desig.
zapoteca Cresson. South. Ariz., south to Costa Rica. Pollen: Unknown, but visits flowers of
Asclepias, Helenium, Helianthus, Lathyrus, Melilotus, Monarda.
Megachile zapoteca Cresson, 1878. Amer. Ent. Soc., Trans. 7: 128. °.
Megachile tuxtla Cresson, 1878. Amer. Ent. Soc., Trans. 7: 128. é.

Genus MEGACHILE Subgenus MEGACHILE Latreille

Megachile Latreille, 1802. Hist. Nat. Formis, p. 413, 433.
Type-species: Apis centuncularis Linnaeus. Desig. by Curtis, 1828.
Anthophora Fabricius, 1804. Systema piezatorum, p. 372. Preoce.
Type-species: Apis centuncularis Linnaeus. Desig. by Michener, 1951.
Anthemois Robertson, 1903. Amer. Ent. Soc., Trans. 29: 168.
Type-species: Apis centuncularis Linnaeus. Monotypie and orig. desig. (=Megachile
infragilis Cresson).
Cyphopyga Robertson, 1903. Amer. Ent. Soc., Trans. 29: 169.
Type-species: Megachile montivaga Cresson. Monotypic and orig. desig.
Megalochila Schulz, 1906. Spolia Hym., p. 64. Emend.
centuncularis (Linnaeus). Holarctic, in America principally northern, but occurs south to Fla.
Mo., Colo., Nev., Ariz. Ecology: Nests both below and above ground in preexisting holes
or cavities. Parasite: Coeliorys modesta Sm., C. moesta Cress., C. octodentata Say,
Dibrachys sp., Melittobia chalybii, M. megachilidis (Pack.), Monodontomerus
montivagus Ashm., Ptinus sp. nr. hirtellus Sturm. Pollen: Apparently polylectiec, visits a
wide variety of flowers, especially Compositae and Leguminosae; visitation records
include Althaea rosea, Aster, Carduus undulatus, Centaurea cyanus, C. jacea,
Epilobium angustifolium, Fendlera, Gladiolus, Grindelia squarrosa, Helianthus
annuus, H. maximillianus, Inula heleniwm, Medicago sativa, Melilotus alba, M.
officinalis, Psoralea, Pyracantha, Sedum, Solidago, Taraxacum taraxacum, Trifolium,
Vernonia fasciculata, Zinnia.
Apis centuncularis Linnaeus, 1758. Syst. Nat., Ed. 10, p. 575. 2.
Apis rotundata Fabricius, 1787. Mantissa Insectorum, v. 1, p. 303. d.
Megachile infragilis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 127. 6.
Megachile leoni Titus, 1906. Ent. Soc. Wash., Proc. 7: 150. 2.
Perezia maura Ferton, 1914. Soc. Ent. France, Ann. 83: 233. Intersex.

Taxonomy: Hurd, 1967. Ent. Medd. 35: 5-6 (synonymy). —Pasteels, 1969. Soc. Ent. France,
Bul. 74: 248 (intersex, synonymy).

Biology: Gentry, 1874. Canad. Ent. 6: 171-175 (nest, supersedure). —Packard, 1874. Guide to
the study of insects, 4th ed., pp. 136-137 (nest). —Hicks, 1926. Colo. Univ. Studies 15: 231
(nest, as infragilis). —Michelbacher and Hurd, 1954. Pan-Pacific Ent. 20: 146 (nest,
parasite). —Medler, 1959. Canad. Ent. 91: 113-115, 1 fig. (nest). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 325-326 (life history, nest architecture, parasite).
—Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 20-21 (life history, parasite).
—Holm and Skou, 1972. Ent. Seand. 3: 169-180, 6 figs., 8 tables (life history, nest
architecture, sex ratio, rate of emergence, parasite, diseases).

inermis Provancher. N. S. to Ga., west to B. C., south to Calif., Utah., Colo., N. Mex. and Tex.
Ecology: Nests in decaying poplar and in borings in wood. Parasite: Coelioxys funeraria
Sm., Dibrachys maculipennis Szelenyi, Leucospis affinis Say. Pollen: Apparently

2056 Hymenoptera in America North of Mexico

polylectic, especially Compositae and Leguminosae; visitation records include Baptisia
tinctoria, Carduus undulatus, Inula helenium, Lathyrus venosus, Medicago sativa,
Nepeta cararia, Penstemon, Polymnia uvedalia, Rhododendron, Rubus strigosus, R.
villosus, Sanicula marylandica, Taraxacum officinale, Tephrosia virginiana, Vernonia.
Megachile simplex Provancher, 1882. Nat. Canad. 13: 229. 6. Preoce.
Megachile inermis Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 3238. 3.
Megachile simplicissima Dalla Torre, 1896. Cat. Hym., v. 10, p. 449. N. name.
Megachile sapellonis Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 7. 2.
Megachile temporalis Friese, 1903. Ztschr. System. Hym. Dipt. 3: 247. 3 (2 misdet.).
Megachile decipiens Lovell and Cockerell, 1907. Psyche 14: 19. 6.

Taxonomy: Titus, 1906. Ent. Soc. Wash., Proc. 7: 150 (synonymy).

Biology: Stephen, 1955. Econ. Ent., Jour. 48: 543 (effectiveness as pollinator of alfalfa).
—Stephen, 1956. Pan-Pacific Ent. 32: 98-101 (nest). —Koerber and Medler, 1958. Wis.
Acad. Sci., Arts and Letters 47: 56 (life history). —Medler, 1958. Canad. Ent. 90: 325-327, 1
fig. (life history). —Medler, 1958. Ent. News 66: 21 (parasite). —Fye, 1965. Canad. Ent. 97:
874-875, tables 2-4 (life history, nest architecture, sex ratio, parasite). —Medler and
Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 22-23 (life history, parasite).

montivaga Cresson. N. S. to N. C., west to B. C. and Calif.; Mexico. Ecology: Uses petals of
Clarkia for nest construction. Parasite: Coeliorys insita Cress., C. rufitarsis Sm.,
Leucospis affinis Say, Nemognatha dubia LeC., Phaenaera sp. Pollen: Polylectic,
especially Compositae, visits flowers of Argemone intermedia, A. platyceros, Asclepias,
Aster parishii, Blephilia, Brauneria, Campanula, Carduus undulatus, Centaurea jacea,
C. solstitialis, Chaenactis artemisiaefolia, Chilopsis linearis, Chrysopsis villosa,
Chrysothamnus, Cirsium californicum, C. lanceolatum, Clarkia amoena huntiana, C.
biloba, C. cylindrica, C. dudleyana, C. elegans, C. rubicunda, C. unguiculata, C.
viminea, C. williamsoni, Cleome, Convolvulus occidentalis, Coreopsis stellata,
Cryptantha intermedia, Dianthera, Echinacea, Epilobium, Euphorbia albomarginata,
Geranium maculatum, Gilia, Grindelia camporum, G. hallii, Gutierrezia, Haplopappus
bloomeri var. angustatus, Hedonia, Helenium bigelovii, Helianthus annuus, H.
gracilentus, Iris hartwegii, Lactuca, Lotus scoparius, Marrubium vulgare, Medicago
sativa, Melilotus, Monarda, Monardella, Oenothera, Opuntia, Palafoxia linearis,
Parthenium, Penstemon, Phacelia, Polymnia uvedalia, Rosa, Rudbeckia, Salix, Salvia
carduacea, S. mellifera, Scrophularia, Senecio douglasii, Sidalcea, Silphium, Solidago,
Trifolium, Verbena lasiostachys, Verbesina encelioides, Vernonia, Viguiera laciniata, V.
multiflora.

Megachile montivaga Cresson, 1878. Amer. Ent. Soc., Trans. 7: Zan Or

Biology: Hicks, 1926. Colo. Univ. Studies 15: 232 (nest).
nivalis Friese. Alaska, south to Oreg., east to Que. and Maine. Parasite: Anthrax irroratus
Say. Pollen: Unknown, but visits flowers of Elaeagnus, Taraxacum.
Megachile nivalis Friese, 1903. Ztschr. System. Hym. Dipt. 3: 246. 2 (d misdet.).
Megachile (Anthemois) santiamensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 311. @.

Taxonomy: Mitchell, 1942. Pan-Pacific Ent. 18: 115. d, °.
relativa Cresson. Mackenzie to Newfoundland, south to Calif., Ariz., Ill, Tenn., and Ga.

Ecology: Nests in borings in wood. Parasite: Coeliorys banksi Cwfd., C. funeraria Say,
C. modesta Sm., C. moesta Cress., C. porterae Ckll., Dibrachys maculipennis Szeleny),
Leucospis affinis Say, Melittobia chalybii Ashm., Pteromalus venustus Walker. Pollen:
Apparently polylectic, visits flowers of Amorpha, Apocynum, Aster punicus, A.
umbellatus, Baptisia, Besseya plantagineae, Brassica, Chamaenerion angustifolium,
Chrysanthemum leucanthemum, E pilobium, Erigeron coulteri, Frasera, Geranium,
Gilia, Houstonia, Iris, Melilotus, Mertensia, Phacelia, Physalis, Prenanthes,
Ranunculus, Rhodora, Rosa, Rubus, Rudbeckia hirta, Senecio subnudus, Solidago,
Trifolium, Valeriana.

Megachile relativa Cresson, 1878. Amer. Ent. Soc., Trans. 7: 126. °.

Megachile (Xanthosarus) exclamans Viereck, 1916. Conn. State Geol. and Nat. Hist.
Survey, Bul. 22: 743. °.

Megachile aspera Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 158. ?.

Superfamily APOIDEA 2057

Biology: Medler and Koerber, 1958. Ent. Soc. Amer., Ann. 51: 337-344, 3 figs. (life history,
parasite). —Koerber and Medler, 1958. Wis. Acad. Sci., Arts and Letters 47: 56 (life
history, parasite).

Genus MEGACHILE Subgenus EUTRICHARAEA Thomson

Megachile subg. Eutricharaea Thomson, 1872. Hym. Seand., v. 2, p. 228.
Type-species: Apis argentata Fabricius. Monotypic.
Megachile subg. Paramegachile Friese, 1898. Termesz. Fus. 21: 198.
Type-species: Apis argentata Fabricius. Desig. by Mitchell, 1934.
Megachile subg. Paramegalochila Schulz, 1906. Spolia Hym., p. 71. Emend.
Androgynella Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 7: 313.
Type-species: Megachile detersa Cockerell. Monotypic and orig. desig.
Megachile subg. Neoeutricharaea Rebmann, 1967. Ent. Ztschr. (N. S.) 17: 36.
Type-species: Megachile rotundata (F.). Orig. desig. (name of type-species placed
originally in quotation marks).

Taxonomy: Rebmann, 1967. Ent. Ztschr. (n. s.) 77: 33-38. —Hurd, 1967. Ent. Medd. 35: 3-10.
—Rebmann, 1967. Deut. Ent. Ztschr. (n. s.) 15: 21-48. —Rebmann, 1967. Ent. Ztschr. 77:
169-171.

Biology: Parker, Torchio, Nye and Pedersen, 1976. Jour. Apicult. Res. 15: 89-92 (field-cage
studies).
apicalis Spinola. Canada, N. J., Va. Presumably introduced from Europe. Pollen: Unknown, but

visits flowers of Centaurea cyanus.

Megachile apicalis Spinola, 1808. Insectorum Liguriae, v. 2, p. 259. 2.

Megachile mixta Costa, 1863. Accad. delle Sci. Fis. e Mat. Napoli, Atti 1 (2): 44. 2.

Megachile dimidiati-ventris Dours, 1873. Rev. Mag. Zool. (3) 1: 300. 2.

Megachile virginiana Mitchell, 1926. Amer. Ent. Soc., Trans. 52: 113. 9.

concinna Smith. Holarctic; Pa. and Ohio south to Fla. and Ala., Kans., Okla., Ariz., Nev., Calif.,

Utah, Wash.; Mexico. Ecology: Nests in borings in wood. Presumably introduced from
West Indies after World War II and was probably introduced from Africa into the West
Indies during the early part of the nineteenth century. Parasite: Anthrax cintalapa
Cole, Coelioxys moesta Cress., Nemognatha lurida LeC., Tetrastichus megachilidis
Burks. Pollen: Polylectic, visits a variety of native and introduced flowers including
Acacia, Asclepias, Aster, Baccharis, Bidens, Centromadia pungens, Citrus, Croton
californicus, Euphorbia albomarginata, Heliotropium curassavicum, Hemizonia
pungens, Ipomoea, Lepidium, Lippia, Lotus purshianus, Medicago sativa, Melilotus
alba, M. indica, Onobrychis vicaefolia, Polygonum aubertii, Prosopis, Raphanus
sativus, Senecio, Sicyos, Tamarix, Trifolium repens, Vernonia, Wislizenia refracta.
Predator: Trogoderma sp.

Megachile concinna Smith, 1879. Deser. n. spp. Hym., p. 79. °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 121-122, fig. 39
(redescription, other possible synonyms).

Biology: Butler and Wargo, 1963. Pan-Pacifie Ent. 39: 201-206, 1 fig. (life history, nest
architecture, parasite, predator). —Butler and Ritchie, 1965. Pan-Pacifie Ent. 41: 153-157
(life history, pollination). —Krombein, 1967. Trap-nesting wasps and bees, pp. 326-327 (life
history, nest architecture, predator).

pacifica (Panzer). Holarctic; Mass. to Va., west to B. C., Wash., Oreg. and Calif. Ecology:
Nests in borings in wood as well as in a wide variety of native and artificial burrows.
Presumably introduced some time during or just after World War IJ; also adventive in
S. Amer. (Chile and Argentina). Parasite: Coeliorys funeraria Sm., C. gilensis Ckll., C.
moesta Cress., C. novomexicana CkIl., C. octodentata Say, C. sodalis Cress., Dibrachys
maculipennis Szelenyi, Melittobia acasta Walker, M. hawatiensis Perkins,
Monodontomerus montivagus Ashm., M. obscurus Westw., Sapyga pumila Cress.,
Tetrastichus albipes Crosby. Pollen: Polylectic, visits a wide variety of both native and
introduced flowers and is an exceptionally valuable pollinator of alfalfa; visitation
records include Asclepias, Centromadia pungens, Cichorium intybus, Cosmos,

2058 Hymenoptera in America North of Mexico

Euphorbia albomarginata, Heliotropium curassavicum var. oculatum, Lotus
corniculatus, Medicago sativa, Melilotus alba, Phacelia ramosissima, Polygonwm
aubertii, Senecio douglasii, Solidago, Veronica. The name adopted by the International
Bee Research Association for this species is Megachile pacifica (Panzer); however, an
application pending before the International Commission on Zoological Nomenclature
requests that the Commission use its plenary powers to set aside all type selections for
Apis rotundata Fabricius, 1793, and to rule that the species is to be interpreted by
reference to a newly established neotype specimen. Thus, if the Commission rules
affirmatively on this application, the correct name for this species will be Megachile
rotundata (Fabricius) which is currently considered to be a synonym of Megachile
centuncularis (Linnaeus) and is so treated in this catalog.

Apis pacifica Panzer, 1798. Faunae Ins. German., v. 55, p. 16. @.

Megachile imbecilla Gerstaecker, 1869. Stettin. Ent. Ztg. 30: 359. 2.

Taxonomy: Krombein, 1948. Ent. Soc. Wash., Proc. 50: 14 (as rotundata). —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 122-124, figs. 38-39 (redescription, possible identity, as
rotundata?). —Hurd, 1967. Ent. Medd. 35: 3-10 (tax. status of rotundata Fabr.).
—Rebmann, 1967. Ent. Ztschr. 77: 169-171 (tax. status of rotundata Fabr. and pacifica
Panz.). —Holm and Skou, 1972. Ent. Scand. 3: 169-170 (review of tax. status of rotundata
Fabr. and pacifica Panz.). —Roberts, 1974. Zool. Nomencl., Bul. 30: 190-192 (proposed
suppression of lectotype and designation of neotype in accord with Megachile rotundata
auct.). —Anonymous, 1975. Bee World 56: 164 (identity problems, adoption by
International Bee Research Association of Megachile pacifica (Panzer) as correct name for
this species).

Biology: Daly, 1952. Ent. News 63: 210-211 (geogr. and floral records in midwest. U. S.).
—Hurd, 1954. Ent. News 65: 93-95 (geogr. and floral records in Calif.). —Stephen and
Torchio, 1961. Pan-Pacific Ent. 37: 85-93 (nest sites, emergence, cells, adult behavior,
leaf-cuttings, larval development, historical notes). —Stephen, 1961. Econ. Ent., Jour. 54:
989-998 (artificial nest sites and propagation for alfalfa pollination). —Bohart, 1962. Utah
Agr. Expt. Sta., Cire. 144: 1. (management for alfalfa pollination). —Stephen, 1962. Oreg.
Agr. Expt. Sta., Bul. 586, 16 pp., 9 figs. (propagation for alfalfa seed production).
—Johansen, Jaycox and Hutt, 1963. Wash. Agr. Expt. Sta., Cire. 418:1. (effect of
pesticides). —Torchio, 1963. Utah Farm Home Sci. 24: 70-71 (parasite). —Nye and Bohart,
1964. Utah State Univ. Agr. Expt. Sta., Cire. 145: 1-7, 6 figs. (equipment for making
nesting holes). —Stephen, 1965. XII. Int. Congr. Ent. London, Proc. p. 350 (circadian
rhythms). —Stephen and Osgood, 1965. Econ. Ent., Jour. 58: 284-286, 1 table (induction of
emergence). —Bacon, Barton, MacSwain, Marble and Stanger, 1965. Calif. Univ. Agr. Ext.
Serv., AXT 160: 1-13 (management for alfalfa pollination). —Hobbs, 1965. Canada Dept.
Agr., Pub. 1209: 1-11, 5 figs. (importation and management for alfalfa pollination).
—Stephen and Osgood, 1965. Econ. Ent. Jour. 58: 965-968 (effect of tunnel size on sex
ratio). —Johansen and Eves, 1966. Wash. State Univ. Agr. Expt. Sta., Cire. 469: 1-12
(parasites and nest destroyers). —Hobbs, 1967. Canada Dept. Agr., Pub. 1313: 1-19
(domestication). —Krombein, 1967. Trap-nesting wasps and bees, pp. 327-329 (life history,
nest architecture). —Bohart and Knowlton, 1967. Utah. Ext. Serv., EL 104 (rev): 1-7, 1 pl.
(management for higher alfalfa seed yields). —Medler and Lussenhop, 1968. Wis. Univ.
Res. Bul. 274: 29-30 (life history, parasites). —Waters, 1968. Idaho Agr. Expt. Sta.,
Current Information Series 97: 1-4 (beetle excluder trap). — Williams, 1968. Idaho Coll.
Forestry, Wildlife and Range Sci., Station Note 10: 1-4 (nesting boards). —Hobbs, 1968.
Canad. Ent. 100: 781-784 (control of insect enemies). —Johansen and Eves, 1969. Wash.
Agr. Ext. Series, EM 2631 (rev.): 1-10 (control of natural enemies). —Klostermeyer and
Gerber, 1969. Ent. Soc. Amer., Ann. 62: 1321-1325 (nesting behavior monitored by an event
recorder). —Eves, 1970. Melanderia 4: 1-18, 7 figs., 11 tables (parasite). —Torchio, 1970.
Ark. Agr. Ext. Serv., Mis. Publ. 127: 84-90 (parasite). —Johansen and Eves, 1971. Wash.
Agr. Ext. Serv., EM 2631 (rev.): 1-10 (control of natural enemies). —Waters, 1971. Idaho
Univ. Agr. Expt. Sta., Current Information Ser. 163: 1-4 (insect enemies and their control).
—Hobbs and Krunic, 1971. Canad. Ent. 103: 674-685 (parasites). —Holm and Skou, 1972.
Ent. Scand. 3: 169-180, 6 figs., 8 tables (life history, nest architecture, sex ratio). —Torchio,
1972. Melanderia 10: 1-22, 55 figs., 1 table (Sapyga pumila Cress., a parasite). —Torchio,

Superfamily APOIDEA 2059

1972. Melanderia 10: 23-30, 7 figs. (control of Sapyga pumila Cress.). —Telford, Johansen
and Eves, 1972. Mededel. Fakult. Landbouwwetensch. Gent 37: 776-783 (management
practices and insecticide poisoning). —Szabo and Smith, 1972. Apiculture Res., Jour. 11:
157-165 (influence of light and temperature on adult activity). —Hobbs, 1972. Bee World
53: 167-173 (beekeeping with alfalfa leafeutter bees in Canada). —Thorp and Briggs, 1972.
Environ. Ent. 1: 399-401 (mortality of immatures in relation to alfalfa saponins).
—Klostermeyer, Mech and Rasmussen, 1973. Kans. Ent. Soc., Jour. 46: 536-548 (sex and
weight of progeny with provision weights). —Santis, 1973. Cienc. e Abejas 2: 15-19
(parasites). —Hobbs, 1973. Canada Dept. Agr., Publ. 1495: 1-30 (use in alfalfa pollination).
—Torchio, 1974. Utah State Univ. Agr. Expt. Sta., UMC 48, Res. Rept. 16: 1-13, 18 figs.
(biology and control of Sapyga pumila Cress., a parasite). —Parker and Pedersen, 1975.
Environ. Ent. 4: 103-104 (effect of alfalfa saponins on larval mortality). —Tasei, 1975.
Apidologie 6: 1-57 (adaptation of N. Amer. population introduced into France). — Davis,
Johansen, and Eves, 1975. Apiculture Research, Jour. 14: 101-104, table (synthetic
attractants). —Stephen and Undurraga, 1976. Jour. Apicult. Res. 15: 81-87, 4 figs.
(x-radiography, an analytical tool in population studies). —Hobbs and Richards, 1976.
Canad. Ent. 108: 165-167 (selection for univoltine strain). —Batra, 1976. Kans. Ent. Soc.,
Jour. 49: 18-22, tables 1-2 (comparative efficiency in alfalfa pollination).

Morphology: Gerber and Akre, 1969. Melanderia 1: 1-36 (external). —Wachmann, Richter and
Schricker, 1973. Ztschr. f. Morph. Tiere 76: 109-128 (fine structure of compound eye).

Genus MEGACHILE Subgenus DELOMEGACHILE Viereck

Megachile subg. Delomegachile Viereck, 1916. Conn. State Geol. and Nat. Hist. Survey,
Bul. 22: 745.
Type-species: Megachile frigida Smith. Monotypic. (=Megachile vidua Smith).
addenda Cresson. Que., Ont. and N. H. to Fla., west to Mich., Kans., Tex., and Calif. Pollen:
Apparently polylectic, visits flowers of Amorpha canescens, Asclepias, Baptisia
tinctoria, Coreopsis, Dianthera, Encelia californica, Gillenia, Hieracium, Hypericum,
Oenothera, Opuntia vulgaris, Penstemon, Polycodium, Psoralea floribunda, Rosa,
Rubus, Tephrosia virginiana, Vaccinium.
Megachile addenda Cresson, 1878. Amer. Ent. Soc., Trans. 7: 124. 2, ¢.
Megachile manumuskin Viereck, 1902. Canad. Ent. 34: 328. 2, d.

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc. 3: 160-162 (nest).
frigida appalachensis Mitchell. N. Y. to Ga. in the Appalachian Mts. Pollen: Unknown, but
visits flowers of Apocynum, Baptisia, Coreopsis, Galax, Koellia, Penstemon,
Pycnanthemum, Rhododendron catawabiense, Vernonia glauca.
Megachile (Delomegachile) vidua var. appalachensis Mitchell, 1935. Amer. Ent. Soc.,
Trans. 61: 205. 2, 6.
frigida frigida Smith. Alaska and Canada, south to Pa., Mich., and Nebr. and in the mts. to N.
Mex., Ariz. and cent. Calif. Ecology: Nesting in decaying poplar and also in borings in
wood. Parasite: Coelioxys funeraria Sm., C. moesta Cress., C. porterae Ckll., C. sodalis
Ckll. Pollen: Unknown, but visits flowers of Apocynum, Astragalus, Campanula,
Castilleia, Chamaenerion angustifolium, Epilobium, Iris, Malvastrum, Marrubium
vulgare, Medicago sativa, Melilotus, Mimulus, Monarda, Penstemon, Phacelia,
Polemonium, Robinia, Rosa, Symphoricarpos, Trifolium, Verbena, Vicia.
Megachile frigida Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 193. d.
Megachile monardarum Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 11. °.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 338. —Cockerell, 1905. Amer. Ent.
Soe., Trans. 31: 337. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 133-134, figs.
43-45 (redescription).

Biology: Stephen, 1956. Pan-Pacific Ent. 32: 95-98 (life history). —Medler and Lussenhop,
1968. Wis. Univ. Res. Bul. 274: 21-22 (life history, parasite).

gemula cressonii Dalla Torre. Wash., Nev., Ariz.
Megachile carbonaria Cresson, 1879. Amer. Ent. Soc., Trans. 7: 208. 2. Preoce.
Megachile cressonii Dalla Torre, 1896. Cat. Hym., v. 10, p. 427. N. name.
Megachile vandykei Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 205. °.

2060 Hymenoptera in America North of Mexico

gemula gemula Cresson. MacKenzie to N. S., south to Ga., west to IIl., Nebr., N. Mex., and

Calif. north to B. C. and N. W. T., principally montane in south. part of range. Pollen:
Unknown, but visits flowers of Apocynum androsaemifolium, Asclepias speciosa,
Baptisia tinctoria, Campanula rotundifolia, Chrysanthemum leucanthemum, Clematis,
Coreopsis, Geranium maculata, Gerardia, Hydrangea, Philadelphus, Pycnanthemum,
Rhododendron catawabiense, Rubus, Rudbeckia, Solidago lanceolata, Trifolium,
Vaccinium, Vicia.

Megachile gemula Cresson, 1878. Amer. Ent. Soc., Trans. 7: 118. ¢ (2 misdet.).

Megachile avaria Cresson, 1878. Amer. Ent. Soc., Trans. 7: 123. 3.

Megachile Vancouveriensis Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p.
424. 6.

Megachile albula Lovell and Cockerell, 1907. Psyche 14: 18. 6.

Megachile (Delomegachile) gemula var. fulvogemula Mitchell, 1936. Amer. Ent. Soc.,
Trans. 61: 185. 9.

Taxonomy: Titus, 1906. Ent. Soc. Wash., Proc. 7: 151 (synonymy).

Biology: Fye, 1965. Canad. Ent. 97: 876-877, fig. 6, tables 2-4 (nest). —Medler and Lussenhop,
1968. Wis. Univ. Res. Bul. 274: 22 (nest).
giliae Cockerell. Alaska, N. W. T., B. C., Alta., Colo. Pollen: Unknown, but visits flowers of
Gilia.
Megachile giliae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 452. ¢.
ingenua Cresson. Pa. to Fla., west to III. Pollen: Possibly oligolectie on Tephrosia including T.
virginiana, but also visits flowers of Lupinus and Vaccinium presumably for nectar.
Megachile ingenua Cresson, 1878. Amer. Ent. Soc., Trans. 7: 122. d.
Megachile tephrosiana Mitchell, 1927. Psyche 34: 179. °.
melanophaea calogaster Cockerell. B. C. and Idaho to mts. of cent. Calif. Pollen: Apparently
polylectic, visits flowers of Astragalus, Hackelia jessicae, Haplopappus aparagoides,
Horkelia fusca, Lotus oblongifolius, Lupinus lyallii lyallii, L. superbus, Mimulus
moschatus, M. tillingi, Pedicularis frigida, P. groenlandica, Phyllodoce breweri,
Polemonium occidentale, Streptanthus tortuosus.
Megachile calogaster Cockerell, 1898. Acad. Nat. Sci. Phila., Proc. 50: 55. ¢, 2.
melanophaea melanophaea Smith. MacKenzie to Newfoundland, south to Calif., Ariz., Colo.,
Penn. and Ga.? Parasite: Coeliorys rufitarsis Sm., C. sodalis Cress. Pollen: Apparently
polylectic, visits a wide variety of flowers including Agastache occidentatis, Apocynum,
Astragalus bisulcatus, Azalea, Campanula rotundifolia, Chamaenerion angustifolium,
Cypripedium reginae, Epibolium angustifolium, Helianthus, Hemilobus tenellus,
Lupinus argenteus, L. rootkatensis, Medicago sativa, Mimulus, Nolina, Phacelia,
Psoralea argophylla, Ranunculus, Raphanus, Rhodora, Robinia, Rosa, Rubus,
Rudbeckia, Solidago, Symphoricarpos occidentalis, Sisymbrium, Taraxacum, Vicia
eracea.
Megachile melanophaea Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 191. 2, ¢.
Megachile canadensis Friese, 1903. Ztschr. System. Hym. Dipt. 3: 248.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 336 (synonymy).

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 162-163 (life history, parasite).
melanophaea rohweri Cockerell. Colo., Utah, Ariz., N. Mex. Ecology: Uses pieces of
Symphoricarpos leaves for nesting material. Pollen: Presumably polylectic, visits
flowers of Cirsium, Erysimum, Lupinus, Nolina, Robinia, Sisymbrium, Trifolium.
Megachile wootoni rohweri Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 453. 9.
Megachile pseudolatimanus Strand, 1917. Arch. f. Naturgesch. 83 (abt. A, h. 11): 65. ¢.
Megachile tuala Strand, 1917. Arch. f. Naturgesch. 83 (abt. A, h. 11): 66. 9.
melanophaea submelanophaea Mitchell. South. Calif., mts., Maine, Mich. Pollen: Presumably
polylectic, visits flowers of Dicentra chrysantha, Lupinus.
Megachile (Delomegachile) melanophaea var. submelanophaea Mitchell, 1935. Amer. Ent.
Soc., Trans. 61: 197. °.

Superfamily APOIDEA 2061

melanophaea wootoni Cockerell. Mont., Nebr., Colo., N. Mex. Parasite: Coeliorys sodalis
Cress., C. rufitarsis Sm. Pollen: Presumably polylectic, visits flowers of Astragalus,
Medicago sativa.
Megachile wootoni Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 1: 125. 6.
mucida Cresson. N. J., N. C., Ga., Tex. Pollen: Unknown, but visits flowers of Lupinus,
Polycodium, Rubus, Tephrosia virginiana.
Megachile mucida Cresson, 1878. Amer. Ent. Soc., Trans. 7: 118. ¢ (2 misdet.).
Megachile mucida semimucida Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 26. 2, d.
Megachile audax Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 161. ¢.

Genus MEGACHILE Subgenus PHAENOSARUS Mitchell

Megachile subg. Phaenosarus Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 303, 309.
Type-species: Megachile fortis Cresson. Orig. desig.
agustini Cockerell. Colo., N. Mex., Utah, Ariz. Pollen: Unknown, but visits flowers of Baileya,
Helianthus, Heterotheca, Verbesina, Viguiera, Zexmenia.
Megachile agustini Cockerell, 1905. Ent. News 16: 82. ¢.
Megachile (Phaenosarus) subfortis Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 145. 3.
fortis Cresson. Ill., lowa, Wis., S. Dak., Nebr., Kans., La., Tex., Colo., N. Mex., Ariz. Parasite:
Coelioxys rufitarsis Sm. Pollen: Unknown, but visits flowers of Dicrophyllum
marginatum, Helianthus peliolaris, H. subrhomboideus, Silphiuwm integrifolium,
Solidago canadensis, Vernonia fasciculata.
Megachile fortis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 262. d.
Megachile emoryi Cockerell, 1904. Entomologist 37: 7. 2.
Megachile fortis var. vestali Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 530. ¢.

Biology: Hicks, 1926. Colo. Univ. Studies 15: 229 (nest, parasite).

Morphology: Fischer, 1957 (1956). Canad. Ent. 88: 657-673, 9 figs. (musculature of male
metasoma and genitalia).

Genus MEGACHILE Subgenus MEGACHILOIDES Mitchell

Megachiloides Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 154.
Type-species: Megachiloides oenotherae Mitchell. Monotypic.

The bees of this subgenus appear to be specialists of the Onagraceae, collecting pollen espe-
cially from the large flowered species of the genus Oenothera. Apparently the species also
differ in that some excavate their own nesting burrows while at least one species appropriates
preexisting tunnels of other bees.

Taxonomy: Bohart and Youssef, 1972. Royal Ent. Soc. London, Trans. 124: 18 (biol.
characters).
amica Cresson. Kans., Tex. Pollen: Unknown, but visits flowers of Anogra pallida, Monarda
punctata, Opuntia, Verbesina encelioides.
Megachile amica Cresson, 1872. Amer. Ent. Soc., Trans. 4: 265. d.
oenotherae (Mitchell). N. J., N. C., Okla., Tex. Ecology: Nests in preexisting burrows of
Andrena macra Mitchell in sandy loam soil, using cut leaves for cells and caps them
with cut piecies of petals from Oenothera laciniata. Parasite: Coelioxys piercei Cwfd.?.
Pollen: Collects pollen from the flowers of Oenothera laciniata, but also visits flowers of
Ceanothus americanus, Meriolex drummondiana, Penstemon.
Megachiloides oenotherae Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 154. °, ¢.

Biology: Sivik, 1954. Ent. News 65: 256 (nest, parasite).
umatillensis (Mitchell). Wash., Utah, Colo. Ecology: Excavates nesting burrows in partially
stabilized sand dunes, uses Oenothera petals for cell walls and folds cell walls over at
top to cap the cell. Parasite: Coeliorys mesae Ckll., Nemognatha lutea LeC. Pollen:
Collects pollen from flowers of Oenothera pallida.
Megachiloides wmatillensis Mitchell, 1927. Psyche 34: 118. d, °.

Biology: Bohart and Youssef, 1972. Royal Ent. Soe. London, Trans. 124: 1-19, 23 figs. (life
history, foraging behavior, nest architecture, parasites).

2062 Hymenoptera in America North of Mexico

Genus MEGACHILE Subgenus DEROTROPIS Mitchell

Megachile subg. Derotropis Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 119, 156.
Type-species: Megachile pascoensis Mitchell. Orig. desig.

Taxonomy: Mitchell, 1944. Pan-Pacific Ent. 20: 142-144 (Key to spp.).

alamosana Mitchell. Oreg., Wyo., Colo.
Megachile (Xeromegachile) alamosana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 329. ¢.
anograe Cockerell. Mont., Wyo., Nebr., Colo., Kans., Tex., Oreg., Calif. Ecology: Excavates
nesting burrows in sandy soil. Pollen: Collects pollen from flowers of Opuntia and
Oenothera albicaulis, but visits other flowers presumably for nectar including Anogra
coronopifolia, Astragalus.
Megachile anograe Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 261. 9.
astragali Mitchell. Calif. (Mojave Desert). Pollen: Unknown, but visits flowers of Astragalus
lentiginosus var. fremontii.
Megachile (Derotropis) astragali Mitchell, 1938. Pan-Pacific Ent. 14: 174. 5, @.
gravita Mitchell. Wash., Oreg., Calif. Pollen: Collects pollen from flowers of Clarkia including
C. amoena amoena, C. amoena huntiana, C. cylindrica, C. dudleyana, C. gracilis
albicaulis, C. purpurea quadrivulnera, C. rubicunda, C. speciosa nitens, C. speciosa
polyantha, C. speciosa speciosa, C. unguiculata, C. viminea, C. williamsonii, but visits
other flowers for nectar including Chaenactis glabriuscula, Malacothamnus densiflorus.
Megachile (Xeromegachile) gravita Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 322. 3.
Megachile (Xeromegachile) astata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 345. 9°.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 20-23 (floral
relationships).
laurita Mitchell. Wash., Oreg., Utah. Pollen: Collects pollen from flowers of Oenothera pallida,

O. trichocalyx.
Megachile laurita Mitchell, 1927. Psyche 34: 115. 9.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 41 (floral
relationships).
melanderi Mitchell. Tex., Calif. (Inyo Co.).
Megachile (Derotropis) melanderi Mitchell, 1944. Pan-Pacific Ent. 20: 140. °.
pascoensis Mitchell. Mont., Idaho, Wash., Oreg., Calif. Parasite: Lines cells with pieces of
petals cut from flowers of Clarkia cylindrica and C. speciosa. Pollen: Oligolege of
Clarkia including C. amoena, C. biloba, C. bottae, C. cylindrica, C. dudleyana, C.
elegans, C. gracilis albicaulis, C. pulchella, C. purpurea, C. rhomboidea, C. rubicunda,
C. speciosa, C. unguiculata, C. xantiana, but also visits other flowers for nectar
including Chamaebatia foliolosa, Convolvulus villosus, Cordylanthus pilosus, Sidalcea
malvaeflora, Trifolium, Wyethia angustifolia.
Megachile (Xeromegachile) bakeri Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 316. 3.
Preoce.
Megachile (Xeromegachile) pascoensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 320. 3.
Megachile (Xeromegachile) gabrielensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 346. °.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 23-26 (floral
relationships).

semilaurita Mitchell. Utah.
Megachile laurita semilaurita Mitchell, 1927. Psyche 34: 116. °.

subanograe Mitchell. Tex., Wyo., Utah, Nev., east. Calif. Pollen: Unknown, but visits flowers
of Sphaeralcea ambigua.
Megachile (Megachiloides) subanograe Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 344. 2.
sublaurita Mitchell. Utah.
Megachile laurita sublaurita Mitchell, 1927. Psyche 34: 117. °.

xerophila Cockerell. South. Calif., deserts., Ariz. Pollen: Unknown, but visits flowers of
Baileya, Chaenactis stevioides, Encelia, Geraea canescens, Helianthus niveus, Larrea

Superfamily APOIDEA = 2068

tridentata, Melilotus, Oenothera deltoides, Palafoxia linearis, Sphaeralcea ambigua, S.
emoryi, S. rosacea.
Megachile xerophila Cockerell, 1933. Pan-Pacific Ent. 9: 27. °.
yumensis Mitchell. Ariz. (Wickenburg).
Megachile (Derotropis) ywmensis Mitchell, 1944. Pan-Pacifie Ent. 20: 141. 2.

Genus MEGACHILE Subgenus XEROMEGACHILE Mitchell

Megachile subg. Xeromegachile Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 302, 309.
Type-species: Megachile integra Cresson. Orig. desig.
alata Mitchell. South. and east. Calif., Ariz.; Mexico (Baja California). Pollen: Unknown, but
visits principally flowers of the Compositae including Bebbia juncea, Chrysothamnus,
Encelia californica, E. farinosa, Eschscholzia, Helianthus gracilentus, Prosopis,
Viguiera parishii.
Megachile (Xeromegachile) alata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 324. d.
Megachile (Xeromegachile) redlandica Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 331. 2.
boharti Mitchell. South. Calif., Ariz.; Mexico (Sonora). Pollen: Unknown, but visits flowers of
Cercidium, Prosopis.
Megachile (Xeromegachile) boharti Mitchell, 1942. Pan-Pacific Ent. 18: 74. °.
bradleyi Mitchell. Utah, Nev., Calif. (Antioch).
Megachile (Xeromegachile) bradleyi Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 340. 2.
brimleyi Mitchell. N. C., Fla. Pollen: Unknown, visits principally flowers of Galactia, but has
also been found at flowers of Erigeron volubilis.
Megachile brimleyi Mitchell, 1926. Amer. Ent. Soc., Trans. 52: 114. ¢.
bruneri Mitchell. Colo., east. Calif.; Mexico (Baja California). Pollen: Unknown, but visits
flowers of Chrysothamnus.
Megachile (Xeromegachile) bruneri Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 328. d.
casadae Cockerell. S. Dak., Nebr., Okla., Tex., Wyo., Colo., N. Mex., Utah, Calif., Ariz. Pollen:
Unknown, but visits flowers of Helianthus, Opuntia, Populus.
Megachile casadae Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 1: 127. 6.
Megachile populi Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 17. 2.
Megachile opuntiarwm Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 229. 2.
Megachile austinensis Mitchell, 1927. Psyche 34: 105. °.
coloradensis Mitchell. Colo.
Megachile (Xeromegachile) coloradensis Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 379. °.
dakotensis Mitchell. Minn., Iowa, IIl., Mont., N. Dak., S. Dak., Nebr., Tex. Pollen: Unknown, but
visits flowers of Amorpha canescens, Cleome serrulata, Helianthus, Kuhnistera
candida, K. oligophylla, Petalostemon violaceus, Verbena.
Megachile dakotensis Mitchell, 1926. Psyche 33: 164. d, °.
deflexa Cresson. N. C., Fla., Nebr., Kans., Okla., Tex. Pollen: Unknown, but visits flowers of
Bidens, Clethra, Coreopsis, Erigeron, Gaillardia amblydon, Geobanus pallidus,
Petalostemon, Phaseolus, Rubus, Strophostyles.
Megachile deflexa Cresson, 1878. Amer. Ent. Soc., Trans. 7: 125. d.
Megachile megagyna Cockerell, 1909. Canad. Ent. 41: 394. °.
Megachile indianorum Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 533. d.
Megachile (Xeromegachile) coreopsana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 319. d.
dulciana Mitchell. South. Calif.
Megachile (Xeromegachile) dulciana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 343. °.
fucata Mitchell. Colo., N. Mex., Utah, Ariz., Calif., deserts. Pollen: Possibly polylectic, is known
to collect pollen from Larrea tridentata, but visits other flowers including Argemone
platyceros, Aster abatus, A. tortifolius, Baileya, Cercidium, Encelia frutescens,
Palafoxia linearis, Prosopis, Senecio douglasti var. longilobus.
Megachile (Xeromegachile) fucata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 323. d.
Megachile (Xeromegachile) histrata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 334. °.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 38 (floral relationships).

2064 Hymenoptera in America North of Mexico

hilata Mitchell. Colo., Utah, east. Calif.
Megachile (Xeromegachile) hilata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 332. °.

hookeri Cockerell. Colo., Utah.
Megachile hookeri Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 533. °.
impartita Mitchell. Colo., east. Calif.
Megachile (Xeromegachile) impartita Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 333. °.
instita Mitchell. N. Mex., Calif:, Ariz.; north. Mexico. Pollen: Possibly an oligolege of the
Leguminosae, visits flowers of Acacia, Cercidium, Cirsium, Prosopis.

Megachile (Xeromegachile) instita Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 326. ¢.
integra Cresson. N. J. to Fla., west to Ill, Kans., and Tex. Pollen: Unknown, but visits flowers
of Erigeron, Galactia, Glycina, Koellia, Phaseolus, Pycnanthemum, Strophostyles

umobellata.
Megachile integra Cresson, 1878. Amer. Ent. Soc., Trans. 7: 121. ¢.
Megachile strophostylis Robertson, 1904. Canad. Ent. 36: 277. °.
integrella Mitchell. N. C., Fla. Pollen: Unknown, but visits flowers of Hypericum,
Oxydendrum.
Megachile integrella Mitchell, 1926. Amer. Ent. Soc., Trans. 52: 116. @.
inyoensis Mitchell. Calif., Ariz. Pollen: Unknown, but visits flowers of Encelia.
Megachile (Xeromegachile) inyoensis Mitchell, 1942. Pan-Pacific Ent. 18: 117. 2.
laguniana Mitchell. South. Calif.
Megachile (Xeromegachile) laguniana Mitchell, 1937. Amer. Ent. Soc., Trans. 63: 419. °.
latita Mitchell. Wyo., Colo.
Megachile (Xeromegachile) latita Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 335. °.
legalis Cresson. Colo., Nev., Ariz., Wash., Oreg., east. Calif. Pollen: Unknown, but visits
flowers of Cirsium, Encelia farinosa, Iris, Senecio.
Megachile legalis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 209. 6.
Megachile (Xeromegachile) couleeana Mitchell, 1938. Pan-Pacific Ent. 14: 171. 9.
macneilli Mitchell. Calif. (Riverside Co.).

Megachile (Xeromegachile) macneilli Mitchell, 1957. Pan-Pacifie Ent. 33: 24. d.
manifesta Cresson. Alta., Mont., N. Dak. to N. Mex., Ariz., south. Calif. Pollen: Unknown, but
visits flowers of Aster, Chrysothamnus nauseosus speciosus, Cleome, Grindelia

squarrosa, G. sarothrae, Haplopappus, Helianthus, Viguiera.
Megachile manifesta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 122. 6.
Megachile chrysothamni Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 262. 9.
maurata Mitchell. Wyo., east. Calif., Ariz. Pollen: Unknown, but visits flowers of Encelia,
Sphaeralcea. This is probably the male of M. (Derotropis) subanograe Mitchell.
Megachile (Xeromegachile) maurata Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 369. ¢.
micheneri Mitchell. Colo., Calif. Pollen: Unknown, but visits flowers of Vigwiera multiflora.
Megachile (Xeromegachile) micheneri Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 3738. ¢.
mojavensis Mitchell. South. and east. Calif., Ariz. Pollen: Unknown, but visits flowers of Aster,
Chaenactis, Encelia, Phacelia.
Megachile (Xeromegachile) mojavensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 340. °.
mucorosa Cockerell. Nebr., Kans., Tex., Wyo., Colo., N. Mex., Utah, Ariz. Pollen: Unknown, but
visits flowers of Eustoma issenianum, Helianthus, Monarda, Ratibida columnaris.
Megachile mucorosa Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 265. d.
Megachile (Argyropile) nebraskana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 347. °.
nelsoni Mitchell. Colo.
Megachile (Xeromegachile) nelsoni Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 381. °.
nevadensis Cresson. Mont. and Wyo. to N. Mex., west to Wash. and Calif., La. (Winnfield).
Pollen: Apparently an oligolege of autumnal flowering Compositae, including Chrysopsis
villosa, Chrysothamnus bernardina, C. nauseosus, C. n. consimilis, C. n. mohavensis, C.
n. speciosus, C. pumilis, C. viridulus, C. viscidiflorus typicus, Grindelia camporum,
Haplopappus acradenius, H. bloomeri var. angustatus, H. linearifolius, H. vernonioides,
Helianthus, Heterotheca grandiflora, Pluchea camphorata, Senecio douglasti, Solidago

Superfamily APOIDEA 2065

californica, S. confinis, but also visits flowers of Cleome and Melilotus presumably for
nectar.
Megachile nevadensis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 209. 2, d.
oslari Mitchell. Colo.
Megachile (Xeromegachile) oslari Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 330. 2.
pagosiana Mitchell. Colo.
Megachile (Xeromegachile) pagosiana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 333. 2.
palmensis Mitchell. South. Calif., Ariz. Pollen: Unknown, but visits flowers of Bebbia juncea,
Cirsium, Echinocactus engelmannii, Echinocereus, Encelia farinosa, Sphaeralcea
rosacea, Viguiera.
Megachile (Xeromegachile) palmensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 317. 6.
Megachile (Xeromegachile) smithi Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 337. °.
parksi Mitchell. Tex.
Megachile (Xeromegachile) parksi Mitchell, 1936. Amer. Ent. Soc., Trans. 62: 346. °.
pseudolegalis Mitchell. Calif. (Riverside Co.). Pollen: Unknown, but visits flowers of Larrea
tridentata.
Megachile (Xeromegachile) pseudolegalis Mitchell, 1957. Pan-Pacific Ent. 33: 22. d.
pseudonigra Mitchell. Oreg., Calif., Ariz. Pollen: Unknown, but visits flowers of Chaenactis
glabriuscula, Cirsium, Oenothera campestris.
Megachile pseudonigra Mitchell, 1927. Psyche 34: 112. 2.
rubi Mitchell. N. C. to Fla. Ecology: Nests in sandy loam. Pollen: Unknown, but visits flowers
of Crataegus, Cuscuta, Ilex, Rubus, Senecio.
Megachile rubi Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 159. 2.
Megachile neglecta Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 160. d.

Biology: Sivik, 1954. Ent. News 65: 255-256 (nest).
seducta Mitchell. Calif. Pollen: Unknown, but visits flowers of Lotus.

Megachile (Xeromegachile) sedwcta Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 342. 2.
soledadensis Cockerell. Tex., N. Mex., Ariz., south. Calif. Parasite: Coelioxys soledadensis Ckll.
Pollen: Apparently an oligolege of Compositae, visits flowers of Baccharis, Baileya
multiradiata, Bebbia juncea, Chrysothamnus, Encelia, Haplopappus, Heterotheca,

Hymenothrix, Verbesina, Viguiera.
Megachile soledadensis Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 12. d, 9.

stoddardensis Mitchell. Calif. (San Bernardino Co.)., Ariz.
Megachile (Xeromegachile) stoddardensis Mitchell, 1957. Pan-Pacifie Ent. 33: 25. d.
subnigra angelica Mitchell. Oreg., Calif., Ariz. Pollen: Unknown, although principally visits
flowers of the Compositae, records include Amsinckia intermedia, Achillea millefolium,
Aster, Baccharis viminea, Balsamorrhiza deltoides, Bebbia juncea, Chaenactis
artemisiaefolia, C. fremontii, C. glabriuscula, Coreopsis lanceolata, Cryptantha
intermedia, Encelia actoni, E. californica, E. farinosa, Eriogonum fasciculatum, E.
confertiflorum, Erysimum, Ferrocactus acanthodes, Haplopappus cooperi, H.
linearifolius, H. pinifolius, Hesperochiron californicus, Isomeris arborea, Lupinus
nanus, Oenothera campestris, Salvia, Sonchus oleraceus, Swertia parryi, Viguiera
laciniata.
Megachile (Xeromegachile) angelica Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 318. 6.
Megachile (Xeromegachile) blaisdelli Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 336. °.
Megachile (Xeromegachile) moschata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 338. 9°.
subnigra subnigra Cresson. Mont., Wyo., Colo., Utah, Idaho, Nev., B. C., Wash., Oreg., Calif,,
restricted to high mts. in south. part of range. Pollen: Unknown, but visits flowers of
Lupinus densiflorus.
Megachile subnigra Cresson, 1879. Amer. Ent. Soc., Trans. 7: 208. 2, d.

toscata Mitchell. Colo.
Megachile (Xeromegachile) toscata Mitchell, 1934, Amer. Ent. Soc., Trans. 59: 341. 9.

victoriana Mitchell. Tex. (Victoria).
Megachile (Xeromegachile) victoriana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 315. ¢.

2066 Hymenoptera in America North of Mexico

wheeleri Mitchell. Sask., N. Dak., Colo. to B. C. and mts. of Calif. and Ariz. Pollen: Unknown,
but visits flowers of Aster foliaceus, Chaenactis glabriuscula, Chenopodium,
Chrysopsis, Chrysothamnus nauseosus speciosus, Cryptantha, Encelia, Grindelia
maritima, G. squarrosa, Lasthenia, Layia glandulosa, Lygodesmia juncea, Medicago
sativa, Solidago, Sphaeralcea ambigua.
Megachile wheeleri Mitchell, 1927. Psyche 34: 107. 2.
Megachile spokanensis Mitchell, 1927. Psyche 34: 109. 6.
wyomingensis Mitchell. Wyo.
Megachile (Xeromegachile) wyomingensis Mitchell, 1934. Amer.’Ent. Soc., Trans. 59: 327.
é.

Genus MEGACHILE Subgenus ARGYROPILE Mitchell

Megachile subg. Argyropile Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 302, 308.
Type-species: Megachile parallela Smith. Orig. desig.

Taxonomy: Mitchell, 1943. Pan-Pacific Ent. 19: 16 (Key to spp.).

asterae Mitchell. Utah. Pollen: Unknown, but visits flowers of Aster.
Megachile (Argyropile) asterae Mitchell, 1943. Pan-Pacifie Ent. 19: 13. 9.
parallela Smith. Transcont. U.S., south. Canada; Mexico. Ecology: Nests in ground, uses

leaf-cuttings from Spiraea and Trifoliwm, but also accepts trap-nests for nesting. Pollen:
Unknown, but visits a wide variety of flowers, notably of the family Compositae,
including Agoseris, Asclepias, Aster, Baccharis, Boltonia, Brassica geniculata,
Calycadenia multiglandulosa, Ceanothus, Centaurea melitensis, C. solstitialis,
Cephalanthus, Chrysanthemum, Chrysothamnus, Cirsium, Clarkia williamsonii,
Coreopsis grandiflora, C. lanceolata, C. tinctoria, Corethrogyne, Encelia farinosa,
Ericameria parishu, Erigeron pygmaeus, Eriophyllum confertiflorum, Gaillardia
pulchella, Gilia, Gossypium, Grindelia camporum, Gutierrezia californica, G. sarothrae,
Haplopappus squarrosus, H. vernonioides, Helenium bigelovii, Helianthus annuus, H.
atrorubens, H. gracilentus, H. nuttallii, H. petiolaris, Heliopsis, Hemizonia lobbi, H.
wrightii, Heterotheca grandiflora, H. subaxillaris, Hypericum, Lepachys, Lotus
scoparius, Malacothrix tenuifolia, Medicago sativa, Melilotus alba, Palafoxia linearis,
Phaseolus, Petalostemon, Ratibida, Rudbeckia, Senecio douglasii, silphium,
Stephanomeria exigua, Verbena, Verbesina, Viguiera, Xanthocephalum, Zexmenia.

Megachile parallela Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 191. 6.

Megachile facunda Cresson, 1872. Amer. Ent. Soc., Trans. 4: 266. d. N. syn.

Megachile sexdentata Robertson, 1895. Amer. Ent. Soc., Trans. 22: 125. ¢.

Megachile verbesinae Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 264. @.

Megachile (Argyropile) parallela var. rita Mitchell, 1937. Amer. Ent. Soc., Trans. 63: 53. 9.

Biology: Fischer, 1951. Kans. Ent. Soc., Jour. 24: 49-50 (nest). —Medler and Lussenhop, 1968.
Wis. Univ. Res. Bul. 274: 26 (nest).
rossi Mitchell. Ariz. Pollen: Unknown, but visits flowers of Aster, Bidens, Cercidium, Encelia,

Verbesina, Viguiera.
Megachile (Argyropile) rossi Mitchell, 1943. Pan-Pacific Ent. 19: 14. 9.

sabinensis Mitchell. Ariz., Tex. Pollen: Unknown, but visits flowers of Aster, Baileya,
Eriogonum, Haplopappus, Helianthus.
Megachile (Argyropile) sabinensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 348. 9.
Megachile (Xeromegachile) pararubi Mitchell, 1957. Pan-Pacific Ent. 33: 21. 9.
subparallela Mitchell. Ariz. Ecology: Nests in ground under Brayulinea densa, uses entire
leaves of Desmodium for cell construction. Pollen: Unknown, but visits flowers of
Eriogonum, Haplopappus, Heterotheca.
Megachile (Argyropile) subparallela Mitchell, 1944. Pan- Pacific Ent. 20: 132. d.
townsendiana Cockerell. N. C. to Fla., west to Okla., Tex., Colo., N. Mex., Ariz., and southeast.
Calif., Mexico. Ecology: Nests in sandy soil. Pollen: Unknown, but visits flowers of
Actinella, Baileya pleniradiata, Chrysopis, Encelia frutescens, Geraea canescens,
Helianthus debilis, Hoffmanseggia, Medicago sativa, Melanthera parviflora, Prosopis,
Verbesina, Wislizenia refracta.
Megachile townsendiana Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 1: 129. 6.

Superfamily APOIDEA 2067

Megachile bishoppi Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 535. 6.

tulariana Mitchell. Calif. (Lemon Cove).
Megachile (Argyropile) parallela var. tulariana Mitchell, 1937. Amer. Ent. Soc., Trans. 63:
Seer

Genus MEGACHILE Subgenus XANTHOSARUS Robertson

Xanthosarus Robertson, 1903. Amer. Ent. Soc., Trans. 29: 168.
Type-species: Megachile latimanus Say. Monotypic and orig. desig.
cochisiana Mitchell. N. Mex., Ariz., Nev., south. and east. Calif., Mexico. Ecology: Uses pieces
cut from leaves of grape for nest construction. Pollen: Unknown, but visits flowers of
Cercidium, Lotus americanus, Medicago sativa.
Megachile (Xanthosarus) cochisiana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 313. ¢.
Megachile (Xanthosarus) pallidiana Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 314. °.

Taxonomy: Mitchell, 1942. Pan-Pacific Ent. 18: 116. 9.
comata Cresson. Kans., Tex., Colo., N. Mex., Ariz., Calif.; Mexico. Pollen: Unknown, but visits
flowers of Asclepias, Gilia, Lotus, Marrubium vulgare, Phacelia, Senecio.
Megachile comata Cresson, 1872. Amer. Ent. Soe., Trans. 4: 262. d.
Megachile armata Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 80. 6.
Megachile paratexana Mitchell, 1930. Amer. Ent. Soc., Trans. 56: 226. 2.

dentitarsus Sladen. Alta., Wash., Mont., N. Dak., S. Dak., Wyo., Nebr., Colo., Utah, N. Mex.,
Ariz. Ecology: Cuts pieces of grape leaves for nesting material. Pollen: Unknown, but
visits flowers of Aster, Cleome serrulata, Grindelia squarrosa, Gutierrezia sarothrae,
Helianthus petiolaris, Medicago sativa, Melilotus, Solidago, Trifolium pratense,
Verbena.
Megachile diligens Sladen, 1918. Agr. Gaz. Canada 5: 125. ¢, 2. Preoce.
Megachile dentitarsus Sladen, 1919. Canad. Ent. 51: 85. N. name.

Taxonomy: Sladen, 1918. Canad. Ent. 50: 303.

innupta Cockerell. Colo.
Megachile innupta Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 534. 9.
latimanus Say. Alta. to N.S., south to Ga., Kans., Wyo. and Colo. Parasite: Coeliorys
funeraria Sm., C. rufitarsis Sm. Pollen: Polylectic, visits a wide variety of flowers
including Abutilon, Agastache, Althaea rosea, Amorpha, Apocynum androsaemifolium,
Arabis, Asclepias, Aster, Astragalus, Baptisia, Bidens, Blephilia, Boltonia, Campanula
petiolata, Carduus, Cassia, Cephalanthus, Chamaenerion, Chrysopsis, Cirsium, Cleome,
Convolvulus, Coreopsis, Epilobium angustifolium, Eupatorium, Gaillardia, Gerardia,
Grindelia, Helenium, Helianthus, Hieracium, Hydrophyllum, Kuhnistera, Lactuca,
Lepachys, Lespedeza, Liatris, Malvastrum, Medicago sativa, Melilotus alba, Mentzelia,
Monarda, Penstemon cyanthus, Petalostemon, Physostegia, Polemonium, Potentilla,
Psoralea, Pycnanthemum, Rosa, Rudbeckia, Ruellia, Solidago, Sonchus, Stachys,
Strophostyles, Taraxacum, Teucrium, Traumeria, Trifolium repens, Verbena,
Verbesina, Vernonia, Vicia.
Megachile latimanus Say, 1823. West. Quart. Rptr. 2: 81. ¢.
Megachile femorata Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 188. 6.
Megachile acuta Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 192. °.
Megachile vidua Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 192. 2.

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 162-163, figs. 4, 4a, 5 (life history, nest
architecture, parasite). —Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 24 (nest,

parasite).

Morphology: Mitchell, 1932. Elisha Mitchell Sci. Soc., Jour. 47: 52-54, pl. 4 (gynandromorph).
perihirta Cockerell. Alta. to Nebr. and Tex., west to B. C. and Calif., Mich.?; Mexico. Parasite:
Coelioxys grindeliae Ckll., C. octodentata Say, C. rufitarsis Sm., Dasymutilla fulvohirta
(Cress.) Nemognatha lutea LeC., Physocephala fronto (Will.). Pollen: Polylectic, visits
flowers of many families, especially those of the family Compositae; visitation records
include Arnica, Asclepias galioides, Aster adscendens, A. canescens, A. foliaceus,

2068

Hymenoptera in America North of Mexico

Astragalus parishu, Bidens laevis, Castilleia miniata, Calendula, Centaurea
solstitialis, Chrysopsis villosa, Chrysothamnus nauseosus, C. n. speciosus, C.
viscidiflorus typicus, Cirsium andersonii, C. californicum, C. lanceolatum, C.
tioganum, Clarkia viminea, Cleome serrulata, Coreopsis lanceolata, Cosmos
bipinnatus, Dicentra chrysantha, Encelia farinosa, Epilobium angustifolium, Erigeron
glaucus, Eriogonum fasciculatum, E. f. var. polifolium, E. latifolium var. nudum,
Eschscholzia californica, Eupatorium occidentale, Gormania obtusata, Grindelia
platyphylla, G. stricta var. procumbens, Gutierrezia sarothrae, Haplopappus
apargioides, H. vernonioides, Helenium bigelovii, Helianthus annuus, H. petiolaris,
Horkelia bernardina, Lathyrus odoratus, Lotus argophyllus, L. glaber, L. oblongifolius,
Lotus scoparius, Lupinus paynei, Medicago sativa, Melilotus officinalis, Microseris
nutans, Mimulus tillingii, Phacelia frigida, P. ramosissima, Pluchea camphorata,
Potentilla, Robinia, Rosa californica, Senecio douglasii, S. integerrimus, Solidago
californica, S. confinis, S. multiradiata, S. occidentalis, Spiraea densiflora,
Stephanomeria virgata, Symphoricarpos, Taraxacum officinale, Trichostema
lanceolatum, T. involucratum, Verbena, Vicia, Wyethia.

Megachile perihirta Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 1: 126. 6.
Megachile latimanus grindeliarum Cockerell, 1904. Ent. News 15: 33. @.

Biology: Hicks, 1926. Colo. Univ. Studies 15: 233 (nest, parasite). —Hicks, 1936. Canad. Ent.
68: 49 (nest, parasite). —Hobbs, 1957 (1956). Canad. Ent. 88: 625-631 (life history,
parasites).

Genus MEGACHILE Subgenus LEPTORACHIS Mitchell

Megachile subg. Leptorachis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 301, 308.

Type-species: Megachile petulans Cresson. Orig. desig.

petulans Cresson. N. J. to Fla., west to N. Dak., Nebr., and Ariz., Mexico. Pollen: Polylectic,

visits many flowers, especially Compositae, Labiatae, Leguminosae; visitation records
include flowers of Achillea, Apocynum, Asclepias, Aster, Baptisia, Bidens, Blephilia,
Buddleia, Brauneria, Campanula rotundifolia, Cassia, Cephalanthus, Chamaecrista,
Chrysanthemum leucanthemum, Cicuta, Cirsium, Coreopsis stellata, Crotalaria,
Desmodium, Eryngium, Flaveria, Galactia, Gerardia, Grindelia, Helenium, Helianthus
atrorubens, Houstonia purpurea, Hydrolea, Hypericum, Hyptis, Ilex, Koellia, Lespedeza
repens, Liatris, Lobelia, Ludwigia, Lycopus, Lythrum, Melilotus alba, Nepeta,
Pastinaca, Phaseolus, Polygonum, Psoralea, Pycnanthemum, Rhododendron, Rhus,
Rubus, Rudbeckia, Senecio, Silphium, Solidago, Strophostyles, Tephrosia, Trifolium,
Verbena, Verbesina, Vernonia glauca, Vicia.

Megachile petulans Cresson, 1878. Amer. Ent. Soc., Trans. 7: 127. d.
Megachile mexicana Cresson, 1878. Amer. Ent. Soc., Trans. 7: 127. 2, 3.

Genus MEGACHILE Subgenus PSEUDOCENTRON Mitchell

Megachile subg. Pseudocentron Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 303, 307.

Type-species: Megachile pruina Smith. Orig. desig.

bidentata (Fabricius). Supposedly North American, but probably Neotropical.

Andrena bidentata Fabricius, 1775. Syst. Ent., p. 377. 3.

morio Smith. “North America.”

Megachile morio Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 189. 2. Probably a form of

pruina.

pruina nigropinguis Mitchell. Tex.

Megachile (Pseudocentron) pruina var. nigropinguis Mitchell, 1937. Amer. Ent. Soc.,

Trans. 63: 68. °.

pruina pruina Smith. N. C., Ga., Fla., Bermuda. Pollen: Unknown, but visits flowers of

Asclepias tuberosa, Crotalaria, Croton linearis, Cuscuta, Dalbergia ecastophyllum,
Gaillardia, Galactia floridana, Helianthus, Melanthera parviflora, Ocimum, Rhus,
Strophostyles, Vernonia blodgettii.

Megachile pruina Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 190. °.
Megachile pinguis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 125. 6.

Superfamily APOIDEA 2069

Megachile floridana Robertson, 1895. Amer. Ent. Soc., Trans. 22: 125. d.
Megachile shermani Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 163. ¢.
Megachile pruina var. bermudensis Mitchell, 1929. Psyche 36: 93. 9.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 337.
sidalceae Cockerell. Tex., N. Mex., Ariz., south. Calif., Mexico. Ecology: Uses pieces of leaves

cut from Amaranthus, Boerhaavia, Fraxinus, alfalfa, chinaberry, cotton, cottonwood,
rose, and umbrella tree for nesting material. Parasite: Coelioxrys novomexicana Ckll.
Pollen: Polylectic, visits a wide variety of flowers including Acacia, Aloysia, Asclepias,
Asparagus, Baccharis, Cephalanthus, Cercidium, Cleome, Condalia, Croton
californicus, Encelia, Eriogonum, Ferocactus, Franseria, Funastrum, Gutierrezia,
Haplopappus, Helianthus, Hoffmanseggia, Hymenothrix, Kallstroemia grandiflora,
Larrea tridentata, Lepidium, Lotus, Medicago sativa, Melilotus, Mimosa, Mortonia,
Olneya tesota, Opuntia, Pectis papposa, Prosopis glandulosa var. glandulosa,
Psilostrophe cooper, Salix, Salsola, Sapindus, Senecio douglasii, Sidalcea malvaeflora,
Tamarix, Verbesina, Wislizenia.

Megachile sidalceae Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 158. d.

Megachile abducta Mitchell, 1926. Amer. Ent. Soc., Trans. 52: 111. ¢.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 38 (floral relationships with
Larrea).

Genus MEGACHILE Subgenus ACENTRON Mitchell

Megachile subg. Acentron Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 303, 307.
Type-species: Megachile albitarsis Cresson. Orig. desig.
albitarsis Cresson. Mich., Ind., N. C., Ga., Fla., Ala., Miss., La., Tex., N. Mex., Ariz.; Mexico.

Pollen: Apparently polylectic, visits flowers of Asclepias, Aster, Bidens, Cassia,
Cephalanthus, Chrysopsis, Coreopsis, Cyrilla, Eryngium, Flaveria, Gaillardia,
Galactia, Helenium, Helianthus, Ilex, Ipomoea, Kallstroemia grandiflora, Lespedeza
repens, Liatris, Melilotus, Phaseolus, Pycnanthemum hypsopifolia, Rhus, Rudbeckia,
Solidago, Stokesia, Verbena, Xyris caroliana.

Megachile albitarsis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 263. d.

Megachile optiva Cresson, 1872. Amer. Ent. Soc., Trans. 4: 268. °.

Megachile newelli Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 262. 2.

Megachile kallstroemiae Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 264. °.

Genus MEGACHILE Subgenus MELANOSARUS Mitchell

Megachile subg. Melanosarus Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 303, 307.
Type-species: Megachile xylocopoides Smith. Orig. desig.
bahamensis Mitchell. Fla.; Bahamas. Pollen: Unknown, but visits flowers of Bidens leucantha,
Flaveria, Melanthera brevifolia, Poinsettia cyathophora.
Megachile bahamensis Mitchell, 1927. Psyche 34: 47. °.
Megachile (Melanosarus) floridensis Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 349. d, °.
xylocopoides Smith. Md., N. C., Ga., Fla., Ala., Miss., La., Tex. Parasite: Coelioxrys dolichos
Fox, Leucospis affinis Say, Melittobia possibly megachilis (Pack.), Tetrastichus
megachilidis Burks. Pollen: Polylectic, visits flowers of Bidens leucantha, Borrichia,
Cephalanthus, Citrus limonum, Elephantopus carolinianus, Gaillardia, Helenium,
Helianthus, Ilex, Itea, Mikania, Oxypolis, Phaseolus, Poinsettia cyathophora,
Polygonum, Rhus, Rubus, Rudbeckia hirta, Senecio, Solidago, Trifolium, Vernonia.
Megachile xylocopoides Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 189. 9, d.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 329-331, pl. 20, fig. 98 (life
history, nest architecture, supersedure, parasites).
Genus MEGACHILE Subgenus SAYAPIS Titus

Gnathocera Provancher, 1883. Nat. Canad. 13: 232. Preoce.
Type-species: Megachile pugnatus Say. Monotypic. (=Gnathocera cephalica

Provancher).

2070 Hymenoptera in America North of Mexico

Ceratias Robertson, 1903. Amer. Ent. Soce., Trans. 29: 168. Preoce.
Type-species: Megachile pugnatus Say. Orig. desig.

Sayapis Titus, 1905. Ent. Soc. Wash., Proc. 7: 154. Proposed to replace Gnathocera

Provancher and Ceratias Robertson.
dentipes Vachal. South. Tex. to Paraguay.
Megachile dentipes Vachal, 1909. Rev. de Ent. 28: 12. 3.
Megachile poculifera Cockerell, 1919. U. S. Natl. Mus., Proe. 55: 217. 6.
fidelis Cresson. Mont. and S. Dak., Nebr., N. Mex., west to Idaho, Oreg., Calif. Pollen:

Polylectic, visits a wide variety of flowers, notably those of the family Compositae;
visitation records include Aster canescens, A. delectabilis, Calycadenia multiglandulosa,
Ceanothus leucodermis, Chrysopsis fastigiata, C. villosa, Chrysothamnus nauseosus, C.
viscidiflorus typicus, Cirsium, Cleome serrulata, Cleomella, Coreopsis lanceolata,
Croton californicus, Cosmos, Daucus carota, Erigeron divergens, E. stenophyllus,
Eriogonum fasciculatum var. polifolium, E. latifolium var. nudum, E. subscaposum, E.
wright, Grindelia camporum, G. stricta var. procumbens, Gutierrezia californica, G.
sarothrae, Haplopappus arborescens, H. bloomeri var. angustatus, Helenium bigelovii,
H. puberulum, Helianthus gracilentus, H. petiolaris, Heliopsis, Heterotheca, Lotus
scoparius, Lupinus, Marrubium vulgare, Mentha pulegium, Monarda, Phacelia,
Rhamnus californica, Rudbeckia californica, Senecio douglasii, Solidago californica, S.
confinis, Verbena, Viguiera.

Megachile fidelis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 120. 9, 6.

Megachile fidelis var. concinnula Cockerell, 1899. Entomologist 32: 158. 6.

frugalis frugalis Cresson. Pa. and N. J., to Fla., west to Calif.; Mexico. Pollen: Polylectie, visits
flowers of Croton, Erigeron, Eriogonum fasciculatum, Helianthus, Lotus, Marrubium
vulgare, Opuntia, Perezia microcephala, Tephrosia virginiana.
Megachile frugalis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 266. d.
frugalis pseudofrugalis Mitchell. Ariz., Calif. Pollen: Polylectic, visits a wide variety of
flowers including Acacia greggii, Asclepias erosa, Aster spinosus, Bebbia juncea,
Cercidium, Chaenactis, Chilopsis linearis, Clarkia, Cirsium vulgare, Cissus, Cleome
serrulata, Croton californicus, Cryptantha, Dalea emoryi, Encelia californica, E.
farinosa, Eriodictyon crassifolium, Eriogonum elongatum, E. fasciculatum, E. f. var.
polifolium, E. wrightu, Grindelia, Gutierrezia camporum, G. sarothrae, Haplopappus
squarrosus, Helianthus gracilentus, Hemizonia fasciculata, Hyptis emoryi, Lotus
hamatus, L. scoparius, Marrubium vulgare, Melilotus, Perezia microcephala,
Penstemon, Petalonyx thurberi, Prosopis glandulosa var. torreyanum, P. pubescens,
Salvia clevelandii, Sapindus, Senecio douglasii, Tamarix gallica, Trichostema parishii,
Verbesina encelioides, Wislizenia refracta.
Megachile (Sayapis) frugalis var. pseudofrugalis Mitchell, 1937. Amer. Ent. Soc., Trans.
63: 186. 3, °.
helianthi Cockerell. Colo. Pollen: Unknown, but visits flowers of Helianthus lenticularis.
Megachile helianthi Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 259. 9.
inimica inimica Cresson. Fla. to Tex., Ariz.; Mexico south to Guatemala. Ecology: Nests in
holes in mesquite trees and mesquite fence posts, and uses leaves of Monisia pallida to
line their nests; also nests in trap nests which were plugged with 5mm. of firmly
agglutinated sand by this species. Parasite: Melittobia chalybii Ashm. Pollen: Polylectic,
visits flowers of Achyranthes, Avicennia, Bidens leucantha, Borrichia frutescens,
Coccoloba, Crotalaria pumila, Helenium, Heliopsis, Heterotheca, Lacinaria punctata,
Melilotus, Ocimum, Salvia ballotaeflora, Sphaeralcea, Vitex negundo var. incisa,
Ximensia encelioides.
Megachile inimica Cresson, 1872. Amer. Ent. Soc., Trans. 4: 267. 9.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 331-332 (life history, nest
architecture, parasite).
inimica jacumbensis Mitchell. Calif.; Mexico (Baja California). Pollen: Apparently polylectic,
visits flowers of Chrysothamnus, Eriogonum fasciculatum, Haplopappus arborescens,
Heterotheca grandiflora, Grindelia camporum, Senecio douglasii, Solidago.
Megachile inimica var. jacumbensis Mitchell, 1927. Psyche 34: 109.

Superfamily APOIDEA 2071

inimica sayi Cresson. Pa. to Fla., west to Idaho and Calif.; Mexico. Ecology: Nests in borings,

and closing plug is 5mm. thick consisting of leaf cuttings, pebbles, and leaf pulp. Pollen:
Polylectic, especially on flowers of the family Compositae (Heliantheae, Cynarieae,
EKupatorieae), visits a wide variety of these and other flowers including Arctium, Aster,
Bidens, Blephilia, Brauneria, Cephalanthus, Chrysopsis mariana, Chrysothamnus
nauseosus consimilis, Cleome, Coreopsis stellata, Gilia, Gutierrezia californica,
Haplopappus, Helenium, Helianthus petiolaris, Heliopsis helianthoides, Heterotheca,
Kallstroemia grandiflora, Lepachys, Liatris, Lythrum, Marrubium vulgare, Melilotus
alba, Penstemon, Petalostemon, Pycnanthemum, Rudbeckia, Senecio, Silphium,
Solidago, Strophostyles, Teucrium, Verbena, Verbesina encelioides, Vernonia
fasciculata, V. glauca, Vicia, Viguiera, Zexmenia.

Megachile sayi Cresson, 1878. Amer. Ent. Soc., Trans. 7: 119. 6, 9.

Megachile heterodonta Cockerell, 1900. Ann. and Mag. Hist. (7) 6: 18. 2.

Megachile sayi paludicola Cockerell, 1911. U. S. Natl. Mus., Proce. 40: 254. 9°.

Megachile sayi var. sancta Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 10: 487. 3.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 332-334 (life history, nest
architecture, supersedure). —Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 23-24
(life history, nest).

mellitarsis Cresson. B. C., Wash., Oreg., Calif., Nev., Colo., N. Mex., Ariz. Pollen: Apparently
polylectic, visits flowers of Aster, Chrysothamnus, Encelia, Eriogonum, Phacelia,
Solidago multiradiata.
Megachile mellitarsis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 121. 6.
Megachile terrestris Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 1: 260. 2. Preoce.
Megachile geophila Cockerell, 1908. Canad. Ent. 40: 460. N. name.
newberryae Cockerell. N. Mex., Ariz., Calif., Mexico, deserts. Ecology: Nests in holes in
Prosopis and uses pieces of leaves cut from Celtis for cell construction. Pollen:
Apparently polylectic, visits flowers of Acacia greggii, Asclepias, Aster, Cercidium
torreyanum, Encelia, Funastrum, Helenium, Larrea tridentata, Melilotus alba, Olneya
tesota, Opuntia, Pittosporum, Prosopis glandulosa var. torreyanum, P. juliflora, P.
pubescens, Senecio douglasii, Verbesina encelioides.
Megachile newberryae Cockerell, 1900. Entomologist 33: 244. 6.
Megachile howardi Cockerell, 1919. U. S. Natl. Mus., Proc. 55: 216. ¢ (2 misdet.).

Biology: Butler, 1965. Ariz. Agr. Exp. Sta. Tech. Bul. 187: 14 (nest).
policaris Say. Fla., La., Nebr., Kans., Okla., Tex., Colo., N. Mex., Ariz., Calif.; Mexico. Ecology:

Uses leaflets from Eysenhardtia polystachya, Mimosa biuncifera, and Prosopis for plug
construction; nests in trap nests, the larvae develop amicably in a single large brood cell
with no apparent cannibalism. Parasite: Coeliorys texana Cress., Leucospis affinis Say,
Melittobia chalybii Ashm. Pollen: Polylectic, especially flowers of the Compositae
(Heliantheae), but in some localities uses pollen solely from Prosopis for nest provision;
visitation records include Acacia, Amorpha fruticosa, Asclepias, Baccharis, Bidens,
Brauneria pallida, Chrysothamnus, Condalia, Coreopsis lanceolata, Croton
californicus, Encelia californica, Erigeron, Eriogonum inflatum, Gutierrezia sarothrae,
Haplopappus acradenius, Helianthus, Larrea tridentata, Melilotus, Oenothera, Opuntia,
Palafoxia linearis, Petalostemon, Salsola, Salvia ballotaeflora, Senecio douglasii,
Silphium, Solidago, Tamarix, Verbena, Verbesina encelioides, Viguiera multiflora.
Predator: Pyemotes ventricosus (Newport).

Megatchile policaris Say, 1831. Deser. New Spp. No. Amer. Ins. Found in La. by J.
Barabino, p. 17. °.

Megachile pollicaris Say, 1837. Boston Jour. Nat. Hist. 1: 406. ¢. Emend.

Megachile grandis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 268. 2. Preoce.

Megachile pollicaris var. perexima Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 12. ¢.

Megachile vallorum Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 18. 2.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 334-337, pl. 19, figs. 92-97 (life
history, nest architecture, larval food, supersedure, parasite). —Hurd and Linsley, 1975.
Smithsn. Contrib. Zool. 193: 38 (floral relationships).

2072 Hymenoptera in America North of Mexico

pugnata pomonae Cockerell. Nev., Calif., Ariz. Pollen: Presumably polylectic, visits flowers of

Arctium, Chrysothamnus, Cirsium, Dicentra chrysantha, Mimulus, Raillardella
scaposa, Senecio, Solidago, Sphenosciadium capitellatum, Symphoricarpos, Taraxacum
officinale, Viguiera multiflora, Wyethia mollis.

Megachile pugnata var. pomonae Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 278. 9.

pugnata pugnata Say. U.S. and south. Canada except lower Mississippi Valley, Gulf Coast,

and Calif., principally at moderate to high altitudes in southern part of range. Ecology:
Nests in borings in sumac twigs. Parasite: Coelioxys alternata Say. Pollen: Polylectic,
especially flowers of Compositae (Heliantheae; Cynarieae), visits flowers of Achillea
lanulosa, Arctium minus, Asclepias, Aster, Besseya plantaginea, Blephilia, Brauneria
pallida, Carduus plattinus, C. undulatus, Chrysanthemum leucanthemum, Cicuta,
Cirsium californicum, Cleome, Coreopsis, Dianthera, Erigeron, Helianthus tuberosus,
Lepachys, Marrubium vulgare, Melilotus, Nepeta, Phacelia, Rudbeckia hirta, R.
occidentalis, Senecio, Silphium, Solidago, Taraxacum officinale, Trifolium repens,
Verbena, Verbesina, Vernonia.

Megachile pugnatus Say, 1837. Boston Jour. Nat. Hist. 1: 408. d, @.

Megachile scobiculata Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 191. ¢.

Megachile bucephala Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 193. °.

Megachile disparilis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 264. ¢.

Megachile lucrosa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 267. 2.

Gnathocera cephalica Provancher, 1882. Nat. Canad. 13: 233. 9.

Megachile temporalis Friese, 1903. Ztschr. System. Hym. Dipt. 3: 247. 2 (¢ misdet.).

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 337.

Biology: Medler, 1964. Canad. Ent. 96: 918-921, 1 fig. (life history, nest architecture,
parasite). —Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 27 (nest, parasites).

NOMINA NUDA IN MEGACHILE

Megachile coloradensis Uhler, 1877. U.S. Geol. Geog. Survey, Bul. 3: 784.
Megachile consimilis Evans, 1896. Canad. Ent. 28: 13.

Genus CHALICODOMA Lepeletier

This genus, which is represented in North America by two subgenera, contains a number of
subgenera in the Old World and one subgenus is present in South America. The bees of this
genus do not use pieces of leaves or petals in the construction of their nests, but rather use
resin, mud and other such materials.

Revision: Mitchell, 1937. Amer. Ent. Soc., Trans. 63: 381-426, pls. XX VI-X XIX (Part VIII,
treats Nearctic spp. of the subgenus Chelostomoides; addenda and index to previous
parts).

Taxonomy: Mitchell, 1956. Pan-Pacific Ent. 32: 129-138 (Key to Nearctic and Neotropical
spp.). —Michener, 1962. N. Y. Ent. Soc., Jour. 70: 17-29 (classification). —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 181-190, figs. 59-62, table 6 (eastern U. S. spp.).
—Pasteels, 1965. Mus. Royal l’ Afrique Centr., Ann. Sci. Zool. 137: ix and 579 pp.
(classification). —Butler, 1965. Ariz. Agr. Expt. Sta. Tech. Bul. 187: 1-19 (Ariz. spp.).
—Michener, 1965. Amer. Mus. Nat. Hist., Bul. 130: 185-186 (classification). —Stephen,
Bohart and Torchio, 1969. The biology and external morphology of bees, pp. 53-54
(classification).

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 337-344 (life histories, nests,
associates). —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 38-39, fig. 16 (Larrea
visiting spp.).

Genus CHALICODOMA Subgenus CHALICODOMA Lepeletier

Chalicodoma Lepeletier, 1841. Hist. Nat. Ins., Hym. 2: 309.
Type-species: Apis muraria Fabricius. Desig. by Girard, 1879.

The typical subgenus does not occur in North America.

Superfamily APOIDEA 2073

Genus CHALICODOMA Subgenus CHELOSTOMOIDES Robertson

Chelostomoides Robertson, 1901. Canad. Ent. 33: 231.
Type-species: Chelostoma rugifrons Smith. Monotypic and orig. desig. (~Megachile
rufimanus Robertson).
Oligotropus Robertson, 1903. Amer. Ent. Soc., Trans. 29: 168.
Type-species: Oligotropus campanulae Robertson. Monotypic and orig. desig.
Gnathodon Robertson, 1903. Amer. Ent. Soc., Trans. 29: 168. Preoce.
Type-species: Megachile georgica Cresson. Monotypic and orig. desig.
Sarogaster Robertson, 1918. Ent. News 29: 92. Proposed to replace Gnathodon Robertson.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1059, figs. 162-164 (larva.) —Mitchell,
1956. Pan-Pacifie Ent. 32: 134-138 (revised key to spp.). —Medler and Lussenhop, 1968.
Wis. Univ. Res. Bul. 274: 30-32 (Wis. spp.).

adelphodonta (Cockerell). Ariz.; Mexico (Baja California). Pollen: Unknown, but visits flowers
of Allionia, Kallstroemia grandiflora.
Megachile adelphodonta Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 548. 6.
Megachile (Chelostomoides) tucsonensis Mitchell, 1956. Pan-Pacific Ent. 32: 133. °.

angelarum (Cockerell). B. C., Wash., Oreg., Calif., Nev., Ariz. Parasite: Sapyga angustata

(Cress.). Pollen: Polylectic, visits a wide variety of flowers including Achillea
millefolium, Adenostegia villosa, Amorpha californica, Aster adscendens, A. canescens,
Astragalus parishii, Brassica incana, Calycadenia multiglandulosa, Centaurea
solsitialis, Cercidium torreyanum, Chamaenerion angustifolium, Chrysopsis fastigiata,
C. fremonti, C. villosa, Chrysothamnus nauseosus, Clarkia unguiculata, Cirsiwm
vulgare, Collinsia tinctoria, Cordylanthus filifolius, C. nevinii, C. pilosus, C. rigidus,
Corethrogyne filaginifolia, Cryptantha ambfgua, C. denticulata, C. intermedia, Dicentra
chrysantha, Encelia farinosa, Eriodictyon, Eriogonum fasciculata, E. f. var. polifolium,
E. latifolium var. nudum, E. subscaposum, E. wrightii, Erysimum asperum, Franseria,
Geranium richardsonii, Gilia capitata, Glycyrrhiza lepidota, Grindelia camporum, G.
hallti, Haplopappus arborescens, Helianthus gracilentus, Heliotropium curvassavicum,
Horkelia bernardina, H. bolanderi var. parryi, Lathyrus, Lessingia leptoclada, Lonicera
interrupta, Lotus americanus, L. argophyllus, L. crassifolius, L. davidsonii, L. glaber, L.
humistratus, L. nevadensis, L. oblongifolius, L. scoparius, L. torreyi, Lupinus
austromontanus, L. longipes, Marrubium vulgare, Melilotus alba, M. officinalis,
Mentzelia laevicaulis, Monardella lanceolata, M. linoides, M. stricta, Penstemon
bridgesii, P. grinnellii, P. palmer, P. rothrocki, Perideridia gairdneri, Phacelia
heterophylla, P. imbricata, P. ramosissima, Potentilla bernardina, P. glandulosa,
ranunculus, Rhamnus californica, Rudbeckia californica, Salvia apiana, S.
pachyphylla, Solidago californica, S. confinis, S. occidentalis, Stachys albens, S.
pycnantha, Swertia parryi, Symphoricarpos, Trichostema laxum, T. parishiu, Trifolium
variegatum, Verbena californica, V. prostrata, Vitex.

Megachile angelarum Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 70. °.

browni (Mitchell). Ariz., south. Calif.; Mexico (Baja California). Pollen: Unknown, but visits

flowers of Acacia greggii, Asclepias, Canotia, Dalea, Frankenia palmeri, Opuntia,
Prosopis juliflora.

Megachile (Chelostomoides) browni Mitchell, 1934. Amer. Ent. Soc., Trans. 53: 354. d.

Megachile (Chelostomoides) felipiana Mitchell, 1937. Amer. Ent. Soc., Trans. 63: 405. 2.

Taxonomy: Timberlake, 1957. Pan-Pacific Ent. 33: 1382 (synonymy).
campanulae campanulae (Robertson). Que. and Mass. to Fla., west to Iowa, Wis., Minn.,
Nebr., Kans. and Tex. Ecology: Nests in borings. Pollen: Unknown, but visits flowers of
Asclepias, Baptisia tinctoria, Campanula americana, C. campanulae, C. rotundifolia,
Desmodium canadense, Epilobium angustifolium, Galactia, Helianthus, Lobelia,
Lythrum, Malva sylvestris, Melilotus, Nepeta, Oenothera, Psoralea, Pycnanthemum,
Rudbeckia, Solidago, Strophostyles, Symphoricarpos, Verbena, Veronica stricta.
Oligotropus campanulae Robertson, 1903. Amer. Ent. Soc., Trans. 29: 171. °, ¢.

Biology: Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 202 (nest). —Krombein, 1967.
Trap-nesting wasps and bees, p. 341 (life history, nest architecture). —Medler and
Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 31 (nest).

2074 Hymenoptera in America North of Mexico

campanulae wilmingtoni (Mitchell). Fla. to Va., coastal. Ecology: Nests in borings. Parasite:
Coelioxys modesta Sm. Pollen: Unknown, but visits flowers of Galactia, Melilotus,

Pontederia, Solidago, Strophostyles.
Oligotropus wilmingtoni Mitchell, 1924. Elisha Mitchell Sci. Soc., Jour. 40: 156. 2, d.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 341-342 (life history, nest
architecture, parasite).
chilopsidis (Cockerell). Tex., N. Mex., Ariz., Calif., Mexico, deserts. Parasite: Nemognatha

scutellaris Lec. Pollen: Apparently polylectic, visits flowers of Acacia greggii,
Cercidium floridum, C. microphyllum, C. torreyanum, Chilopsis linearis, Cirsium,
Dalea californica, D. spinosa, Encelia farinosa, Helianthus, Heliotropium
curassavicum, Hoffmanseggia, Larrea tridentata, Marrubium vulgare, Medicago sativa,
Melilotus alba, Mimosa, Olneya tesota, Prosopis glandulosa var. torreyanum, Sapindus,
Senecio, Vauquelinia, Verbesina encelioides.

Lithurgus oblongus Fox, 1893. Calif. Acad. Sci., Proc. (2) 4: 20. ?. Preoce.

Megachile chilopsidis Cockerell, 1900. Ann. and Mag. Hist. (7) 6: 17. 9.

Megachile longula Fox, 1902. Ent. News 13: 137. N. name.

Megachile pratti Cockerell, 1913. Ann. and Mag. Nat. Hist. (8) 11: 541. ¢ (¢ misdet.).

Taxonomy: Mitchell, 1956. Pan-Pacific Ent. 32: 129-130 (synonymy). —Timberlake, 1957.
Pan-Pacific Ent. 33: 132 (synonymy).
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 38 (floral relationships).
davidsoni (Cockerell). South. Calif., Ariz. Pollen: Possibly oligolectic on Dicentra including D.
chrysantha, but visits other flowers including Fremontia, Lupinus austromontanus,
Penstemon, Senecio.

Megachile davidsoni Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 70. °.
Megachile occidentalis var. leucotricha Cockerell, 1902. South. Calif. Acad. Sci., Bul. 1: 187.

3
discorhina (Cockerell). Calif., Ariz., N. Mex.; Mexico (Baja California). Parasite: Lewcospis
affinis Say. Pollen: Polylectic, visits flowers of Acacia greggii, Baileya, Cercidium
floridum, C. torreyanum, Coursetia, Dalea schottii, Encelia farinosa, Hyptis emoryi,
Larrea tridentata, Lepidium, Medicago sativa, Melilotus, Prosopis glandulosa var.
torreyanum, Senecio, Sphaeralcea ambigua.
Megachile discorhina Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 549. °.

Taxonomy: Mitchell, 1956. Pan-Pacific Ent. 32: 131 (tax. status).
Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 38-39, fig. 16 (floral
relationships).
exilis exilis (Cresson). Tex., N. Mex., Ariz. Pollen: Unknown, but visits flowers of Baptisia
tinctoria, Chilopsis, Coreopsis, Dalea, Lespedeza repens, Melilotus, Polygala, Tephrosia
virginiana.
Megachile exilis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 265. 6.
Megachile studiosa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 269. °.
exilis parexilis (Mitchell). Md., Va., N. C., Ga., Fla., Ind., Ala., Miss., Ark., Tex., Ariz. Ecology:
Nests in borings. Pollen: Unknown, but visits flowers of Asclepias, Baccharis, Baptisia,
Ceanothus, Chilopsis, Condalia, Crotalaria, Erigeron, Galactia, Hydrolea, Ilex,
Melilotus, Mentha, Monarda, Oenothera, Phaseolus, Rhus, Tephrosia, Trifolium,

Vaccinium, Vicia.

Megachile (Chelostomoides) exilis var. parexilis Mitchell, 1937. Amer. Ent. Soc., Trans. 63:
393. 3.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 339-340 (life history, nest
architecture).
georgica (Cresson). N. J. to Fla., west to Ark. and Tex. Ecology: Nests in borings. Parasite:

Coelioxys modesta Sm., Leucospis affinis floridana Cress. Pollen: Unknown, but visits
flowers of Afzelia cassinoides, Amorpha, Baptisia tinctoria, Chrysopsis, Clethra
alnifolia, Crotalaria, Desmodium, Erigeron, Galactia, Helenium, Hypericum, Lobelia,

Superfamily APOIDEA 2075

Melilotus, Penstemon, Phaseolus, Psoralea, Pycnanthemum hypsifolia, Stachys,
Strophostyles, Tephrosia virginiana, Vaccinium.

Megachile georgica Cresson, 1878. Amer. Ent. Soe., Trans. 7: 123. 2, 6.

Megachile penicillata Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 536. °.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 337-339 (life history, nest
architecture, supersedure, parasite).
lobatifrons (Cockerell). N. Mex., Ariz., south. and east. Calif., Mexico. Pollen: Unknown, but
visits flowers of Acacia greggii, Cercidium, Dalea emoryi, Eriogonum fasciculatum, E.
trichopes, Larrea tridentata, Olneya tesota, Prosopis glandulosa var. torreyanum, P.
pubescens, Wislizenia refracta.
Megachile lobatifrons Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 547. 2, 6.
manni (Mitchell). Ariz.

Megachile (Chelostomoides) manni Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 355. 2, 6.
occidentalis (Fox). Tex., N. Mex., Ariz., south. Calif.; Mexico; deserts. Ecology: Nests in adobe
and brick walls as well as in dead wood of a pepper tree and trap nests. Parasite:

Nemognatha lurida LeC. Pollen: Unknown, but visits flowers of Caleopsis, Cercidium

torreyanum, Chilopsis linearis, Cleome, Croton californicus, Duranta plumieri, Geraea
canescens, Helianthus, Hoffmanseggia densiflora, Larrea tridentata, Lotus, Marrubiwm
vulgare, Medicago sativa, Nolina parryi, Phacelia, Phyla nodiflora, Pluchea
camphorata, Prosopis, Schinus molle, Ruellia, Senecio, Tamarix gallica, Verbesina,
Vernonia, Vicia cracca, Wislizenia refracta.

Megachile occidentalis Fox, 1894. Calif. Acad. Sci., Proc. (2) 4: 117. 6.

Megachile prosopidis Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 6: 16. 9.

Megachile prosopidis var. testudinis Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 550. °.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 343-344 (life history, nest
architecture, supersedure). —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 39
(floral relationships).

odontostoma (Cockerell). N. Mex., Ariz., south. Calif., Mexico, deserts. Pollen: Unknown, but
visits flowers of Acacia greggii, Asclepias, Baileya, Cercidium floridum, Chaenactis
stevioides, Encelia farinosa, Helianthus, Heliotropium, Lepidium, Medicago sativa,
Melilotus, Palafoxia linearis, Prosopis glandulosa var. torreyanum, Senecio, Verbesina
encelioides.
Megachile odontostoma Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 27: 550. 2.
Megachile (Chelostomoides) duplexa Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 353. ¢.

Taxonomy: Mitchell, 1956. Pan-Pacific Ent. 32: 131 (synonymy).
rugifrons (Smith). N. C., Ga., Fla., Mich., Ill., Nebr., Kans., Ark., Tex. Pollen: Unknown, but
visits flowers of Apocynum, Blephilia, Coreopsis, Erigeron, Hypericum, Lobelia,
Nepeta, Pontederia, Psoralea, Tephrosia, Vaccinium, Verbena, Veronicastrum
virginicum.
Chelostoma rugifrons Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 220. °.
Chelostomoides rugifrons Cockerell, 1904. Canad. Ent. 36: 301. 2.
Megachile rufimanus Robertson, 1891. Amer. Ent. Soc., Trans. 18: 65. 2, d.
spinotulata (Mitchell). Tex., N. Mex., Ariz., south. Calif. Parasite: Lewcospis affinis Say.
Pollen: Polylectic, visits flowers of Adenostegia filifolia, Allionia, Asclepias erosa,
Boerhaavia spicata, Calochortus splendens, Cercidium, Chrysopsis fastigiata,
Cordylanthus filifolius, C. nevinii, C. rigidus, Croton californicus, Cryptantha, Dalea
californica, Eriodictyon, Eriogonum fasciculatum, E. inflatum, E. swbscaposum, E.
trichopes, Geranium, Helianthus, Lotus americanus, L. argophyllus, L. Hamatus, L.
scoparius, Marrubium vulgare, Mentzelia laevicaulis, Phacelia ramosissima, Robinia,
Senecio, Sapindus, Swertia parryi.
Megachile (Chelostomoides) spinotulata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 357. 2,
é.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 39 (floral relationships).

subexilis (Cockerell). Nebr., Colo., N. Mex., Utah, Ariz., Nev., Calif. Ecology: Nests in adobe
wall and also in borings. Parasite: Coelioxys gilensis Ckll., Dioxys pomonae Ckll. Pollen:

2076 Hymenoptera in America North of Mexico

Unknown, but visits flowers of Amorpha, Chamaebatia millefolium, Geranium,
Helianthus, Lathyrus odoratus, Lotus, Marrubium vulgare, Medicago sativa, Melilotus,
Monarda, Opuntia, Phacelia, Phaseolus, Psoralea tenuiflora, Salvia, Trifolium,
Verbesina encelioides, Vicia.

Megachile subexilis Cockerell, 1908. Entomologist 41: 292. 2.

Megachile semiexilis Cockerell, 1908. Entomologist 41: 292. 9, d.

Biology: Hicks, 1927. Ent. News 38: 17-21 (nest architecture, parasite). —Hurd, 1958. Calif.
Univ. Pubs. Ent. 14: 273 (parasite). —Krombein, 1967. Trap-nesting wasps and bees, p. 343
(life history, nest architecture).

subspinotulata (Mitchell). Ariz.
Megachile (Chelostomoides) subspinotulata Mitchell, 1934. Amer. Ent. Soc., Trans. 59: 360.
OF
texensis (Mitchell). Tex. (Southmost in Cameron Co.). Pollen: Unknown, but visits flowers of
Parkinsonia.
Megachile (Chelostomoides) texensis Mitchell, 1956. Pan-Pacific Ent. 32: 132. 2.

Genus CHALICODOMA Subgenus PSPEUDOMEGACHILE Friese

Megachile subg. Pseudomegachile Friese, 1899. Die Bienen Europas 5: 36.
Type-species: Megachile ericetorum Lepeletier. Desig. by Alfken, 1933.
Megachile subg. Archimegachile Alfken, 1933. Konowia 12: 56.
Type-species: Megachile flavipes Spinola. Orig. desig.
Taxonomy: Pasteels, 1965. Mus. Royal l’Afrique Cent., An. ser. 8, Zool. 137: 377 (synonymy).
lanata (Fabricius). Fla. (Miami). Ecology: Builds clay cells in cavities. An Indian species,
probably introduced into U.S. from Cuba, where it is also adventive.
Apis lanata Fabricius, 1775. Systema Ent., p. 385. °.
Apis purpurea Christ, 1791. Naturgesch. Insekt. Bienen, Wespen u. Ameisengeschl., p. 168,
pl. 13, fig. 7.
Megachile Martindalei Fox, 1891. Amer. Ent. Soc., Trans. 18: 344. 2, 3.

Taxonomy: Moure, 1960. Studia Ent. 3: 108-109 (notes on type). —Pasteels, 1965. Mus. Royal
Afrique Cent., An. ser. 8, Zool. 137: 402-404 (synonymy).

Biology: Horne, 1870. Zool. Soc. London, Trans. 7: 176-177, pl. 19, figs. 11, lla, 11b. —Bodkin,
1918. Ent. Soe. London, Trans. p. 303.

Genus COELIOXYS Latreille

These bees are primarily cleptoparasitic in the nests of Megachile, but some species have been
reared from the nests of Chalicodoma as well as other bees including Xylocopa in India.

Revision: Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. 129 pp., 61 figs., frontis.
(subgenera of Western Hemisphere). —Baker, 1975. Kans. Univ. Sci. Bul. 50: 649-730, 49
figs., 1 table (Nearctic spp. of subg. Boreocoeliorys, Coelioxys, Schizocoelioxys,
Xerocoelioxys).

Taxonomy: Cockerell, 1905. Psyche 12: 87. —Cockerell, 1912. Canad. Ent. 44: 168.
—Crawford, 1914. Ent. Soc. Amer., Ann. 7: 148. —Sladen, 1915. Canad. Ent. 47: 205
(Ontario spp.). —Cockerell, 1921. Amer. Mus. Novitates 21: 7. —Cockerell, 1925.
Pan-Pacific Ent. 1: 150. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 192-282, figs.
1, 63-66, table 7 (eastern U. S. spp.). —Baker, 1971. Kans. Ent. Soc., Jour. 44: 225-235, 40
figs; 1 table (larval development and sexual dimorphism).

Biology: Graenicher, 1927. Ent. News 38: 231-235, 273-276 (life history)

Morphology: Pasteels and Pasteels, 1971. Acad. Sci. Paris, Compt. Rend. 273: 1580-1581
(structure of tergal plates).

Genus COELIOXYS Subgenus COELIOXYS Latreille
Coelioxys Latreille, 1809. Gen. Crust. Ins., v. 4, p. 166.

Superfamily APOIDEA 2077

Type-species: Apis quadridentata Linnaeus. Desig. by Latreille, 1810.
(=Anthophora conica Fabricius = Apis conica Linnaeus).
Coelioxys subg. Paracoelioxys Gribodo, 1884. Soc. Ent. Ital., Bol. 16: 274.
Type-species: Coeliorys montandoni Gribodo. Monotypic.
Paracoelioxys Radoszkowski, 1893. Soc. Ent. Rossica, Horae 27: 53. Preoce.
Type-species: Paracoelioxys barrei Radoszkowski. Desig. by Sandhouse, 1943.
Coelioxys subg. Liothyrapis Cockerell, 1911. U. S. Natl. Mus., Proce. 40: 246.
Type-species: Coelioxrys apicata Smith. Monotypic. (=Coelioxys decipiens Spinola).
ed ia Hemicoelioxys Pasteels, 1968. Mus. Roy. l’ Afr. Centr., Ann. Sci. Zool. (8)
167: 133.
Psa Pea Coelioxys (Hemicoelioxys) gracilis Pasteels. Monotypic and orig.
esig.
hirsutissima Cockerell. Tex. to Calif.
Coelioxys hirsutissima Cockerell, 1912. Canad. Ent. 44: 168. d.
immaculata Cockerell. Mass. to Fla., Ind., Kans., Okla., Ark., Tex.
Coelioxys immaculata Cockerell, 1912. Canad. Ent. 44: 165. 6.
Coelioxys sculptifrons Crawford, 1914. Ent. Soc. Amer., Ann. 7: 153. 2.
mitchelli Baker. N. C., Fla., Kans., Nev., Calif.
Coelioxys (Coelioxys) mitchelli Baker, 1975. Kans. Univ. Sci. Bul. 50: 726, figs. 42B, 43,
46H-I. 2, 3.
serricauda Baker. Wash., Oreg., Calif.
Coelioxys (Coelioxys) serricauda Baker, 1975. Kans. Univ. Sci. Bul. 50: 728, figs. 45B, F,
H, 46J, 49. 2, 6.
sodalis Cresson. Alaska and N. W. T., south to Calif., Ariz. and N. Mex. in the west and Great
Lakes and New England states in the east. Host: Megachile frigida Sm., M.
melanophaea Sm., M. melanophaea wootoni Ckll., M. pacifica (Panz.), M. texana Cress.
Coelioxys sodalis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 99. d.
Coelioxys ribis Cockerell, 1900. Canad. Ent. 32: 301. 9.
Coelioxys ribis var. kincaidii Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 18: 33. 2.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 226 (searching behavior, as quadridentata).
—Graenicher, 1927. Ent. News 38: 233, 273 (host, as ribis). —Graenicher, 1935. Ent. Soe.
Amer., Ann. 28: 300, 304 (life history, as ribis). —Medler and Lussenhop, 1968. Wis. Univ.
Res. Bul. 274: 58 (host). —Hobbs, 1968. Canad. Ent. 100: 781 (host).

Genus COELIOXYS Subgenus BOREOCOELIOXYS Mitchell

Coelioxys subg. Boreocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p. 37.
Type-species: Coeliorys rufitarsis Smith. Orig. desig.
banksi Crawford. Que. to N. C., west to B. C., south to N. Mex., Ariz. and Calif. Host:
Megachile relativa Cress.
Coelioxys banksi Crawford, 1914. Ent. Soc. Amer., Ann. 7: 155. °.
Coelioxys angulifera Cockerell, 1916. Pomona Col. Jour Ent. Zool. 8: 61. 2, d.

Biology: Medler and Koerber, 1958. Ent. Soc. Amer., Ann. 51: 343 (life history, host).
insita Cresson. Nebr. to Tex., west to Colo., N. Mex. and east. Ariz.
Coelioxys insita Cresson, 1872. Amer. Ent. Soc., Trans. 4: 273. °.
Coelioxys rudis Cockerell, 1934. Amer. Mus. Novitates 732: 3. 2, 6.
moesta Cresson. N. S. south to Va., west to N. W. T. and Alaska, south to Calif., Ariz., and N.
Mex. Host: Megachile centuncularis (Linn.), M. concinna Sm., M. frigida Sm., M.
pacifica (Panz.), M. relativa Cress., M. texana Cress.
Coelioxys moesta Cresson, 1864. Ent. Soc. Phila., Proc. 2: 403. ?.
Coelioxys tristis Provancher, 1882. Nat. Canad. 13: 241. °.
Coelioxys maesta(!) Dalla Torre, 1896. Cat. Hym., v. 10, p. 488.
Coelioxys lutzi Cockerell, 1921. Amer. Mus. Novitates 21: 5. °, d.

Taxonomy: Bohart, 1970. Utah State Univ. 41st Faculty Honor Lecture, p. 9 (immatures).

2078 Hymenoptera in America North of Mexico

Biology: Graenicher, 1927. Ent. News 38: 233, 274 (life history, host). —Medler and Koerber,
1958. Ent. Soe. Amer., Ann. 51: 337, 343 (life history, host). —Medler, 1959. Canad. Ent. 9:
114-115 (life history, host). —Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 54
(host). —Hobbs, 1968. Canad. Ent. 100: 781 (host).

novomexicana Cockerell. Tex. west to Calif. and Oreg.; Mexico (Chihuahua, Durango, Nuevo
Leon and Tamaulipas). Host: Megachile brevis Say, M. gentilis Cress., M. pacifica
(Panz.), M. sidalceae Ckll.
Coelioxys sayi var. novomexicana Cockerell, 1909. Ent. News 20: 9. 9.

Taxonomy: Cockerell, 1921. Amer. Mus. Novitates 21: 1. 6.

Biology: Bechtel, 1958. Pan-Pacifie Ent. 34: 12-13 (host).

octodentata Say. Transcont. U.S. and south. Canada; Mexico. Host: Megachile brevis Say, M.
centuncularis (Linn.), M. mendica Cress., M. onobrychidis Ckll., M. pacifica (Panz.), M.
perihirta Ckll., M. texana Cress.

Coelioxys 8-dentata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 353. d.
Coelioxys brevis Cresson, 1864. Ent. Soe. Phila., Proc. 2: 402. 9, ¢. Preoce.

Coelioxys altilis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 219. N. name for brevis Cresson.
Coelioxys cressoni Dalla Torre, 1896. Cat. Hym., v. 10, p. 485. N. name for brevis Cresson.
Coelioxys coquilletti Crawford, 1914. Ent. Soc. Amer., Ann. 7: 157, fig. °.

Coelioxys megatricha Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 80: 60. 3.

Coelioxys crassula Cockerell, 1919. Canad. Ent. 51: 27. 2.

Coelioxys mediata Cockerell, 1925. Pan-Pacific Ent. 1: 146. 9, d.

Coelioxys catalinica Cockerell, 1940. South. Calif. Acad. Sci., Bul. 38: 136. ¢.

Coelioxys atlantica Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 198, fig. 65. 2, d.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1060, figs. 159-161, 165, 166 (larva).
—Stephen, Bohart and Torchio, 1969. The biology and external morphology of bees, p. 29
(immatures). —Baker, 1971. Kans. Ent. Soc., Jour. 44: 225-235, figs. (immatures).

Biology: Fox, 1900. Ent. News 11: 553 (host). —Hicks, 1926. Colo. Univ., Studies 15: 227 (life
history, host). —Robertson, 1926. Psyche 33: 116 (host). —Graenicher, 1935. Ent. Soc.
Amer., Ann. 28: 300, 304 (host). —Michener, 1953. Kans. Univ. Sci. Bul. 35: 1737-1742, figs.
24-26 (life history, host, immature stages). —Linsley, 1958. Hilgardia 27: 582 (ecology).
—Medler, 1965. Ent. Soc. Wash., Proc. 67: 113-115 (life history, host). —Medler and
Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 55 (hosts).

porterae Cockerell. Newfoundland, south to N. C., west to B. C., south to Calif., Ariz. and N.
Mex. Host: Megachile frigida Sm., M. relativa Cress.
Coelioxys porterae Cockerell, 1900. Canad. Ent. 32: 298. 9.
Coelioxys dubitata var. melanopoda Viereck, 1917 (1916). Conn. State Geol. and Nat. Hist.
Survey, Bul. 22: 747. 6.
Coelioxys hypodonta Cockerell, 1925. Pan-Pacifie Ent. 1: 150. d.
pratti Crawford. Tex. (Kerrville); Mexico and Cent. Amer.
Coelioxys pratti Crawford, 1914. Ent. Soc. Amer., Ann. 7: 159. °.
rufitarsis Smith. Transcont. U. S. and south. Canada; Mexico. Host: Megachile fortis Cress., M.
latimanus Say, M. melanophaea Sm., M. m. wootoni Ckll., M. montivaga Cress., M.
perihirta Ckll., M. texana Cress.
Coelioxys rufitarsus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 271. d.
Coelioxys dubitata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 272. 2.
Coelioxys comstockii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 96. °.
Coelioxys coloradensis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 98. ¢.
Coelioxys rufitarsis rhois Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 452. °.
Coelioxys rufitarsis claripes Cockerell, 1925. Pan-Pacific Ent. 1: 146. d.

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 162-163 (host). —Graenicher, 1906.
Wis. Nat. Hist. Soc., Bul. 4: 138 (life history, host). —Robertson, 1926. Psyche 33: 116
(host). —Hicks, 1926. Colo. Univ., Studies 15: 230, 233 (as coloradensis and rufitarsis).
—Graenicher, 1927. Ent. News 38: 233 (life history, host). —Graenicher, 1935. Ent. Soe.
Amer., Ann. 28: 300, 304 (host, as dubitata). —Medler and Lussenhop, 1968. Wis. Univ.
Res. Bul. 274: 57 (life history).

Superfamily APOIDEA 2079

salinaria Cockerell. Utah. Host: Megachile brevis Say.
Coelioxys salinaria Cockerell, 1925. Pan-Pacifie Ent. 1: 148. d.
sayi Robertson. N. Y. to Fla., west to Nebr., Colo. and Ariz. Host: Megachile brevis Say, M.
mendica Cress.
Coelioxys sayi Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 346. 2, d.
Coelioxys mendacina Cockerell, 1921. Amer. Mus. Novitates 21: 3. 9, d.

Biology: Krombein, 1967. Trap-nesting wasps and bees, p. 483 (host).

Genus COELIOXYS Subgenus XEROCOELIOXYS Mitchell

Coelioxys subg. Xerocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p. 44.
Type-species: Coelioxys edita Cresson. Orig. desig.
aperta Cresson. Colo., N. Mex., Ariz.; Mexico (Jalisco).
Coelioxys aperta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 95, fig. °.

Taxonomy: Cockerell, 1921. Amer. Mus. Novitates 21: 4. ¢.

bisoncornua Hill. Nebr., Minn., Kans.
Coelioxys bisoncornua Hill, 1936. Ent. News 47: 205. @, d.

boharti Mitchell. Fla. (Cocoa and Labelle), Tex. (College Station).
Coelioxys boharti Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 206. °.

edita Cresson. Fla., Ill., Mo., Ark., and Tex., west to Alta., Wash., Oreg. and Calif.; Mexico.
Coelioxys edita Cresson, 1872. Amer. Ent. Soc., Trans. 4: 272. d.
Coelioxys deplanata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 96. °.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 160-161, fig. 7 (sleep).

galactiae Mitchell. N. C., Fla., Ill.
Coelioxys galactiae Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 204, figs. 65, 66. °,
3

grindeliae Cockerell. N. Dak. to N. Mex., west to B. C., Wash., Oreg. and Calif. Host:
Megachile perihirta Ckll.
Coelioxys grindeliae Cockerell, 1900. Canad. Ent. 32: 300. °, é.
Coelioxys grindeliae denverensis Cockerell, 1912. Canad. Ent. 44: 166. 6.
Coelioxys lamellicauda Cockerell, 1921. Amer. Mus. Novitates 21: 6. d.

Biology: Hicks, 1925. Colo. Univ., Studies 15: 237 (host).
mesae Cockerell. Colo., Utah, Idaho, Oreg. and Wash. Host: Megachile wmatillensis Mitchell.
Coelioxys mesae Cockerell, 1921. Amer. Mus. Novitates 21: 6. d.
Coelioxys flagrata Baker, 1972. In Bohart and Youssef, Royal Ent. Soc. Lond., Trans. 124:
1, 13-16, 19, figs. 2, 16-19. 9. N. syn.
Biology: Bohart and Youssef, 1972. Royal Ent. Soe. London, Trans. 124: 1, 18-16, 19, figs. 2,
16-19 (host, life history, immature stages, as flagrata).
nodis Baker. IIl., Kans. and Nebr.
Coelioxys (Xerocoelioxys) nodis Baker, 1975. Kans. Univ. Sci. Bul. 50: 716. d.
piercei Crawford. Tex. (Cotulla).
Coelioxys piercei Crawford, 1914. Ent. Soc. Amer., Ann. 7: 152, fig. 2.
soledadensis Cockerell. N. Mex., Ariz. Host: Megachile soledadensis Cockerell?
Coelioxys soledadensis cockerell, 1909. Ent. News 20: 9. d.

Genus COELIOXYS Subgenus SCHIZOCOELIOXYS Mitchell

Coelioxys subg. Schizocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p.

50.
Type-species: Coelioxys funeraria Smith. Orig. desig. and monotypic.
funeraria Smith. Alaska, Yukon and B. C., east to N. S., south to IIl., Ind. Pa. and Ga. in the

east, and south to Calif., Utah, and N. Mex. in the west. Host: Megachile frigida Sm., M.
inermis Prov., M. latimanus Say, M. pacifica (Panz.), M. relativa Cress.

Coelioxys funeraria Smith, 1854. Cat. Hym. Brit. Mus,., v. 2, p. 272. 6.

Coelioxys lateralis Cresson, 1864. Ent. Soc. Phila., Proc. 2: 405. 6.

2080 Hymenoptera in America North of Mexico

Coelioxys lucrosa Cresson, 1878. Amer. Ent. Soc., Trans. 7: 97. 2.
Coelioxys hicksi Cockerell, 1934. Amer. Mus. Novitates 732: 3. °.

Biology: Graenicher, 1905. Nat. Hist. Soc. Wis., Bul. 3: 160 (host, as lwcrosa). —Graenicher,
1927. Ent. News 38: 233 (host, life history, as lwcrosa). —Graenicher, 1935. Ent. Soe.
Amer., Ann. 28: 300, 304 (host, life history, as wcrosa). —Medler, 1958. Canad. Ent. 90:
326-327 (host, life history, as lwcrosa). —Medler and Koerber, 1958. Ent. Soc. Amer., Ann.
51: 348 (host, life history, as lucrosa). —Hobbs, 1968. Canad. Ent. 100: 783 (host).

Genus COELIOXYS Subgenus SYNOCOELIOXYS Mitchell

Coelioxys subg. Synocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p. 57.
Type-species: Coelioxys texana Cresson. Orig. desig.
alternata Say. Que. to N. C., west to B. C., Wash., Utah and Ariz. Host: Megachile pugnata
Say.
Coelioxys alternata Say, 1837. Boston Jour. Nat. Hist. 1: 401. 2, d.
Coelioxys texana vegana Cockerell, 1912. Canad. Ent. 41: 166. ¢.
Coelioxys cockerelli Crawford, 1915. Insecutor Inscitiae Menstruus 3: 108. 2.
Coelioxys wisconsinensis Cockerell, 1925. Pan-Pacific Ent. 1: 145. 9.

Biology: Medler and Lussenhop, 1968. Wis. Univ. Res. Bul. 274: 51-52 (host, life history).
apacheorum Cockerell. B. C. to Calif., east to S. Dak., Colo., and Tex.
Coelioxys apacheorum Cockerell, 1900. Canad. Ent. 32: 299. °.
Coelioxys fragariae Cockerell, 1912. Canad. Ent. 44: 167. ¢.
Coelioxys quercina Cockerell, 1912. Canad. Ent. 44: 167. ¢.
erysimi Cockerell. B. C., Idaho, Oreg., Calif., Nev. and Colo.
Coelioxys erysimi Cockerell, 1912. Canad. Ent. 44: 166. 6.
hunteri Crawford. N. Y. to Fla., west to Oreg., Utah and Ariz.; Mexico. Possibly this is the
female of C. floridana Cress.
Coeliorys hunteri Crawford, 1914. Ent. Soc. Amer., Ann. 7: 151, figs. °.
texana Cresson. Fla., Mo., Kans., Okla. and Tex., west to south. Calif.; Mexico and Cent. Amer.
Host: Megachile policaris Say.
Coelioxys texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 272. 2.
Coelioxys texana sonorensis Cockerell, 1914. Entomologist 47: 116. d.

Biology: Schwarz, 1896. Ent. Soc. Wash., Proc. 4: 24 (sleep). —Banks, 1902. N. Y. Ent. Soc.,
Jour. 10: 212 (sleep). —Robertson, 1926. Psyche 33: 177 (phenology). —Krombein, 1967.
Trap-nesting wasps and bees, pp. 483-484 (host, life history).

Genus COELIOXYS Subgenus NEOCOELIOXYS Mitchell

Coelioxys subg. Neocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p. 64.
Type-species: Coelioxys assumptionis Schrottky. Orig. desig.

menthae Cockerell. N. Mex., Ariz.; Mexico to E] Salvador.
Coelioxys menthae Cockerell, 1897. Canad. Ent. 29: 120. d.

slossoni arenicola Crawford. N. C. to Ga., west to Tex.; Mexico to El Salvador.
Coelioxys slossoni var. arenicola Crawford, 1914. Ent. Soc. Amer., Ann. 7: 156. 2, 6.

slossoni slossoni Viereck. Fla.
Coelioxrys slossoni Viereck, 1902. Canad. Ent. 34: 327. 9, ¢.

Genus COELIOXYS Subgenus MELANOCOELIOXYS Mitchell
Coelioxys subg. Melanocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p.
78.

Type-species: Coelioxys tolteca Cresson. Orig. desig.

dolichos Fox. N. C. to Fla. Host: Megachile xylocopoides Sm.
Coelioxys dolichos Fox, 1890. Ent. News 1: 107. °.

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 482-483 (host, life history).

Superfamily APOIDEA 2081

Genus COELIOXYS Subgenus HAPLOCOELIOXYS Mitchell

Coelioxys subg. Haplocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p. 85.
Type-species: Coelioxrys mexicana Cresson. Orig. desig.
mexicana Cresson. N. C. to Fla., west to Tex.; Mexico.
Coelioxys mexicana Cresson, 1878. Amer. Ent. Soe., Trans. 7: 99. 2, 3.
Coelioxys asteris Crawford, 1914. Ent. Soc. Amer., Ann. 7: 156, fig. 2.

Genus COELIOXYS Subgenus GLYPTOCOELIOXYS Mitchell
Coelioxrys subg. Glyptocoelioxys Mitchell, 1973. N. C, State Univ., Contrib. Dept. Ent. p.
92.

Type-species: Coelioxys vidua Smith. Orig. desig.

germana Cresson. Ill. to N. J., south to Fla.
Coelioxys germana Cresson, 1878. Amer. Ent. Soc., Trans. 7: 102. ?.

Genus COELIOXYS Subgenus CYRTOCOELIOXYS Mitchell

Coelioxys subg. Cyrtocoelioxys Mitchell, 1973. N. C. State Univ., Contrib. Dept. Ent. p.
106.
Type-species: Coelioxys costaricensis Cockerell. Orig. desig.
angelica Cockerell. Calif.
Coelioxys angelica Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 201. 9.
deani Cockerell. Colo., Calif.
Coelioxys deani Cockerell, 1909. Ent. News 20: 8. 3.
floridana Cresson. Ind. to Fla., west to Tex. This is possibly the male of C. hunteri Cwfd.
Coelioxys floridana Cresson, 1878. Amer. Ent. Soc., Trans. 7: 98. 6.
gilensis Cockerell. Ariz., Calif., Utah. Host: Chalicodoma subewilis (Ckll.).
Coelioxys gilensis Cockerell, 1896. N. Mex. Univ., Bul. 1: 62. 3.

Biology: Hicks, 1927. Ent. News 38: 17 (host).
gonaspis Cockerell. Calif.; Mexico (Baja California).
Coelioxys gonaspis Cockerell, 1924. Calif. Acad. Sci., Proce. (4) 12: 560. °.
modesta Smith. Que. and New England States, west to Nebr., south to Fla. and Tex. Host:
Chalicodoma campanulae wilmingtoni (Mitchell), C. georgica (Cress.), Megachile
centuncularis (Linn.), M. relativa Cress.
Coelioxys modesta Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 271. 9.

Biology: Graenicher, 1927. Ent. News 38: 233, 274 (host). —Fye, 1965. Canad. Ent. 97: 876
(host). —Krombein, 1967. Trap-nesting wasps and bees, pp. 484-486 (host, validity of
previously published host records, life history).
obtusiventris Crawford. Fla.

Coelioxys obtusiventris Crawford, 1914. Ent. Soc. Amer., Ann. 7: 150. 9.
scitula Cresson. Tex.

Coelioxys scitula Cresson, 1872. Amer. Ent. Soc., Trans. 4: 273. 2, d.

Genus COELIOXYS Subgenus UNASSIGNED

asclepiadis Cockerell. Ariz. Possibly belongs to the subgenus Cyrtocoelioxys.
Coelioxys asclepiadis Cockerell, 1925. Pan-Pacific Ent. 1: 149. ¢.

Family ANTHOPHORIDAE

This is one of the very largest, if not the largest, family of bees in the world. It is present on
all the continents, although it is neither as abundant nor as well developed in the Australian and
Oriental regions as it is in the Holarctic, Ethiopian and Neotropical regions. The family is excep-
tionally well represented in the New World by numerous species and is perhaps the most
diverse and largest assemblage of these bees in the world. The Anthophoridae contain three
subfamilies, the Nomadinae which are cleptoparasites in the nests of pollen-collecting bees, the

2082 Hymenoptera in America North of Mexico

Anthophorinae which are chiefly pollen-collecting species, and the Xylocopinae which are also
largely pollen-collecting bees. Apart from the cleptoparasites, most anthophorids make their
nests in the ground although the majority of the Xylocopinae and some others (e.g., Clisodon)
nest in wood of various sorts. While many and perhaps most of the pollen-collecting
anthophorids are clearly polylectic in their intrafloral relationships, a number of species as well
as certain groups of species (e.g., Melitomini, Peponapis, Xenoglossa, etc.) have established an
oligolectic relationship with the flora.

There are more than two dozen tribes of anthophorid bees and, with the exception of the
Palaearctic Ammobatoidini and Ancylini, all of these tribes either contain representatives in the
New World (11 tribes) or are found only here (14 tribes). Of the tribes found in the New World
only the Neotropical Caenoprosopidini, Canephorulini, Eucerinodini, Osirini, Rathymini and
Tetrapediini are not known to be present in America north of Mexico.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 233-512, figs. 67-127, tables
8-17 (eastern U. S. spp.).

SUBFAMILY NOMADINAE

This is a very large, diverse and widespread group of cleptoparasitic bees being found on all
the continents and many of the islands. It is especially well represented in the Holarctic,
Ethiopian, and Neotropical regions. Members of this subfamily are parasitic in the nests of all
families of bees, except the Megachilidae (including the Fideliinae) and the Apidae. All the spe-
cies of some genera (e.g., Epeolus, Neolarra and Triopasites) are apparently parasitic only in
the nests of certain genera of pollen-collecting bees. However, the more usual pattern is that a
particular genus (e.g., Nomada and Triepeolus) has established through its component species a
broad spectrum of host relationships involving several genera or families of host bees.

The subfamily contains nearly a dozen tribes and all except the Ammobatoidini,
Caenoprosopidini and the Osirini are endemic to or represented in the Nearctic Region.

This subfamily occupies an anomalous position within the family Anthophoridae and perhaps
would be better considered as an independent family, a status formerly accorded this group by
Linsley and Michener (1939. Amer. Ent. Soc., Trans. 65: 265-305, pls. XV-X VIII). Subsequently
it has generally been recognized and treated alternatively as a number of independent tribes as-
signable either to the subfamily Anthophorinae (e.g., Michener, 1944. Amer. Mus. Nat. Hist., Bul.
82: 270-271) or to the subfamily Nomadinae in the family Anthophoridae (e.g., Michener, 1974.
The social behavior of bees, Chapter 2, fig. 2.2). The limits of several tribes are imperfectly
known (e.g., Epeolini, Osirini, etc.) and therefore a thorough reevaluation of the position and
classification of these cleptoparasitic bees treated here as a subfamily of the Anthophoridae is
clearly indicated. Superficially at least, it appears that these bees bear the same relationship
within the superfamily Apoidea as do the Sapygidae within the superfamily Scolioidea.

Revision: Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: 265-305 (classification).
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 338-487, figs. 98-107, tables
11-13 (eastern U.S. spp.). —Rozen, 1966. Amer. Mus. Novitates 2244: 1-38, 83 figs. (larva).

TRIBE BIASTINI

Included in this tribe are two genera, Biastes which occurs in the Palaearctic Region and
Neopasites which is found only in the Nearctic Region.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 33-36, figs. 72-78 (larva).
Genus NEOPASITES Ashmead
Revision: Linsley, 1943. Amer. Ent. Soc., Trans. 69: 141 (as Gnathopasites).
Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1067, figs. 180-185 (larva).
Genus NEOPASITES Subgenus NEOPASITES Ashmead

Neopasites Ashmead, 1898. Psyche 8: 284.
Type-species: Phileremus fulviventris Cresson. Monotypic and orig. desig.
Gnathopasites Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: 272.
Type-species: Phileremus fulviventris Cresson. Orig. desig.

Superfamily APOIDEA 2083

fulviventris (Cresson). Calif. Host: Dufourea dentipes Bohart?
Phileremus fulviventris Cresson, 1873. Amer. Ent. Soc., Trans. 8: 83. ¢.
sierrae (Linsley). Calif.
Gnathopasites (Gnathopasites) sierrae Linsley, 1943. Amer. Ent. Soc., Trans. 6: 144. 2, ¢.

Genus NEOPASITES Subgenus MICROPASITES Linsley

Gnathopasites subg. Micropasites Linsley, 1942. Pan-Pacifie Ent. 18: 130.
Type-species: Neopasites cressoni Crawford. Monotypic and orig. desig.

An undescribed species of this subgenus has been reared from Dufowrea (Halictoides)
trochantera Bohart.

cressoni Crawford. Calif., Ariz. Host: Dufouwrea mulleri (CkIl.).
Neopasites cressoni Crawford, 1916. Insecutor Inscitiae Menstruus 4: 136. d.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 33-36, figs. 72-83 (larva).
Biology: Torchio, Rozen, Bohart and Favreau, 1967. N. Y. Ent. Soc., Jour. 75: 143-145, figs.
7-9 (adult activity, oviposition, egg, host).

mojavensis (Linsley). Calif.
Gnathopasites (Micropasites) mojavensis Linsley, 1943. Amer. Ent. Soc., Trans. 69: 148. 9.

timberlakei (Linsley). Calif.
Gnathopasites (Micropasites) timberlakei Linsley, 1943. Amer. Ent. Soc., Trans. 69: 147. °,

3
TRIBE TOWNSENDIELLINI

Townsendiella is the only genus included in this tribe and, so far as known, the species are
cleptoparasites in the nests of Conanthalictus and Hesperapis.

Genus TOWNSENDIELLA Crawford
Revision: Linsley, 1943. Amer. Ent. Soc., Trans. 69: 93 (included spp.).
Genus TOWNSENDIELLA Subgenus TOWNSENDIELLA Crawford

Townsendiella Crawford, 1916. Insecutor Inscitiae Menstruus 4: 138.
Type-species: Townsendiella pulchra Crawford. Monotypic.

pulchra Crawford. N. Mex., south. Calif.; Mexico (Baja California).
Townsendiella pulchra Crawford, 1916. Insecutor Inscitiae Menstruus 4: 138. °.

Genus TOWNSENDIELLA Subgenus XEROPASITES Linsley

Townsendiella subg. Xeropasites Linsley, 1942. Pan-Pacific Ent. 18: 130.
Type-species: Townsendiella (Xeropasites) rufiventris Linsley. Monotypie and orig.
desig.

rufiventris Linsley. South. Calif.
Townsendiella (Xeropasites) rufiventris Linsley, 1942. Pan-Pacific Ent. 18: 130. 2, d.

Genus TOWNSENDIELLA Subgenus EREMOPASITES Linsley
Townsendiella subg. Eremopasites Linsley, 1942. Pan-Pacifie Ent. 18: 131.
Type-species: Townsendiella californica Michener. Monotypic and orig. desig.
californica Michener. South. Calif. Host: Hesperapis rufipes (Ashm.).
Townsendiella californica Michener, 1936. Ent. News 47: 181. 2, 6.
TRIBE NEOLARRINI
This tribe contains only the Nearctic genus Neolarra whose species are parasitic in the nests
of the andrenid genus Perdita and possibly also Nomadopsis.
Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 36-37, figs. 79-83 (larva).

2084 Hymenoptera in America North of Mexico

Genus NEOLARRA Ashmead
Revision: Michener, 1939. Amer. Ent. Soc., Trans. 65: 347-362.
Genus NEOLARRA Subgenus NEOLARRA Ashmead

Neolarra Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 8.
Type-species: Neolarra pruinosa Ashmead. Monotypic.

abdominalis Michener. Mont.

Neolarra (Neolarra) abdominalis Michener, 1939. Amer. Ent. Soc., Trans. 65: 352. 2, 6.
alba Cockerell. South. Calif.

Neolarra alba Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 285. 6.
californica Michener. South. Calif.

Neolarra (Neolarra) californica Michener, 1939. Amer. Ent. Soc., Trans. 65: 356. ?, 3.
congregata Crawford. Tex., N. Mex.

Neolarra congregata Crawford, 1907. N. Y. Ent. Soc., Jour. 15: 181. 2, 6.
helianthi Cockerell. Colo.

Neolarra congregata helianthi Cockerell, 1936. Amer. Mus. Novitates 831: 5. 6.
linsleyi Michener. South. Calif.

Neolarra (Neolarra) linsleyi Michener, 1939. Amer. Ent. Soc., Trans. 65: 357. 2, d.
pruinosa Ashmead. Alta. to Colo., N. Mex. and Calif. Host: Perdita (Cockerellula) opuntiae

Ckll., P. (Perdita) zebrata zebrata Cress.
Neolarra pruinosa Ashmead, 1890. Colo. Biol. Assoc., Bul. 1: 8. °.
Neolarra vittata Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 285. 6.
Taxonomy: Baker, 1896. Ent. Soc. Wash., Proc. 4: 23. 2, ¢. —Rozen, 1966. Amer. Mus.
Novitates 2244: 36-37 (larva).
Biology: Rozen, 1965. N. Y. Ent. Soc., Jour. 73: 87-88 (adult activity, host).

vandykei Michener. Calif. (Oakley).

Neolarra (Neolarra) vandykei Michener, 1939. Amer. Ent. Soc., Trans. 65: 3538. 2, ¢.

verbesinae (Cockerell). Nebr. to N. Mex.
Phileremus verbesinae Cockerell, 1895. Psyche 7 (sup.): 10. 2, 6.

Genus NEOLARRA Subgenus PHILEREMULUS Cockerell

Phileremulus Cockerell, 1895. Psyche 7 (sup.): 9.
Type-species: Phileremulus vigilans Cockerell. Monotypic and orig. desig.

cockerelli (Crawford). Tex., Tenn., Ga. Host: Perdita (Alloperdita) obscurata Cress.?
Phileremulus cockerelli Crawford, 1916. Insecutor Inscitiae Menstruus 4: 139. 9.
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 490 (redescription, possible
parasite).

coloradensis Michener. Colo.
Neolarra (Phileremulus) coloradensis Michener, 1939. Amer. Ent. Soc., Trans. 65: 352. °,

3

mallochi (Crawford). Alta.
Phileremulus mallochi Crawford, 1912. Canad. Ent. 44: 359. 6.

vigilans (Cockerell). N. Mex., south. Calif. Host: Perdita (Perdita) indioensis Timb.
Phileremulus vigilans Cockerell, 1895. Psyche 7 (sup.): 9. 6, 9.
Phileremulus nanus Cockerell, 1895. Psyche 7 (sup.): 9. °.

TRIBE HOLCOPASITINI

This tribe contains only two genera, Holcopasites from the Nearctic Region and
Schmiedeknechtia of the Palaearctic Region.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 30-33, figs. 66-71 (larva).

Superfamily APOIDEA 2085
Genus HOLCOPASITES Ashmead

Insofar as known the species of this genus are cleptoparasites in the nests of pollen-collecting
bees belonging to the subfamily Panurginae of the family Andrenidae. Although a definite host
association has been established with the genera Calliopsis, Heterosarus, Hypomacrotera, Pseu-
dopanurgus and Pterosarus, thus far none of the other genera of the formerly recognized tribe
Panurgini (Nomadopsis, Panurginus, Perdita, Protandrena, ete.) which occur within the geo-
graphic range of Holcopasites has been implicated as a possible host.

Revision: Linsley, 1943. Amer. Ent. Soc., Trans. 69: 119 (as Neopasites). —Hurd and
Linsley, 1972. Smithsn. Contrib. Zool. 114: 1-41, 16 figs., 1 table (Nearctic spp., including
biology).

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 30-33, figs. 66-71 (larva).

Biology: Rozen, 1965. N. Y. Ent. Soc., Jour. 73: 88-91, figs. 1-4 (life history). —Shinn, 1967.
Kans. Univ. Sci. Bul. 46: 928-932. —Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114: 2,
table 1 (host associations, summary of literature).

Genus HOLCOPASITES Subgenus TRICHOPASITES Linsley

Neopasites subg. Trichopasites Linsley, 1942. Pan-Pacifie Ent. 18: 127.
Type-species: Neopasites (Trichopasites) insoletus Linsley. Monotypic and orig.
desig.
Neopasites subg. Odontopasites Linsley, 1942. Pan-Pacific Ent. 18: 128.
Type-species: Neopasites (Odontopasites) arizonicus Linsley. Monotypic and orig.
desig.
arizonicus (Linsley). Idaho, Utah, Colo., Ariz., N. Mex., south into Mexico (Baja California and
Zacatecas). Host: Calliopsis coloradensis Cress., C. pectidis Shinn?, Pterosarus occidus
(Timb.)?, P. timberlake Ckll.?
Neopasites (Odontopasites) arizonicus Linsley, 1942. Pan-Pacific Ent. 18: 129. 9, 6.
insoletus (Linsley). Ariz., N. Mex. Host: Pterosarus boylei (CkIl.)?, P. perlaevis (Ckll.)?, P.
timberlakei (Ckll.)?
Neopasites (Trichopasites) insoletus Linsley, 1942. Pan-Pacific Ent. 18: 128. 2.
rozeni Hurd and Linsley. Ariz.; Mexico (Sinaloa and Sonora).
ieee (Tricho pasites) rozeni Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114:
10N2R

Genus HOLCOPASITES Subgenus HOLCOPASITES Ashmead

Holcopasites Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 82.
Type-species: Phileremus illinoiensis Robertson. Desig. by Crawford, 1915.

SPOTTED SPECIES GROUP

apacheorum Hurd and Linsley. Ariz., N. Mex.
Holcopasites (Holcopasites) apachgorum Hurd and Linsley, 1972. Smithsn. Contrib. Zool.
114: 14. 9.
bohartorum Hurd and Linsley. Ariz., Calif. (Blythe).
Holcopasites (Holcopasites) bohartorum Hurd and Linsley, 1972. Smithsn. Contrib. Zool.
114: 18, fig. 8. 2, d.
calliopsidis calliopsidis (Linsley). Mont. to N. Y., south to Tenn., Tex. and Ariz.; Mexico
(Zacatecas). Host: Calliopsis andreniformis Sm., Pseudopanurgus sp.?
Neopasites calliopsidis Linsley, 1943. Amer. Ent. Soc., Trans. 69: 187. 2, d.
Holcopasites pseudocarinatus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 489. 2.

Biology: Ainslie, 1937. Canad. Ent. 69: 99 (as stevensi). —Shinn, 1967. Kans. Univ. Sci. Bul.
46: 928-932 (life history).

calliopsidis carinatus (Linsley). South. Tex.; Mexico (Hidalgo, San Luis Potosi and Veracruz).
Neopasites carinatus Linsley, 1943. Amer. Ent. Soc., Trans. 69: 127. °.

2086 Hymenoptera in America North of Mexico

heliopsis (Robertson). Alta., Mont., Colo. and IIl., south to Ark. and Kans. Host: Calliopsis
nebraskensis Cwfd.?
Ammobates heliopsis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 350. d.
Holcopasites lutzi Cockerell, 1934. Amer. Mus. Novitates 697: 12. 6.
pulchellus (Cresson). Alta., Sask. to N. Dak., south to Tex., N. Mex. and Ariz.; Mexico
(Durango, Michoacan and Zacatecas). Host: Pseudopanurgus sp.?
Phileremus pulchellus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 84. 2, 6.
Neopasites robertsoni Crawford, 1906. Canad. Ent. 38: 283. 2, ¢.
Neopasites robertsoni pubescens Linsley, 1943. Amer. Ent. Soc., Trans. 69: 131. 6.
tegularis Hurd and Linsley. Ariz. Host: Heterosarus nanulus (Timb.).
Holcopasites (Holcopasites) tegularis Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114:
2A ORIG:

BANDED SPECIES GROUP

bigibbosus Hurd and Linsley. Ariz. (Willcox); N. Mex. (Rodeo).
Holcopasites (Holcopasites) bigibbosus Hurd and Linsley, 1972. Smithsn. Contrib. Zool.
M428) figs lla on
cazieri Hurd and Linsley. Ariz., N. Mex.
Holcopasites (Holcopasites) cazieri Hurd and Linsley, 1972. Smithsn. Contrib. Zool. 114:
29, figs. 4, 16. 2, d.
eamia (Cockerell). Tex., Okla.
Neopasites eamia Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 4: 29. ¢.
Holcopasites acanthochilus Crawford, 1915. Insecutor Inscitiae Menstruus 3: 125. °.
Holcopasites texanus Crawford, 1915. Insecutor Inscitiae Menstruus 3: 126. d.
haematurus Cockerell and Hicks. Iowa, Nebr., Kans., Colo.
Holcopasites haematurus Cockerell and Hicks, 1926. Ent. News 37: 107. d.
illinoiensis illinoiensis (Robertson). Ill. to Mass., south to Ga., La. and Tex. Host: Calliopsis
andreniformis Sm.
Phileremus illinoiensis Robertson, 1891. Amer. Ent. Soc., Trans. 18: 64. 2.
Holcopasites pratti Ashmead In Crawford, 1915. Insecutor Inscitiae Menstruus 3: 123. °.
Neopasites punctulatus Linsley, 1943. Amer. Ent. Soc., Trans. 69: 135. 6.
illinoiensis minimus (Linsley). Ariz., N. Mex.; Mexico (Sonora). Host: Hypomacrotera callops
persimilis Ckll., H. callops callops Ckll.?
Neopasites minimus Linsley, 1943. Amer. Ent. Soc., Trans. 69: 126. °.
stevensi Crawford. Alta. N. Dak., Nebr., Tex., N. Mex., Ariz. Host: Calliopsis crypta Shinn, C.
rozent Shinn?
Holcopasites stevensi Crawford, 1915. Insecutor Inscitiae Menstruus 3: 125. 9, ¢.
Neopasites elegans Linsley, 1944. N. Y. Ent. Soe., Jour. 52: 277. 9.
Neopasites knulli Linsley, 1944. N. Y. Ent. Soc., Jour. 52: 278. 2.

TRIBE AMMOBATINI

Although this tribe is represented in the Palaearctic and Ethiopian regions by several genera,
only the Nearctic genus Oreopasites is present in the New World and its species are apparently
all cleptoparasitic in the nests of the andrenid genera Nomadopsis and Hypomacrotera.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 28-30, figs. 56-60 (larvae). —Rozen,
1974. Amer. Mus. Novitates 2551: 1-16, 33 figs. (larvae, pupae).
Genus OREOPASITES Cockerell
Oreopasites Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 442.
Type-species: Oreopasites scituli Cockerell. Monotypic.
Revision: Linsley, 1941. Amer. Ent. Soc., Trans. 66: 307.

Taxonomy: Rozen, 1954. Pan-Pacifie Ent. 30: 203-207, 6 figs. (larva). —Rozen, 1966. Amer.
Mus. Novitates 2244: 28-30, figs. 56-58. —Rozen, 1974. Amer. Mus. Novitates 2551: 12-15,
figs. 26-28 (larvae, pupae).

Superfamily APOIDEA 2087

albinota Linsley. Calif.
Oreopasites albinota Linsley, 1941. Amer. Ent. Soc., Trans. 66: 313. 2.
arizonica Linsley. Ariz.
Oreopasites arizonica Linsley, 1941. Amer. Ent. Soc., Trans. 66: 310. 9.
euphorbiae Cockerell. South. Calif. Host: Nomadopsis euphorbiae (Ckll.).
Oreopasites euphorbiae Cockerell, 1929. Pan-Pacific Ent. 5: 105. 2, d.
scituli Cockerell. Colo. Host: Nomadopsis scitula (Cress.).
Oreopasites scituli Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 442. 2, 6.
vanduzeei diabloensis Linsley. North. Calif. Host: Nomadopsis sp.
Oreopasites vanduzeei diabloensis Linsley, 1941. Amer. Ent. Soc., Trans. 66: 315. 2, d.
vanduzeei melanantha Linsley. North. Calif. (Sierra Nevada). Host: Nomadopsis anthidia
(Fowler).
Oreopasites vanduzeei melanantha Linsley, 1941. Amer. Ent. Soc., Trans. 66: 316. 2, d.
vanduzeei vanduzeei Cockerell. North. Calif. (Cent. Valley). Host: Nomadopsis equina (CkIl.).
Oreopasites vanduzeei Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 202. 9.
vanduzeei vernalis Linsley. South Calif. Host: Nomadopsis sp.
Oreopasites vanduzeei vernalis Linsley, 1941. Amer. Ent. Soc., Trans. 66: 317. °, d.
vanduzeei xerophila Linsley. Ariz., South. Calif. (deserts). Host: Nomadopsis puellae (CkIl.).
Oreopasites vanduzeei xerophila Linsley, 1941. Amer. Ent. Soc., Trans. 66: 318. 2, d.

TRIBE PROTEPEOLINI

This is a small group found only in the Americas. In addition to the genus Protepeolus it con-
tains Isepeolus and possibly one other Neotropical genus.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 10-12, figs. 2-7 (larva).

Genus PROTEPEOLUS Linsley and Michener

Protepeolus Linsley and Michener, 1937. Pan-Pacifie Ent. 13: 75.
Type-species: Protepeolus singularis Linsley and Michener. Monotypic and orig.
desig.
integer Linsley. South Calif. to west. Tex.; Mexico (Durango and Jalisco). Host: Diadasia
ochracea (CkIl.)?
Protepeolus integer Linsley, 1939. Pan-Pacific Ent. 15: 4. d.

Taxonomy: Hurd and Linsley, 1963. Kans. Ent. Soc., Jour. 36: 253-255. 3, 2 (possible
parasite).

singularis Linsley and Michener. N. Mex. Host: Diadasia ochracea (Ckll.).
Protepeolus singularis Linsley and Michener, 1937. Pan-Pacific Ent. 13: 76. 2.

Biology: Eickwort, Eickwort and Linsley, 1977. Kans. Ent. Soc., Jour. 50: 2, 9 (host, as
Diadasia olivacea).

TRIBE EPEOLOIDINI

Although there is a South American species attributed to the genus Epeoloides, the tribe is
apparently represented only in the Holaretic Region by the genus E’peoloides. It is believed that
the species are cleptoparasitic in the nests of the melittid genus Macropis.

Genus EPEOLOIDES Giraud

Epeoloides Giraud, 1863. Zool.-Bot. Gesell, Wien., Verh. 13: 45.
Type-species: Apis coecutiens Fabricius. Monotypic. (~Epeoloides ambiguus
Giraud).
Viereckella Swenk, 1907. Ent. News 18: 298.
Type-species: Viereckella obscura Swenk. Orig. desig.
obscura (Swenk). Nebr.
Viereckella obscura Swenk, 1907. Ent. News 18: 299. °.

2088 Hymenoptera in America North of Mexico

pilosula (Cresson). Canada (Cap. Rouge), New England to Ga., west to Wis. and N. Dak.
Nomada pilosula Cresson, 1878. Amer. Ent. Soc., Trans. 7: 77. d.
Nomia compacta Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 337. d.
Epeolus pilosulus Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 426. 2.
Viereckella ceanothina Cockerell, 1907. Ent. News 18: 300. 2.
Epeoloides nearcticus Ducke, 1909. Rev. Ent. Caen 27: 39. °.

Taxonomy: Crawford, 1917. Ent. Soc. Wash., Proc. 19: 167. —Mitchell, 1962. N. C. Agr. Expt.
Sta. Tech. Bul. 152: 491-492, figs. 115, 116 (redescription, synonymy).

TRIBE EPEOLINI

The members of this tribe share many features with the Nomadini and perhaps, as has been
suggested, they should be placed together in the Nomadini. With the exception of the primarily
Holarctic genus Epeolus and a few Old World species of the predominantly New World genus
Triepeolus, the epeoline bees are centered in the Americas and are especially well represented
in the Neotropical Region by several genera. Only the genera Epeolus and Triepeolus are
present in America north of Mexico, but it appears that a few of our species belong to the genus
Trophocleptria.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 12-22, figs. 8-38 (larva).

Genus EPEOLUS Latreille

Epeolus Latreille, 1802. Hist. Nat. Fourmis, p. 427.

Type-species: Apis variegata Linnaeus. Monotypic. (~Nomada variegata Fabricius).
Pyrrhomelecta Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 66.

Type-species: Epeolus glabratus Cresson. Monotypic and orig. desig.
Argyroselenis Robertson, 1903. Canad. Ent. 35: 284.

Type-species: Triepeolus minimus Robertson. Monotypic and orig. desig.

Insofar as known, the species of this chiefly Holarctic genus are cleptoparasitic in the nests of
the genus Colletes.

Taxonomy: Cockerell, 1928. Colo. Univ., Studies 16: 105 (key). —Michener, 1953. Kans. Univ.
Sci. Bul. 35: 1072, figs. 195, 196, 198 (larva). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech.
Bul. 152: 438-458, figs. 108-110 (eastern U.S. spp.). —Rozen, 1966. Amer. Mus. Novitates
2244: 19-20, figs. 25-32 (larva).

ainsliei Crawford. Iowa.
Epeolus ainsliei Crawford, 1932. Ent. Soc. Wash., Proc. 34: 74. 9.

americanus (Cresson). Canada; Colo., Calif.
Phileremus americanus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 83. ¢, 2.
Taxonomy: Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 33. —Linsley, 1939. Pan-Pacific
Ent. 15: 1.

arciferus Cockerell. Calif.
Epeolus arciferus Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 13: 319. 9.

asperatus Cockerell. Calif.
Epeolus asperatus Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 5: 25. @.
Taxonomy: Cockerell, 1934. Amer. Mus. Novitates 697: 12. °.

australis Mitchell. N. C., Ga.
Epeolus australis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 441, fig. 110. 2, ¢.

autumnalis Robertson. Minn. to Maine, south to N. C. Host: Colletes compactus Cress.?
Epeolus autumnalis Robertson, 1902. Ent. News 13: 81. 2.
Taxonomy: Robertson, 1903. Canad. Ent. 35: 287, 288. °.
banksi (Cockerell). Minn. to N. J. and N. C.
Triepeolus banksi Cockerell, 1907. Entomologist 40: 135. d.
barberiellus Cockerell. N. Mex.
Epeolus barberiellus Cockerell, 1907. Entomologist 40: 266. °.

Superfamily APOIDEA 2089

beulahensis Cockerell. N. Mex., Colo., Iowa.
Epeolus beulahensis Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 40. 9.
bifasciatus bifasciatus Cresson. New England to Fla., west to Minn., Colo., Tex.; Mexico
(northern). Host: Colletes latitarsis Robertson? Another subspecies occurs in Mexico
and Panama.
Epeolus bifasciatus Cresson, 1864. Ent. Soc. Phila., Proc. 3: 38. 6.

Taxonomy: Robertson, 1903. Canad. Ent. 33: 287, 288. 2, 3 (key).

canadensis Mitchell. N. S., Que., N. Y., Ont., Mich., Ind.

Epeolus canadensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 444, fig. 110. 2,

é.

carolinus Mitchell. N. C., Fla.

Epeolus carolinus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 445, fig. 110. 2, ¢.
compactus Cresson. Tex. to Colo., Nev., Calif.

Epeolus compactus Cresson, 1878. Amer. Ent. Soc., Trans. 7. 89. 2, 3.

Taxonomy: Brues, 1903. Ent. News 14: 79 (key).
crucis Cockerell. N. Mex.
Epeolus crucis Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 39. ?.
eastwoodae Cockerell. Calif. (San Miguel Isl.).
Epeolus eastwoodae Cockerell, 1937. Pan-Pacific Ent. 13: 149. 3.
erigeronis Mitchell. N. C., Ga., Fla.
Epeolus erigeronis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 445, fig. 110. 2, 3.
floridensis Mitchell. Fla.
Epeolus floridensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 446, fig. 110. 2, ¢.
fumipennis Say. Tex., north. Mex.
Epeolus fumipennis Say, 1837. Boston Jour. Nat. Hist. 1: 403.
gabrielis Cockerell. Calif.
Epeolus gabrielis Cockerell, 1909. Ann. and Mag. Nat. Hist. (8) 5: 26. ¢.
Epeolus geminatus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 315. 2,
é.

glabratus Cresson. Ga., Fla.

Epeolus glabratus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 90. 6.
hitei Cockerell. Colo.

Epeolus hitei Cockerell, 1908. Entomologist 41: 60. °.
howardi Mitchell. N. C. (Southern Pines). Host: Colletes howardi Sw.?

Epeolus howardi Mitchell, 1962. N. C. Agr. Expt. St. Tech. Bul. 152: 447, fig. 110. 2, 3.
humillimus Cockerell. Wash., Oreg.

Epeolus humillimus Cockerell, 1918. Ann. and Mag. Nat. Hist. 9: 160. 3.
ilicis Mitchell. R. I. and Mass. to Ga., Tenn.

Epeolus ilicis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 448, fig. 110. 2, 6.
interruptus Robertson. N. J. to Ga., west to Colo., Calif. and Tex. Host: Colletes aestivalis

Patton?
Epeolus interruptus Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 55. 2.

Taxonomy: Robertson, 1903. Canad. Ent. 33: 287, 288. 2, d (key).
lanhami Mitchell. Maine, Pa., Mich.
E'peolus lanhami Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 450, fig. 110. 2, 6.
lectiformis Cockerell. Colo.
Epeolus lectiformis Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 623. 9.
lectoides Robertson. Ill. to New England, south to Ga.
Epeolus lectoides Robertson, 1901. Canad. Ent. 33: 231. °@.
Epeolus semilectus Cockerell, 1907. Entomologist 40: 136. ¢.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 287, 288. 2°, d (key).

lectus Cresson. Kans., S. Dak.
Epeolus lectus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 88. 2.

2090 Hymenoptera in America North of Mexico

Epeolus agnathus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 89. ¢.
lutzi Cockerell. Colo., Utah, Alta.
E peolus lutzi Cockerell, 1921. Amer. Mus. Novitates 23: 16. d, @.
Epeolus lutzi dimissus Cockerell, 1921. Amer. Mus. Novitates 23: 16. °.
melectimimus Cockerell and Sandhouse. Calif.
Epeolus melectimimus Cockerell and Sandhouse, 1924. Calif. Acad. Sei., Proc. (4) 13: 317.
Se
mercatus Fabricius. “Carolina.”
Epeolus mercatus Fabricius, 1804. Systema Piezatorum, p. 389.
mesillae mesillae (Cockerell). N. Mex. to south. Calif. Host: Colletes clypeonitens Swenk?
Phileremus mesillae Cockerell, 1895. Psyche 7 (sup.): 10. ¢.

Taxonomy: Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 138: 42. 2. —Hurd and Linsley, 1975.
Smithsn. Contrib. Zool. 193:40 (Larrea visitation).
mesillae palmarum Linsley. South. Calif. Host: Colletes clypeonitens Swenk?
Epeolus mesillae palmarum Linsley, 1939. Pan-Pacific Ent. 15: 2. 9.
minimus (Robertson). IIl., Mich., Wis., west to Colo. and Calif. Host: Colletes eulophi
Robertson.
Triepeolus minimus Robertson, 1902. Ent. News 13: 81.

Biology: Graenicher, 1906. Wis. Nat. Hist. Soc., Bul. 4: 135.
montanus (Cresson). Nev.
Phileremus montanus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 83. 6.
novomexicanus Cockerell. N. Mex.
Epeolus novomexicanus Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 10: 487. ¢.
olympiellus Cockerell. Wash.
Epeolus olympiellus Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 41. ¢.
oswegoensis Mitchell. N. Y. (Oswego).
Epeolus oswegoensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 453. 6.
pilatei Cockerell. Calif.
Epeolus pilatei Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 13: 320. 2.
piscatoris Cockerell. South. Calif. Is.
Epeolus piscatoris Cockerell, 1939. Calif. Acad. Sci., Proc. (4) 23: 482. 2.
pusillus Cresson. Maine to Fla., west to Tex., Calif., Utah, Colo. and Wyo. Host: Colletes
americanus Cresson?, C. ciliatoides Stephen, C. compactus compactus Cress., C.
deserticola Timb.
Epeolus pusillus Cresson, 1864. Ent. Soc. Phila., Proc. 2: 393. °.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 287, 288. 2, d (key). —Brues, 1903. Ent. News
14: 80, 82 (key). —Rozen, 1966. Amer. Mus. Novitates 2244: 19-20, figs. 28-32 (larva).
rubrostictus Cockerell and Sandhouse. Calif.
Epeolus rubrostictus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 318. @.
rufomaculatus Cockerell and Sandhouse. Calif., Utah, Colo.
Epeolus rufomaculatus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 314.
ETOr
rufulus Cockerell. Colo.
Epeolus rufulus Cockerell, 1941. Canad. Ent. 73: 36. 9.
scelestus Cresson. Tex.
Epeolus scelestus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 86. °.
Epeolus scelestus var. tubercularis Brues, 1903. Ent. News 14: 79. 9.
scutellaris Say. N. S. to N. C. and Fla., west to Minn. and Tex.
Epeolus scutellaris Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 355. 9.
Epeolus vernoniae Cockerell, 1907. Entomologist 40: 136. ¢.

Taxonomy: Cresson, 1864. Ent. Soc. Phila., Proc. 2: 397. 2. —Robertson, 1897. Acad. Sci. St.
Louis, Trans. 7: 343. @, 6.

Superfamily APOIDEA 2091

tristicolor Viereck. B. C.

Epeolus tristicolor Viereck, 1905. Canad. Ent. 37: 280. °.
vernalis Mitchell. N. C. (Holly Shelter).

Epeolus vernalis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 455. 9.
weemsi Mitchell. Fla. (Alachua Co.).

Epeolus weemsi Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 455. d.
zonatus Smith. Fla.

Epeolus zonatus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 257. ¢, @.

Taxonomy: Cresson, 1864. Ent. Soc. Phila., Proc. 2: 397. 2, 6.

Genus TRIEPEOLUS Robertson

Triepeolus Robertson, 1901. Canad. Ent. 33: 231.
Type-species: Epeolus concavus Cresson. Orig. desig.

Triepeolus subg. Synepeolus Cockerell, 1921. Amer. Mus. Novitates 23: 6.
Type-species: Triepeolus (Synepeolus) insolitus Cockerell. Monotypic.

While most of the known host information suggests that these bees are cleptoparasites in the
nests of eucerine bees (e.g., Melissodes, Peponapis, Svastra, and Xenoglossa), at least some spe-
cies are known to be parasites of other Anthophoridae (e.g., Anthophora, Centris) as well as Col-
letidae (Ptiloglossa), Oxaeidae (Protoxaea) and Halictidae (Nomia).

Taxonomy: Cockerell, 1928. Colo. Univ., Studies 16: 107 (key). —Michener, 1953. Kans. Univ.
Sci. Bul. 35: 1070, figs. 190-194, 197 (larva). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech.
Bul. 152: 459-485, fig. 111 (eastern U.S. spp.). —Rozen, 1966. Amer. Mus. Novitates 2244:
12-19, figs. 15-24 (larva).
agaricifer Cockerell. N. Mex.

Triepeolus agaricifer Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 60. d.
alachuensis Mitchell. Fla. (Alachua Co.).

Triepeolus alachuensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 462. 2.
alpestris Cockerell. Colo.

Triepeolus alpestris Cockerell, 1921. Amer. Mus. Novitates 21: 13. °.
amandus Cockerell. Colo.

Triepeolus amandus Cockerell, 1921. Amer. Mus. Novitates 23: 10. d.
ancoratus Cockerell. Calif.

Triepeolus ancoratus Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 63. °.
argyreus (Cockerell). Wash., Oreg., Calif..

Epeolus argyreus Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 60. ¢.
balteatus Cockerell. Colo.

Triepeolus balteatus Cockerell, 1921. Amer. Mus. Novitates 23: 5. d.
bardus (Cresson). Tex. to Minn. Host: Nomia sp.?

Epeolus bardus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 84. °.

Taxonomy: Brues, 1903. Ent. News 14: 80 (key). —Cockerell, 1935. Amer. Mus. Novitates
766: 7. 2, 3 (possible host).
bihamatus (Cockerell). Wash.
Epeolus bihamatus Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 61. ¢.
blaisdelli Cockerell and Sandhouse. Calif. (Mokelumne).
Triepeolus blaisdelli Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 310. d.
brittaini Cockerell. N.S.
Triepeolus brittaini Cockerell, 1931. Canad. Ent. 63: 297. 6.
brunnescens Cockerell and Sandhouse. Calif.
Triepeolus brunnescens Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 313.
3.
brunneus Cockerell. Colo.
Triepeolus brunneus Cockerell, 1921. Amer. Mus. Novitates 23: 7. 9.

2092 Hymenoptera in America North of Mexico

californicus (Cresson). Calif.
Epeolus californicus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 86. °.

callopus Cockerell. Calif.
Triepeolus callopus Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 202. °.
Taxonomy: Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 63-64. d, 2.

charlottensis Mitchell. N. B. (Charlotte Co.).
Triepeolus charlottensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 462, fig. 112.
oO

cirsianus Mitchell. Ind. (Warren Co.).
Triepeolus cirsianus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 463. ¢.

concavus (Cresson). N. C. and Fla., west to IIl., Wis., Colo. and Calif. Host: Svastra obliqua
obliqua (Say). ;
Epeolus concavus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 85. 2.
Taxonomy: Brues, 1903. Ent. News 14: 81, fig. 2, ¢. —Robertson, 1903. Canad. Ent. 33: 284,
285. 2, 3 (key). —Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: pls. 15, 18.
Biology: Custer, 1928. Canad. Ent. 60: 28. —Custer, 1929. Psyche 36: 293.

coquilletti Cockerell. Calif., Miss.
Triepeolus coquilletti Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 106. °.

cressonii cressonii (Robertson). Minn. to New England, south to Tenn. and N. C.
Epeolus cressonii Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 344. d, 9.

cressonii fraserae Cockerell. N. Mex.
Triepeolus cressoni var. fraserae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 39. 6.

custeri Cockerell. Colo.
Triepeolus custeri Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 17: 306. d.

cyclurus Cockerell. Colo.
Triepeolus cyclurus Cockerell, 1923. Ent. News 34: 49. 2.

dacotensis (Stevens). N. Dak., Utah. Parasite: Anthrax limatulus Say.
Epeolus dacotensis Stevens, 1919. Canad. Ent. 51: 210. 2, d.
Biology: Bohart, 1970. The Evolution of Parasitism among bees. Utah State Univ. Ann.
Honor lecture p. 22, fig. 20, 22, 24, 27 (egg, larva, parasite).

denverensis Cockerell. Colo.
Triepeolus denverensis Cockerell, 1910. Entomologist, 43:91. 3.

dichropus Cockerell. Colo.
Triepeolus dichropus Cockerell, 1921. Amer. Mus. Novitates 23: 11. 6.

distinctus (Cresson). Ga., Fla.
Epeolus distinctus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 84. d, 9.

diversipes Cockerell. Calif.
Triepeolus diversipes Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 13: 314. d.

donatus (Smith). Wis. to New England and South. Canada, south to Ga. Host: Melitoma taurea
Say.
Epeolus donatus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 256. 2, d.
Taxonomy: Robertson, 1903. Canad. Ent. 35: 285, 286. °, d (key).

Biology: Lutz and Cockerell, 1920. Amer. Mus. Nat. Hist., Bul. 42: 592 (host).

eldoradensis (Cockerell). Colo.
Epeolus eldoradensis Cockerell, 1910. Psyche 17: 245. d.

eldredi Cockerell. Wash., Utah, Wyo. Host: Melissodes rustica (Say).
Triepeolus eldredi Cockerell, 1907. Canad. Ent. 39: 52. d.
Biology: Clement, 1973. Kans. Ent. Soc., Jour. 46: 516-525 (host).
floridanus Mitchell. Fla. (Gainesville).
Triepeolus floridanus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 468. 6.

fortis Cockerell. Colo., Calif.
Triepeolus fortis Cockerell, 1921. Amer. Mus. Novitates 23: 3. 2, d.

Superfamily APOIDEA 2093

fraseri Cockerell. N. Mex.
Triepeolus fraseri Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 39. 6.

Taxonomy: Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 62. ¢ (key).
georgicus Mitchell. Ga. (Fort Gordon).
Triepeolus georgicus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 469, fig. 112. @.
grindeliae Cockerell. Colo.
Triepeolus grindeliae Cockerell, 1907. Canad. Ent. 39: 51. °.
haematurus Cockerell and Sandhouse. Utah.
Triepeolus haematurus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 311.
OF
helianthi arizonensis Cockerell. Ariz.
Triepeolus helianthi var. arizonensis Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 39.
3.
helianthi grandior Cockerell. Colo., N. Mex.
Triepeolus helianthi grandior Cockerell, 1919. N. Y. Ent. Soc., Jour. 27: 300. 2, ¢.
helianthi helianthi (Robertson). Ind., Ill, and Wis., west to Colo., Wyo. and Ariz. Host:
Melissodes trinodus Robt., M. composita Tucker?
Epeolus helianthi Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 344. 9.

Taxonomy: Cockerell, 1921. Amer. Mus. Novitates 23: 3, 15 (key).
Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 164-166, figs. —Hurd and Linsley,
1959. Ent. News 70: 141-146 (?host).
helianthi pacificus Cockerell. Colo.
Triepeolus helianthi pacificus Cockerell, 1919. N. Y. Ent. Soe., Jour. 27: 300. 6.
Taxonomy: Cockerell, 1921. Amer. Mus. Novitates 23: 4, 15 (key).
heterurus (Cockerell and Sandhouse). Calif., Oreg.
Epeolus heterurus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 316. 2, 3.
hopkinsi Cockerell. Ariz.
Triepeolus hopkinsi Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 184. 3.
insolitus Cockerell. Colo.
Triepeolus (Synepeolus) insolitus Cockerell, 1921. Amer. Mus. Novitates 23: 6. d.
inyoensis Cockerell and Sandhouse. Calif.
Triepeolus inyoensis Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. 4 (13): 309. ¢.
isocomae Cockerell. N. Mex.
Triepeolus isocomae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 38. ¢.
junctus Mitchell. N. Y., N.C.
Triepeolus junctus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 471. 2, 6.
laticaudus Cockerell. Colo.
Triepeolus laticaudus Cockerell, 1921. Amer. Mus. Novitates 23: 12. 2.
lestes Cockerell. Colo., Calif.
Triepeolus lestes Cockerell, 1921. Amer. Mus. Novitates 23: 11. °.

Taxonomy: Cockerell, 1929. Pan-Pacific Ent. 5: 102. 2, 6.
lineatulus Cockerell and Sandhouse. Calif., Ariz. Host: Melissodes tepida timberlakei Ckll.
Triepeolus lineatulus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 306. 2,
Or
loganensis Cockerell. Colo.
Triepeolus loganensis Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 624. d.
lunatus concolor (Robertson). Minn. to Pa., south to Fla. and Tex. Host: Melissodes
bimaculata bimaculata (LeP.).
Epeolus lunatus var. concolor Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 51.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 285, 286. 2, d (key). —Cockerell, 1904. Ann.
and Mag. Nat. Hist. (7) 13: 35. d (key). —Cockerell, 1921. Amer. Mus. Novitates 23: 14
(key).

2094 Hymenoptera in America North of Mexico

lunatus lunatus (Say). Minn. to New England, south to Fla., west to Tex., N. Mex. and Colo.
Epeolus lunatus Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 354. 9, d.

Taxonomy: Brues, 1903. Ent. News 14: 79, 80. 2, ¢d (key). —Robertson, 1903. Canad. Ent. 35:
284, 286. 2, d (key). —Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 62. 3 (key).
—Cockerell, 1921. Amer. Mus. Novitates 23: 2, 14. (key).

lusor Cockerell. Colo.
Triepeolus lusor Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 625. 3.

maculiventris Cockerell. Colo.
Triepeolus maculiventris Cockerell, 1921. Amer. Mus. Novitates 23: 11. 9.

martini (Cockerell). N. Mex., Colo.
Epeolus remigatus var. martini Cockerell, 1900. Canad. Ent. 32: 362. 6, °.
Taxonomy: Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 24 (tax. status).

mensae Cockerell. Oreg.
Triepeolus mensae Cockerell, 1924. Calif. Acad. Sei., Proce. (4) 13: 313. 3.

mesillae Cockerell. Kans., N. Mex. Host: Nomia triangulifera Vachal.
Triepeolus mesillae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 13: 36. 9.
Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 16-17, figs. 19-23 (larva).

michiganensis Mitchell. Conn., N. Y., Mich.
Triepeolus michiganensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 473. 2, 6.

micropygius atripes Mitchell. Pa., N. C., Ga.
Triepeolus micropygius atripes Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 474,
sie, WIPE Oe he

micropygius micropygius Robertson. Il.
Triepeolus micropygius Robertson, 1903. Canad. Ent. 35: 286. °.

mitchelli Hurd, n. name. N. C. (Marion).
Triepeolus sublunatus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 483. 3. Preoce.

mojavensis Linsley. Calif. Host: Anthophora linsleyi Timberlake.
Triepeolus mojavensis Linsley, 1939. Pan-Pacific Ent. 15: 2. 2.
Biology: Linsley and MacSwain, 1942. Amer. Midland Nat. 27: 409.

monardae Mitchell. N. C.
Triepeolus monardae Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 475, fig. 112. 3.

nevadensis (Cresson). N. C. and Ga., west to Nebr., Tex., N. Mex. and Nev.
Epeolus nevadensis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 86. 2.
Taxonomy: Robertson, 1903. Canad. Ent. 35: 285, 286. 2, d (key).
nigriceps (Smith). Calif., Tex.
Epeolus nigriceps Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 103. 2.
nigrihirtus Mitchell. N. C. (Chatham Co.).
Triepeolus nigrihirtus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 476. 6.

norae Cockerell. N. Mex.
Triepeolus norae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 59. °.
Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 161.
obliteratus Graenicher. Wis.
Triepeolus obliteratus Graenicher, 1911. Pub. Mus. City Milwaukee, Bul. 1: 242. 2, 6.
occidentalis (Cresson). Pacific Coast to Kans., N. Mex., Tex. Host: Melissodes mizeae
Cockerell?
Epeolus occidentalis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 87. 6, 2.
Taxonomy: Brues, 1903. Ent. News 14: 80 (key). —Cockerell, 1904. Ann. and Mag. Nat. Hist.
(7) 18: 34, 35. 2, 3 (key).

Biology: Hicks, 1926. Colo. Univ., Studies 15: 225.

paenepectoralis Viereck. B. C. to Wash., Colo.
Triepeolus paenepectoralis Viereck, 1905. Canad. Ent. 37: 278. 9.

Superfamily APOIDEA 2095

Taxonomy: Cockerell, 1921. Amer. Mus. Novitates 23: 15 (key).

pallidiventris Cockerell and Sandhouse. Utah.
Triepeolus pallidiventris Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proe. (4) 13: 308.
é.

pectoralis (Robertson). Maine to Ga., west to Minn., Colo. and Utah. Host: M elissodes rustica
(Say)?
Epeolus pectoralis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 345. °.
Epeolus virginiensis Cockerell, 1907. Entomologist 40: 137. d.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 285. ° (key). —Lovell and Cockerell, 1905.
Psyche 12: 42. ¢.
penicilliferus (Brues). Tex.
Epeolus penicilliferus Brues, 1903. Ent. News 14: 81. 2, 6.
Taxonomy: Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 18: 34. 2.
perelegans Cockerell. Ariz.
Triepeolus perelegans Cockerell, 1921. Amer. Mus. Novitates 23: 8. d.
permixtus (Cockerell). Calif.; Mexico (Baja California).
Epeolus permixtus Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 94. d, °.
pimarum Cockerell. Ariz.
Triepeolus pimarum Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 18: 36. °.
pomonalis Cockerell. Calif.
Triepeolus pomonalis Cockerell, 1916. Canad. Ent. 48: 392. d.
quadrifasciatus atlanticus Mitchell. Md. to Fla., Mo.
Triepeolus 4-fasciatus atlanticus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 479,
figs 2S =
quadrifasciatus quadrifasciatus (Say). Ark., Tex.
Epeolus 4-fasciatus Say, 1823. West. Quart. Rptr. 2: 81.
Taxonomy: Brues, 1903. Ent. News 14: 80 (key).
rectangularis Cockerell. Colo., Utah, Oreg., Calif.
Triepeolus rectangularis Cockerell, 1921. Amer. Mus. Novitates 23: 9. d.
remigatus (Fabricius). N. J. to Fla., west to Minn., Colo. and Calif., south to Tex., N. Mex. and
Ariz.; Mexico (northern). Host: Peponapis pruinosa (Say)?, Xenoglossa strenua (Cress.).
Melecta remigata Fabricius, 1804. Systema Piezatorum, p. 387.
Taxonomy: Cresson, 1864. Ent. Soe. Phila., Proc. 2: 293. 2, d. —Brues, 1903. Ent. News 14:
79 (key). —Robertson, 1903. Canad. Ent. 35: 285, 286. °, d (key). —Rozen, 1966. Amer.
Mus. Novitates 2244: 17-19, fig. 24 (larva).

Biology: Bohart, 1966. Pan-Pacifie Ent. 42: 255-262 (life history, immature stages).

rhododontus Cockerell. Colo.

Triepeolus rhododontus Cockerell, 1921. Amer. Mus. Novitates 23: 5. d.
robustus (Cresson). N. Mex.

Epeolus robustus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 85. 9.
rohweri Cockerell. Colo.

Triepeolus rohweri Cockerell, 1911. Ann. and Mag. Nat. Hist. (8) 8: 668. d.

rufithorax Graenicher. Fla. Host: Svastra obliqua obliqua (Say).
Triepeolus rufithorax Graenicher, 1928. Ent. News 39: 279. ¢, d.

Biology: Rozen, 1964. Amer. Mus. Novitates 2170: 9.

rugosus Mitchell. Fla.
Triepeolus rugosus Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 482, fig. 112. 2.

sandhousei Cockerell. Colo.
Triepeolus sandhousei Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 624. d.

sarothrinus (Cockerell). Calif.
Epeolus sarothrinus Cockerell, 1929. Pan-Pacifie Ent. 5: 103. 2, d.
Epeolus sarothrinus var. confluens Cockerell, 1929. Pan-Pacifie Ent. 5: 104. 6.

2096 Hymenoptera in America North of Mexico

saturninus Cockerell and Sandhouse. Calif.
Triepeolus saturninus Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proce. (4) 13: 312.

3

schwarzi schwarzi Cockerell. Colo.
Triepeolus schwarzi Cockerell, 1921. Amer. Mus. Novitates 23: 4. d.

schwarzi subcalens Cockerell and Sandhouse. Calif.
Triepeolus schwarzi subcalens Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proce. (4)
109 oe

segregatus (Cockerell). N. Mex.
Epeolus occidentalis var. segregatus Cockerell, 1900. Canad. Ent. 32: 361. 3 (2 misdet. ?).
Taxonomy: Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 62. 2, 3d (key).

sequior Cockerell. Colo.
Triepeolus sequior Cockerell, 1921. Amer. Mus. Novitates 23: 8. 6.

simplex Robertson. Wis., Ill. and Mich, south to N. C. and Ga.
Triepeolus simplex Robertson, 1903. Canad. Ent. 35: 285, 286. 2, d.

stricklandi Cockerell. Alta.
Triepeolus stricklandi Cockerell, 1937. Canad. Ent. 69: 86. °.

subalpinus Cockerell. Colo.
Triepeolus subalpinus Cockerell, 1910. Psyche 17: 245. 9.

sublunatus Cockerell. N. Mex.
Triepeolus sublunatus Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 62. 3.

subnitens Cockerell and Timberlake. Calif.
Triepeolus subnitens Cockerell and Timberlake, 1929. Pan-Pacific Ent. 5: 167. d, °.

superbus (Provancher). Calif.
Epeolus superbus Provancher, 1895. Nat. Canad. 22: 190.

tanneri Cockerell. Utah.
Triepeolus tanneri Cockerell, 1928. Psyche 35: 232. 6.

texanus nigripes Cockerell. N. Mex.
Triepeolus texanus var. nigripes Cockerell, 1898. N. Mex. Univ., Bul. 1: 61. 6.

texanus texanus (Cresson). Tex.
Epeolus texanus Cresson, 1878. Amer. Ent. Soc., Trans. 7: 87. d.
Taxonomy: Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 18: 34. 2 (key).

timberlakei Cockerell. Calif., Utah. Host: Melissodes tepida timberlakei Ckll.
Triepeolus timberlakei Cockerell, 1929. Pan-Pacifie Ent. 5: 101. 2, 6.
Triepeolus timberlakei var. heterodoxus Cockerell, 1929. Pan-Pacific Ent. 5: 101. 9.

townsendi Cockerell. N. Mex.
Triepeolus townsendi Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 20: 68. 6, 9.

trichopygus Cockerell and Timberlake. Calif.
Triepeolus trichopygus Cockerell and Timberlake, 1929. Pan-Pacific Ent. 5: 169. 2, 6.

trilobatus Cockerell. Colo.
Triepeolus trilobatus Cockerell, 1921. Amer. Mus. Novitates 23: 7. 6.

utahensis (Cockerell). Utah.
Epeolus utahensis Cockerell, 1921. Amer. Mus. Novitates 23: 15. 6.

vandykei Cockerell and Sandhouse. Calif.
Triepeolus vandykei Cockerell and Sandhouse, 1924. Calif. Acad. Sci., Proc. (4) 13: 307. 2.

verbesinae (Cockerell). N. Mex. to Calif.; Mexico (Baja California).
Epeolus verbesinae Cockerell, 1897. Amer. Ent. Soe., Trans. 24: 156. 6, 9.

Taxonomy: Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 11: 60. 3d (key).

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 161.

wyomingensis Cockerell. Wyo. to Utah.
Triepeolus wyomingensis Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 201. 6.

Superfamily APOIDEA 2097

TRIBE NOMADINI

Although this tribe is represented in the Old World especially by numerous species of
Nomada, it is chiefly an American group inhabiting both North and South America. In America
north of Mexico there are nearly 300 species of Nomada as well as several small genera which
are mostly centered in the more arid parts of the southwestern United States and adjacent Mex-
ico. While there is some evidence to suggest that the members of this tribe and those of the
tribe Epeolini should be grouped together, these tribes are maintained as distinct in this catalog.

On the basis of our current knowledge concerning host relationships most of the species are
cleptoparasites in the nests of the family Andrenidae, especially the genus Andrena, but some of
the species (and even groups of species) also parasitize the nests of Halictidae, Melittidae, and
Anthophoridae.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 22-28, figs. 39-55 (larvae).
Genus NOMADA Scopoli

The bees of this genus are wasp-like in appearance and are present on all the continents and
many of the islands. The genus is represented in the Nearctic Region by numerous species of
several subgenera. These bees are cleptoparasitic in the nests of other bees, principally the
genus Andrena, but are also known to parasitize the nests of certain Halictidae, Melittidae and
Anthophoridae. As with many cleptoparasitic bees, they are frequently encountered either fly-
ing about the nesting sites of their hosts or sipping nectar with them at the same flowers.

Taxonomy: Cresson, 1887. Amer. Ent. Soc., Trans., Sup. (2): 296-297. —Robertson, 1903.
Canad. Ent. 35: 172-179. —Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 559-579,
580-610. —Cockerell, 1904 (1903). Colo. Expt. Sta., Rpt. of Ent., Bul. 94: 65-85. —Cockerell,
1905. Amer. Ent. Soe., Trans. 31: 309-312 (notes on British Museum types). —Cockerell,
1911. U.S. Natl. Mus., Proce. 41: 225-243. —Swenk, 1912. Nebr. Univ., Studies 2 (1): 1-113.
—Rodeck, 1931. Amer. Mus. Novitates 496: 1-11. —Mitchell, 1962. N. C. Agr. Expt. Sta.
Tech. Bul. 152: 338-437, figs. 98-107, tables 11-13 (eastern U. S. spp.).

Morphology: Cockerell and Atkins, 1902. Ann. and Mag. Nat. Hist. (7) 10: 40-44. —Beck,
1933. Utah Acad. Sci., Arts and Letters, Proc. 10: 101, figs. 60-64. —Linsley and Michener,
1939. Amer. Ent. Soc., Trans. 65: 265-305, 4 pls. —Snodgrass, 1941. Smithsn. Mise. Coll. 99:
55-56, pl. 27 F-J.

Genus NOMADA Subgenus NOMADA Scopoli

Nomada Scopoli, 1770. Historico Naturalis, Ann. 4: 44.
Type-species: Nomada ruficornis (Linnaeus). Desig. by Curtis, 18382.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 174-175, 178-179. —Cockerell, 1903. Acad. Nat.
Sci. Phila., Proc. 55: 569-579, 589 (in part), ete. —Viereck et al., 1905. Canad. Ent. 37:
285-287. —Swenk, 1913. Nebr. Univ., Studies 12: 10-11, 15-57. —Swenk, 1915. Nebr. Univ.,
Studies 15: 155-163. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 402-436, figs.
106-107 (eastern U.S. spp.).
accepta Cresson. Colo., Kans.

Nomada accepta Cresson, 1878. Amer. Ent. Soc., Trans. 7: 77. 2, d.

Nomada pacata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 81. 9.
aldrichi Cockerell. Idaho, Wash., B. C.

Nomada vicinalis aldrichi Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 368. 6.
amoena Cresson. IIl.

Nomada amoena Cresson, 1863. Ent. Soe. Phila., Proc. 2: 300. °.
angelarum Cockerell. Calif.

Nomada (Nomada) angelarum Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 574. °.
angulata Swenk. Nebr.

Nomada (Nomada) angulata Swenk, 1913. Nebr. Univ., Studies 12: 40. °.
aprilina Swenk. Nebr.

Nomada (Nomada) aprilina Swenk, 1913. Nebr. Univ., Studies 12: 28. d.

2098 Hymenoptera in America North of Mexico

armatella Cockerell. Mich. to New England.

Nomada armatella Cockerell. 1903. Acad. Nat. Sci. Phila., Proc. 55: 606. 3.
astori Cockerell. Oreg.

Nomada astori Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 206. 2. Var. a= mediana

Swenk.

atrofrontata Cockerell. South. Calif.

Nomada (Nomada) atrofrontata Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 573. 2.
augustiana Mitchell. Ga. (Augusta).

Nomada augustiana Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 402. 9.
azaleae Mitchell. N. C. (Highlands).

Nomada azaleae Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 403, fig. 107. 3.
banksi Cockerell. Ont. and N. Y., south to IIl. and N. C.

Nomada banksi Cockerell, 1907. Entomologist 40: 98. °.
beulahensis Cockerell. N. Mex., N. Dak.

Nomada beulahensis Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 209. ?.
bicrista Swenk. Nebr.

Nomada (Nomada) bicrista Swenk, 1913. Nebr. Univ., Studies 12: 27. 6.
bifurcata Cockerell. Calif.

Nomada (Nomada) bifurcata Cockerell, 1903. Acad. Nat. Sci. Phila., Proce. 55: 577. 6.
bisetosa Swenk. Nebr.

Nomada (Nomada) bisetosa Swenk, 1913. Nebr. Univ., Studies 12: 30. 9.
bisignata Say. “United States.”

Nomada bisignata Say, 1824. In Keating, Narr. Long’s 2nd Exped., v. 2, p. 354. @.

Nomada bicincta(!) Howard, 1902. Insect Book, pl. 3, fig. 32.
californiae Cockerell. Calif.

Nomada (Nomada) Californiae Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 571. 9.
capillata Mitchell. Mass. (Milton).

Nomada capillata Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 406. 3.
carinicauda Cockerell. Colo.

Nomada carinicauda Cockerell, 1921. Amer. Mus. Novitates 24: 7. 9.
ceanothi Cockerell. N. Y. to N. C.

Nomada (Nomada) ceanothi Cockerell, 1907. Entomologist 40: 97. 2.
clarkii Cockerell. Oreg.

Nomada Clarkui Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 208. °.
coloradella Cockerell. Colo.

Nomada coloradella Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 83. 6, °.
composita Mitchell. Conn., Mich.

Nomada composita Mitchell, 1962. N. C. Agr. Expt. Sta., Tech. Bul. 152: 408. @.
cordleyi Cockerell. Oreg.

Nomada Cordleyi Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 445. ¢.
corvallisensis Cockerell. Oreg.

Nomada corvallisensis Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 207. 2.
cressonii cressonii Robertson. Colo., N. Dak. to N. S. south to Tenn. and N. C.

Nomada cressonii Robertson, 1893. Amer. Ent. Soc., Trans. 20: 275. 2, ¢.

Nomada mera Cockerell, 1908. Ent. Soc. Wash., Proc. 10: 83. ¢.
cressonii trevoriana Cockerell. Wash.

Nomada (Nomada) Cressonii Trevoriana Cockerell, 1905. Canad. Ent. 37: 285.
crudelis Cresson. Ga. Host: Andrena obscuripennis Sm?

Nomada crudelis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 82. 2.
cymbalariae Cockerell. Colo.

Nomada cymbalariae Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 439. 2.
davidsoni Cockerell. Calif.

Nomada (Nomada) davidsoni Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 575. 9.

Superfamily APOIDEA 2099

decepta Mitchell. N. Y., Mich.
Nomada decepta Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 411. @.
denticulata Robertson. Minn. to N. Y., south to Ga.
Nomada denticulata Robertson, 1902. Canad. Ent. 34: 49.
Nomada simplex Robertson, 1902. Ent. News 13: 80. 6.
depressa Cresson. Nebr., Mich. to Maine, south to N. C.
Nomada depressa Cresson, 1863. Ent. Soc. Phila., Proc. 2: 302. 9.
Nomada depressicauda Cockerell, 1908. Ent. News 19: 323. 2.
detrita Mitchell. N. H. (Randolph).
Nomada detrita Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 414, figs. 106, 107. d.
dreisbachi Mitchell. Maine, Mass., Mich.
Nomada dreisbachi Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 414. 6.
elegantula Cockerell. Calif., Idaho.
Nomada (Nomada) elegantula Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 578. °.
erythraea Dalla Torre. Calif.
Nomada rubra Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym. p. 427. 2. Preoce.
Nomada erythraea Dalla Torre, 1896. Cat. Hym., v. 10, p. 343. N. name.
flammigera Cockerell. Wash.
Nomada (Nomada) flammigera Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 18: 71. 2.
florilega Lovell and Cockerell. Mich. to Maine, south to N. C.
Nomada (Nomada) florilega Lovell and Cockerell, 1905. Psyche 12: 41. 9.
fontis Cockerell. Colo.
Nomada (Nomada) fontis Cockerell, 1910. Canad. Ent. 42: 367. °.
gibbosa Viereck. Oreg.
Nomada gibbosa Viereck, 1905. Canad. Ent. 37: 285. d.
gracilis Cresson. N. Y. and New England.
Nomada gracilis Cresson, 1863. Ent. Soc. Phila., Proc. 2: 295. ¢.
hoodiana Cockerell. Oreg.
Nomada hoodiana Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 608. 2.
idahoensis Swenk. Idaho.
Nomada (Nomada) idahoensis Swenk, 1913. Nebr. Univ., Studies 12: 53. d.
illinoensis Robertson. Minn. to Mass., south to Ga. Predator: Pselliopus barberi Davis.
Nomada IIlinoiensis Robertson, 1900. Canad. Ent. 32: 294. 2, 6.
Nomada illinoensis Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 657. Emend.
Nomada illinoensis Cockerell, 1921. Amer. Mus. Novitates 24: 7. 2. Emend.; diaeresis
placed over e in illinoensis.

Biology: Bouseman, 1976. Kans. Ent. Soc., Jour. 49: 384 (predator).

indusata Mitchell. N. C., Ga.

Nomada indusata Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 418, fig. 107. d.
inepta Mitchell. Maine to N. C., Minn.

Nomada inepta Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 419. °.
interruptella Fowler. Calif.

Nomada interrupta Fowler, 1899. Ent. News 10: 159. 6. Preoce.

Nomada interruptella Fowler, 1902. Calif. Agr. Expt. Sta., Rpt., 1898-1901, p. 329. N.

name.

itamera Cockerell. Wash.

Nomada itamera Cockerell, 1910. Psyche 17: 95. ¢.
kincaidiana Cockerell. Wash.

Nomada kincaidiana Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 614. 9.
kingstonensis Mitchell. R. I. (Kingston).

Nomada kingstonensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 420. °.
latifrons Cockerell. Calif., Nev.

Nomada (Nomada) latifrons Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 572. 2.

2100 Hymenoptera in America North of Mexico

lehighensis Cockerell. Mich. to N. S., south to Ga.

Nomada lehighensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 605. °, d.
lewisi Cockerell. Oreg.

Nomada Lewisi Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 205. 9.
libata Cresson. Colo.

Nomada libata Cresson, 1878. Amer. Ent. Soce., Trans. 7: 80. 2, ¢.

Nomada limbata(!) Dalla Torre, 1896. Cat. Hym., v. 10, p. 354.
luteopicta Cockereli. Colo.

Nomada luteopicta Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 83. d, °.
malonella Cockerell. Wash.

Nomada (Nomada) malonella Cockerell, 1910. Psyche 17: 98. 2, 5 ?
malonina Cockerell. Wash.

Nomada (Nomada) malonina Cockerell, 1910. Psyche 17: 94. 6.
marginella Cockerell. Calif.

Nomada (Nomada) marginella Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 576. °.
mckenziei Timberlake and Cockerell. Calif.

Nomada mckenziei Timberlake and Cockerell, 1937. Amer. Mus. Novitates 948: 4. 2, 6.
media Mitchell. N. C., Tenn.

Nomada media Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 421, fig. 107. 2, 6.
mediana Swenk. Oreg., Calif.

Nomada (Nomada) mediana Swenk, 1913. Nebr. Univ., Studies 12: 49. 2.
melanosoma Cockerell. Calif.

Nomada melanosoma Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 55. 6.
mendica Mitchell. Vt., N. H.

Nomada mendica Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 422, fig. 107. d.
minima Mitchell. Md. to N. C.

Nomada minima Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 423. ¢.
minuta Swenk. Nebr.

Nomada (Nomada) minuta Swenk, 1913. Nebr. Univ., Studies 12: 44. @.
munda Cresson. Colo.

Nomada munda Cresson, 1878. Amer. Ent. Soc., Trans. 7: 80. °.
nigrociliata Swenk. Nebr.

Nomada (Nomada) nigrociliata Swenk, 1913. Nebr. Univ., Studies 12: 32. °.
nigrocincta Smith. “Arctic America,” Maine, Oreg.

Nomada nigrocincta Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 99. 2.
nigrofasciata Swenk. Nebr.

Nomada (Nomada) nigrofasciata Swenk, 1913. Nebr. Univ., Studies 12: 24. °.
obliquella Fowler. Calif. Host: Andrena swavis Timberlake.

Nomada obliqua Fowler, 1899. Ent. News 10: 160. ¢. Preoce.

Nomada obliquella Fowler, 1902. Calif. Agr. Expt. Sta., Rpt., 1898-1901, p. 329. N. name.

Biology: Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 26 (habits, host).

obscurella Fowler. Calif., Oreg. Host: Andrena caerulea Smith.
Nomada obscura Fowler, 1899. Ent. News 10: 160. ¢. Preoce.
Nomada obscurella Fowler, 1902. Calif. Agr. Expt. Sta., Rpt., 1898-1901, p. 329. N. name.
Nomada Fowleri Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 204. °.

Taxonomy: Rozen, 1966. Amer. Mus. Novitates 2244: 22-24, figs. 39-40 (larva).

Biology: Linsley and MacSwain, 1955. Wasmann Jour. Biol. 13: 275, pl. 1, fig. 4 (life history).

obtusata Swenk. N. Dak.
Nomada (Nomada) obstusata(!) Swenk, 1915. Nebr. Univ., Studies 15: 159. 2. Lapsus
calami, fide Swenk, personal communication with Dr. Hugo Rodeck.
ochrohirta Swenk. Nebr.
Nomada (Nomada) ochrohirta Swenk, 1913. Nebr. Univ., Studies 12: 34. °.

Superfamily APOIDEA 2101

odontocera Cockerell. Calif.

Nomada odontocera Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 57. 6.
opposita Cresson. Calif.

Nomada opposita Cresson, 1878. Amer. Ent. Soc., Trans. 8: 73. 2.
orba Mitchell. N. Y. (Ithaca).

Nomada orba Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 424, fig. 107. d.
orcusella Cockerell. Wash.

Nomada (Nomada) orcusella Cockerell, 1910. Psyche 17: 95. °.
oregonica Cockerell. Oreg., Calif.

Nomada oregonica Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 205. 9, d.
osborni Cockerell. Ariz.

Nomada (Nomada) osborni Cockerell, 1911. U. S. Natl. Mus., Proc. 41: 243. 6.
packardiella Cockerell. Colo., Wash.?

Nomada Packardiella Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 225. 9.
pallidipicta Swenk. Nebr.

Nomada (Nomada) pallidipicta Swenk, 1913. Nebr. Univ., Studies 12: 48. d.
parallela Swenk. Nebr.

Nomada (Nomada) parallela Swenk, 1913. Nebr. Univ., Studies 12: 36. d.
parata Cresson. Colo.

Nomada parata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 81. 2.
parva Robertson. Mich. to New England, south to Miss. and Tex.

Nomada parva Robertson, 1900. Canad. Ent. 32: 294. 2, 6.

Nomada infantula Cockerell, 1907. Entomologist 40: 98. 2, 6d.
propinqua Swenk. Nebr.

Nomada (Nomada) propinqua Swenk, 1913. Nebr. Univ., Studies 12: 46. 2, d.
pulsatillae Cockerell. Colo., Wash.?

Nomada pulsatillae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 18: 69. °.
pygmaea Cresson. Minn. to Maine, south to Va., ?Colo.

Nomada pygmaea Cresson, 1863. Ent. Soc. Phila., Proce. 2: 299. ¢.
rhodosoma rhodosoma Cockerell. Calif., Nev.

Nomada (Nomada) rhodosoma Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 571. 2, 5 ?
rhodosoma rhodosomella Cockerell. Colo.

Nomada rhodosoma var. rhodosomella Cockerell, 1903. Acad. Nat. Sci. Phila., Proe. 55:

Seneca

robertsonella Cockerell. Nev.

Nomada robertsonella Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 618. 9.
rubrica Provancher. Calif. L

Nomada rubrica Provancher, 1896. Nat. Canad. 238: 8. 2.

Taxonomy: Fowler, 1899. Ent. News 10: 162.

salicicola Swenk. Nebr.

Nomada (Nomada) salicicola Swenk, 1913. Nebr. Univ., Studies 12: 35. d.
salicis Robertson. IIl.

Nomada salicis Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 52. d.

sayi Robertson. Minn. to Que. and Maine, south to Ga.
Nomada sayi Robertson, 1893. Amer. Ent. Soc., Trans. 20: 276. °, d.

Taxonomy: Robertson, 1900. Canad. Ent. 32: 293-294. 2, d (redescribed, restricted).

sedi Cockerell. Colo.
Nomada sedae Cockerell, 1919. Ent. News 30: 292. 2.
Nomada sedi Cockerell, 1920. Ann. and Mag. Nat. Hist. (9) 6: 201. Emend.

semirugosa Cockerell. Colo.
Nomada semirugosa Cockerell, 1929. Ann. and Mag. Nat. Hist. (10) 4: 297.

skinneri Cockerell. Conn. to N. C.
Nomada skinneri Cockerell, 1908. Ent. News 19: 323. °.

2102 Hymenoptera in America North of Mexico

sobrina Mitchell. Mich. (Macomb Co.).
Nomada sobrina Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 431, fig. 107. 6.

sphaerogaster Cockerell. N. J., Wis.
Nomada sphaerogaster Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 611. °.

subaccepta Cockerell. Colo.
Nomada subaccepta Cockerell, 1907. Entomologist 40: 267. ¢.

subangusta Cockerell. Calif.
Nomada (Nomada) subangusta Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 570. 3.

subpacata Swenk. N. Dak.
Nomada (Nomada) subpacata Swenk, 1913. Nebr. Univ., Studies 12: 55. °.

subvicinalis Cockerell. Calif.
Nomada (Nomada) subvicinalis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 576. 3.

taraxacella Cockerell. N. Mex., Colo., Nebr., Wash.?
Nomada ultima taraxacella Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 589. °.
Taxonomy: Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 74. —Cockerell, 1911. U. S. Natl.
Mus., Proc. 41: 238. 2, 6. —Swenk, 1913. Nebr. Univ., Studies 12: 22.

‘intinnabelum Cockerell. Calif.
Nomada (Nomada) tintinnabulwm Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 572. 9,

townesi Mitchell. Md. (Takoma Park).
Nomada townesi Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 432. ¢.

tyrrellensis Mitchell. N. C., Fla.
Nomada tyrrellensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 433. 9.

ulsterensis Mitchell. N. Y., D. C.
Nomada ulsterensis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 433, fig. 107. ¢.

ultima Cockerell. Wash., Oreg., Calif.
Nomada ultima Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 206. 9.
Nomada modocorum Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 444. 6.

ultimella septentrionalis Swenk. Wash.
Nomada (Nomada) ultimella septentrionalis Swenk, 1913. Nebr. Univ., Studies 12: 50. 9.

ultimella ultimella Cockerell. Calif.
Nomada (Nomada) ultimella Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 572. 9.

undulaticornis Cockerell. Colo.
Nomada undulaticornis Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 18: 70. 6.

valida Smith. N. S. and N. Y. to B. C.
Nomada valida Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 246. 9.

velutina Swenk. Nebr.
Nomada (Nomada) velutina Swenk, 1913. Nebr. Univ., Studies 12: 39. °.

vicina stevensi Swenk. N. Dak.
Nomada (Nomada) vicina stevensi Swenk, 1913. Nebr. Univ., Studies 12: 56. 2, 6.

vicina vicina Cresson. Mich. to Que., New England, N. J. and Nebr. Host: Andrena vicina Sm.
Nomada vicina Cresson, 1863. Ent. Soe. Phila., Proc. 2: 292. 2 (¢ misdet.).
Biology: Packard, 1878. Guide ... study of insects, 6th ed., p. 142 (life history).

vicinalis infrarubens Cockerell. Oreg.
Nomada vicinalis var. infrarubens Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 84. ¢.

vicinalis vicinalis Cresson. Nebr., Colo.
Nomada vicinalis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 78. ¢.
Taxonomy: Swenk, 1913. Nebr. Univ., Studies 12: 41. d, °.
wootonella Cockerell. N. Mex.
Nomada wootonella Cockerell, 1909. Entomologist 42: 93. d.
wyomingensis Swenk. Wyo.
Nomada (Nomada) wyomingensis Swenk, 1913. Nebr. Univ., Studies 12: 52. °.

ziziae Swenk. N. Dak.
Nomada (Nomada) ziziae Swenk, 1915. Nebr. Univ., Studies 15: 4. 2, ¢.

Superfamily APOIDEA 2103

Genus NOMADA Subgenus GNATHIAS Robertson

Gnathias Robertson, 1903. Canad. Ent. 35: 173.
Type-species: Nomada bella Cresson. Orig. desig.

Taxonomy: Cockerell, 1903. Acad. Nat. Sci. Phila., Proce. 55: 594-603. —Viereck et al., 1905.
Canad. Ent. 37: 282-283. —Swenk, 1913. Nebr. Univ., Studies 12: 11-15, 90-99. —Swenk,
1915. Nebr. Univ., Studies 15: 172-179. — Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul.
152: 346-354, fig. 98 (eastern U.S. spp.).

bella bella Cresson. Minn. to Que. and Maine, south to Fla.
Nomada bella Cresson, 1863. Ent. Soc. Phila., Proc. 2: 287. 3.
Nomada albofasciata Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 100. 3.

Morphology: Snodgrass, 1941. Smithsn. Mise. Coll. 99: 55-56, pl. 27, figs. F-J.

bella callura Cockerell. Colo.

Nomada (Gnathias) bella callura Cockerell, 1911. U. S. Natl. Mus., Proce. 39: 657. 3.
clarescens Cockerell. Colo.

Nomada (Gnathias) clarescens Cockerell, 1921. Amer. Mus. Novitates 24: 10. 9°.
cuneata (Robertson). Minn. to N. S., south to N. C.

Gnathias cuneatus Robertson, 1903. Canad. Ent. 35: 175. 2, 3 ?

Gnathias cuneatus form decemnotatus Robertson, 1903. Canad. Ent. 35: 175. @.

Gnathias cuneatus form octonotatus Robertson, 1903. Canad. Ent. 35: 175. @.

Gnathias cuneatus form sexnotatus Robertson, 1903. Canad. Ent. 35: 176. 9.

Gnathias cuneatus form quadrisignatus Robertson, 1903. Canad. Ent. 35: 176. 9.
custeriana Cockerell. Colo.

Nomada (Gnathias) custeriana Cockerell, 1911. U. S. Natl. Mus., Proce. 41: 241. 3.
debilis Timberlake. Calif.

Nomada (Gnathias) debilis Timberlake, 1954. Pan-Pacifie Ent. 30: 135. 2, 3.

Nomada (Gnathias) debilis var. a Timberlake, 1954. Pan-Pacific Ent. 30: 136. °.
fuscicincta Swenk. N. Dak.

Nomada (Gnathias) fuscicincta Swenk, 1915. Nebr. Univ., Studies 15: 24. 9, d.
grayi eastonensis Cockerell. Wash.

Nomada (Gnathias) grayi eastonensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proce. 55: 597.

OF

grayi grayi Cockerell. Oreg.

Nomada Gray Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 203. @.
heterosticta Cockerell. Idaho.

Nomada (Gnathias) heterosticta Cockerell, 1921. Amer. Mus. Novitates 24: 9. °.
hydrophylli Swenk. N. Dak., Ill., Md.

Nomada (Gnathias) hydrophylli Swenk, 1915. Nebr. Univ., Studies 15: 25. 3.

Gnathias xanthoparius Robertson, 1928. Flowers and Insects, Carlinville, Ill. pp. 9, 101,

111, 152. 6. Nomen nudum.

klamathensis Fox. Oreg.

Nomada (Gnathias) klamathensis Fox, 1926. Pan-Pacifie Ent. 5: 212. 6.
lepida Cresson. Minn. to Maine, south to Ga.

Nomada lepida Cresson, 1863. Ent. Soe. Phila., Proc. 2: 288. d.
leucozona Rodeck. Colo.

Nomada (Gnathias) leucozona Rodeck, 1931. Amer. Mus. Novitates 496: 4. 6.
louisianae Cockerell. Nebr. and Minn. to N. Y., S. C. and La.

Nomada (Gnathias) louisianae Cockerell, 1903. Acad. Nat. Sei. Phila., Proc. 55: 602. °.
maculata Cresson. Minn. to Maine, south to Va.

Nomada maculata Cresson, 1863. Ent. Soc. Phila., Proc. 2: 303. 2, d ?

Nomada volatilis Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 100. d.
opacella Timberlake. Calif. Host: Andrena caerulea Sm., A. swavis Timb.

Nomada (Gnathias) opacella Timberlake, 1954. Pan-Pacifie Ent. 30: 133. 2, d.

2104 Hymenoptera in America North of Mexico

Biology: Linsley and MacSwain, 1955. Wasmann Jour. Biol. 13: 256-270, pl. 1, fig. 1; pls. 2 and
3 (life history, hosts). —Linsley and MacSwain, 1959. Calif. Univ. Pubs. Ent. 16: 25-26
(habits, hosts).
orophila Cockerell. Colo., Wyo.
Nomada (Gnathias) orophila Cockerell, 1921. Amer. Mus. Novitates 24: 8. d, 9.
ovata (Robertson). Wyo., Minn. to Maine, south to S. C.
Gnathias ovatus Robertson, 1903. Canad. Ent. 35: 175. @, d.
Gnathias ovatus form plenus Robertson, 1903. Canad. Ent. 35: 175. @.
Gnathias ovatus form octomaculatus Robertson, 1903. Canad. Ent. 35: 175. °.
Gnathias ovatus form sexmaculatus Robertson, 1903. Canad. Ent. 35: 175. 9.
Gnathias ovatus form quadrimaculatus Robertson, 1903. Canad. Ent. 35: 175. °.
Gnathias ovatus form binotatus Robertson, 1903. Canad. Ent. 35: 175. 2.
Gnathias ovatus form unicolor Robertson, 1903. Canad. Ent. 35: 175. 9°.
Nomada (Gnathias) carolinae Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 602. 9.
perbella (Viereck). Oreg., Wash., B. C.
Gnathias perbella Viereck, 1905. Canad. Ent. 37: 282. °, d.
perplexa Cresson. Minn. to Mass., south to Tenn. and N. C., ?Wyo.
Nomada perplexa Cresson, 1863. Ent. Soc. Phila., Proc. 2: 306. 2°, 5 ?

Taxonomy: Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 600. d, °.

Nomada (Gnathias) cornelliana Cockerell, 1908. Ent. Soc. Wash., Proc. 10: 84. d.
perplexans Cockerell. Wash.

Nomada (Gnathias) perplexans Cockerell, 1910. Psyche 17: 94. d.
physura Cockerell. Nev.

Nomada (Gnathias) physura Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 599. 6.
rhodalis Cockerell. Nev.

Nomada (Gnathias) rhodalis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 600. °.
rhodomelas Cockerell. Oreg., B. C.

Nomada (Gnathias) rhodomelas Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55:

598. 6, 2?

rubi Swenk. N. Dak., Minn.

Nomada (Gnathias) rubi Swenk, 1915. Nebr. Univ., Studies 15: 20. 2, 6.

rubrella Cockerell. Colo.
Nomada (Gnathias) rubrella Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 75. °.
schwarzi contractula Cockerell. N. Mex.
Nomada (Gnathias) schwarzi contractula Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55:
600. 6.

schwarzi schwarzi Cockerell. Colo.
Nomada (Gnathias) schwarzi Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 599. d (2
misdet.).
siouxensis Swenk. Nebr., Wyo.
Nomada (Gnathias) siowxensis Swenk, 1913. Nebr. Univ., Studies 12: 96. 2.
subnigrocincta Swenk. Mass., N. J., Pa., N. C.
Nomada (Nomada) subnigrocincta Swenk, 1915. Nebr. Univ., Studies 15: 155. 2, (d
misdet.).
subrubi Swenk. N. Dak.
Nomada (Gnathias) Subrubi Swenk, 1915. Nebr. Univ. Studies 15: 176. 2, 6.
utensis Swenk. Colo.
Nomada (Gnathias) utensis Swenk, 1913. Nebr. Univ., Studies 12: 97. 2, 6.
vulpis Cockerell. Wyo.
Nomada (Gnathias) vulpis Cockerell, 1921. Amer. Mus. Novitates 24: 10. °.

washingtoni Cockerell. Wash.
Nomada (Gnathias) washingtoni Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 598. °.

Superfamily APOIDEA 2105

Genus NOMADA Subgenus PHOR Robertson

Phor Robertson, 1903. Canad. Ent. 35: 173, 177.
Type-species: Nomada integerrima Dalla Torre. Orig. desig. and monotypic.
(=Nomada integra Robertson) (preocc.).

Taxonomy: Swenk, 1913. Nebr. Univ., Studies 12: 12, 14. —Mitchell, 1962. N. C. Agr. Expt.
Sta. Tech. Bul. 152: 377-379 (eastern U.S. spp.).
integerrima Dalla Torre. Minn. to Que. and New England, south to N. C. and La.
Nomada integra Robertson, 1893. Amer. Ent. Soc., Trans. 20: 276. 2, 6. Preoce.
Nomada integerrima Dalla Torre, 1896. Cat. Hym. v..10, p. 353. N. name.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 174, 175, 177. °, 6.

proxima Cresson. Maine, ?Conn.
Nomada proxima Cresson, 1863. Ent. Soc. Phila., Proc. 2: 294. 3.

Taxonomy: Viereck, 1916. Conn. State Geol. and Nat. Hist. Survey, Bul. 22 (3): 726.
siccorum Cockerell. Colo.
Nomada (Phor) siccorum Cockerell, 1919. Ent. News 30: 291. 6.
subgracilis Cockerell. Calif.
Nomada (Phor) subgracilis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 569. d.

vexator Cockerell. Colo.
Nomada vexator Cockerell, 1909. Entomologist 42: 92. °, d.

Genus NOMADA Subgenus HEMINOMADA Cockerell and Atkins

Nomada subg. Heminomada Cockerell and Atkins, 1902. Ann. and Mag. Nat. Hist. (7) 10:
42.
Type-species: Nomada obliterata Cresson. Orig. desig. and monotypic.
Xanthidium Robertson, 1903. Canad. Ent. 35: 174, 177. Preoce.
Type-species: Nomada luteola Olivier. Orig. desig.

Taxonomy: Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 564-569, 580-587 (part).
—Robertson, 1903. Canad. Ent. 35: 173, 174-175, 177. —Viereck et al., 1905. Canad. Ent.
37: 284-285. —Swenk, 1913. Nebr. Univ., Studies 12: 8-11, 13-15, 57-76. —Swenk, 1915.
Nebr. Univ., Studies 15: 163-169. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
380-400, fig. 105 (eastern U. S. spp.).
agynia Cockerell. Colo.
Nomada agynia Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 81. d.

alpha alpha Cockerell. Colo.

Nomada alpha Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 84. °.

Taxonomy: Swenk, 1913. Nebr. Univ., Studies 12: 71. 2, d.

alpha dialpha Cockerell. Colo. June.

Nomada alpha var. dialpha Cockerell, 1921. Amer. Mus. Novitates 24: 3. 2.
alpha paralpha Cockerell. Colo.

Nomada alpha var. paralpha Cockerell, 1921. Amer. Mus. Novitates 24: 3. 9.
annulata Smith. N. C. Host: Andrena macra Mitch.

Nomada annulata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 248. 2, (d misdet.).
arizonica Cockerell. Ariz.

Nomada (Xanthidium) arizonica Cockerell, 1911. U.S. Natl. Mus., Proce. 41: 242. 6.
ashmeadi Cockerell. Calif.

Nomada (Xanthidium) ashmeadi Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 568. °.
autumnalis Mitchell. Mich. (Oakland Co.).

Nomada (?Xanthidium) autumnalis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:

382. 3.

avalonica Cockerell. Calif.

Nomada (Holonomada) avalonica Cockerell, 1938. Ann. and Mag. Nat. Hist. (11) 2: 152. °.

bethunei Cockerell. Mich., Ohio.
Nomada bethunei Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 607. ¢.

2106 Hymenoptera in America North of Mexico

bishoppi Cockerell. Minn. to Maine, south to Miss. and Fla.
Nomada (Xanthidium) luteola bishoppi Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 655. 9.
calloxantha Cockerell. Wyo.
Nomada (Xanthidium) calloxantha Cockerell, 1921. Amer. Mus. Novitates 24: 4. °.
capitalis Mitchell. N. H., Mich., N. C., Tenn.
Nomada (Heminomada) capitalis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 385.
oF
citrina citrina Cresson. Pacific Coast.
Nomada citrina Cresson, 1878. Amer. Ent. Soc., Trans. 7: 79 °.
citrina flavomarginata Swenk. Nebr.
Nomada (Heminomada) citrina flavomarginata Swenk, 1913. Nebr. Univ., Studies 12:
WEG Os fore
citrina rufula Cockerell. Idaho.
Nomada citrina var. rufula Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 586. °.
civilis civilis Cresson. United States west of Nebr.
Nomada civilis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 78. d.
civilis spokanensis Cockerell. Wash.
Nomada civilis spokanensis Cockerell, 1910. Psyche 17: 92. 9.
collinsiana Cockerell. Colo.
Nomada (Xanthidium) collinsiana Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 79. @.

coloradensis Cockerell. Colo.
Nomada coloradensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 603. °.

Taxonomy: Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 84. 2, 6.
colorata Mitchell. N. Y., Mich.
Nomada (Heminomada) colorata Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 386.
oF
concinnula Cockerell. Colo.
Nomada (Xanthidium) concinnula Cockerell, 1921. Amer. Mus. Novitates 24: 6. 2.
coquilletti Cockerell. Calif., Wash., Idaho.
Nomada (Xanthidium) coquilletti Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 567. ¢.

crawfordi crawfordi Cockerell. Colo., Nebr., Wyo.
Nomada (Xanthidium) crawfordi Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 79. 9°.

Taxonomy: Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 487. 2, d.

crawfordi lachrymosa Cockerell. Wyo.
Nomada (Xanthidium) crawfordi lachrymosa Cockerell, 1921. Amer. Mus. Novitates 24: 5.
OF

decempunctata Cockerell. Calif.
Nomada (Xanthidium) decempunctata Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55:

565. 6.
dentariae (Robertson). Ill. to N. Y.. south to Great Smokies Natl. Park.
Xanthidium dentariae Robertson, 1903. Canad. Ent. 35: 178. d.
dilucida Cresson. Colo.
Nomada dilucida Cresson, 1878. Amer. Ent. Soc., Trans. 7: 80. 9.
ednae Cockerell. Colo.
Nomada (Xanthidium) Ednae Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 537. ¢.

electa Cresson. III., Conn., Mass., Va.

Nomada electa Cresson, 1863. Ent. Soe. Phila., Proc. 2: 290. 2, 3.

Nomada (Xanthidium?) mimula Cockerell, 1908. Ent. Soc. Wash., Proc. 10: 66. 2.
electella Cockerell. R. I. to Ga.

Nomada electella Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 612. 2.

elrodi Cockerell. Mont., Nebr., Colo., Tex.
Nomada (Xanthidium) elrodi Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 583. 3.

Taxonomy: Swenk, 1913. Nebr. Univ., Studies 12: 62. 2, d (variety).

Superfamily APOIDEA 2107

festiva Cresson. Mass., N. J.
Nomada festiva Cresson, 1863. Ent. Soc. Phila., Proc. 2: 289. 9.
fragariae Mitchell. Va., N. C., Fla.
Nomada (Heminomada) fragariae Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 391,
fig. 105. ¢.
fragilis fragilis Cresson. Colo., Nebr., N. Mex., Calif.
Nomada fragilis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 79. 6.
fragilis mesosticta Cockerell. Calif.
Nomada fragilis mesosticta Cockerell, 1939. Ann. and Mag. Nat. Hist. (11) 3: 182. 6.
gillettei Cockerell. Colo.
Nomada gillettei Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 81. ¢.
graenicheri Cockerell. Wis.
Nomada (Xanthidium) Graenicheri Cockerell, 1905. Canad. Ent. 37: 189. 2.

Taxonomy: Graenicher, 1911. Pub. Mus. City Milwaukee, Bul. 1: 238. 2, d.

imbricata Smith. N. Y. to N.S. Host: Andrena vicina Sm., Halictus rubicundus (Chr.)?
Nomada imbricata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 246. 2.

Taxonomy: Viereck, 1916. Conn. State Geol. and Nat. Hist. Survey, Bul. 22 (3): 727. 9.

Morphology: Packard, 1897. N. Y. Ent. Soc., Jour. 5: 83-84.

jocularis Cresson. Nev.
Nomada jocularis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 202. °.
laramiensis Swenk. Wyo.
Nomada (Heminomada) laramiensis Swenk, 1913. Nebr. Univ., Studies 12: 75. d.
luteola Olivier. East. United States.
Nomada luteola Olivier, 1811. Encycl. Meth., v. 8, p. 365.
maculiventer Swenk. N. Dak.
Nomada (Heminomada) maculiventer Swenk, 1915. Nebr. Univ., Studies 15: 163. 2, d.
miniata Smith. Ga.
Nomada miniata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 250. 9.
obliterata Cresson. Minn. to N. Y., south to S. C. Host: Andrena vicina Sm.
2Nomada pulchella Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 247. d.
Nomada obliterata Cresson, 1863. Ent. Soc. Phila., Proc. 2: 301. °.
Nomada viburni Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 341. ¢.
ochlerata Mitchell. Mich. (Ann Arbor).
Nomada (?Heminomada) ochlerata Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
396. ¢.
ornithica Cockerell. Colo.
Nomada (Xanthidium) ornithica Cockerell, 1906. Canad. Ent. 38: 161. ¢.
pallidella Cockerell. Colo.
Nomada pallidella Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 82. d.
perivincta perivincta Cockerell. Colo.
Nomada (Xanthidium) perivincta Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 80. d.
perivincta semirufula Cockerell. Colo.
Nomada (Xanthidium) perivincta var. semirufula Cockerell, 1905. Colo. Agr. Expt. Sta.,
Bul. 94: 80. ?.
placitensis Cockerell. N. Mex.
Nomada placitensis Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 446. °.
pseudops Cockerell. Wis. to Mass.
Nomada (Xanthidium) pseudops Cockerell, 1905. Canad. Ent. 37: 189. °.
pyrrha Cockerell. Calif.
Nomada (Xanthidium) pyrrha Cockerell, 1916. Pomona Col. Jour. Ent. Zool. 8: 55. 2.

rhodoxantha Cockerell. Colo.
Nomada (Xanthidium) rhodoxantha Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 78. 2.

2108 Hymenoptera in America North of Mexico

rivalis Cresson. Calif., Wash.
Nomada rivalis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 79. 6.

Taxonomy: Cockerell, 1910. Psyche 17: 97.
ruidosensis Cockerell. N. Mex., Colo.
Nomada ruidosensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 594. d.

sanctaecrucis Cockerell. Calif.
Nomada (Xanthidium) sanctaecrucis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 566.
3

subrutila Lovell and Cockerell. Mich. to New England, ?Colo.

Nomada (Xanthidium) subrutila Lovell and Cockerell, 1905. Psyche 12: 40. 6.
subsimilis Cockerell. Calif.

Nomada (Xanthidium) subsimilis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 567. 3.
suda Cresson. Nev.

Nomada suda Cresson, 1879. Amer. Ent. Soc., Trans. 7: 203. °.
sulphurata Smith. Minn. to Mass., south to Miss. and Ga., ?Utah.

Nomada sulphurata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 249. °.

Nomada luteoloides Robertson, 1895. Amer. Ent. Soc., Trans. 22: 124. ?.

Taxonomy: Rodeck, 1931. Amer. Mus. Novitates 496: 11 (Utah record).

tricurta Swenk. N. Dak.

Nomada (Heminomada) tricurta Swenk, 1915. Nebr. Univ., Studies 15: 11. 2, d.
truttarum Cockerell. N. Mex.

Nomada (Xanthidium) Truttarum Cockerell, 1909. Entomologist 42: 94. 3.
vallesina honorata Cockerell. Colo.

Nomada (Xanthidium) vallesina var. honorata Cockerell, 1922. Ann. and Mag. Nat. Hist.

(9) 10: 269. °.

vallesina vallesina Cockerell. N. Mex., Colo. Host: Andrena irana Ckll.?

Nomada vallesina Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 226. @.

Biology: Hicks, 1934. Colo. Univ., Studies 21: 267 (host).

vernonensis Cockerell. B. C.

Nomada vernonensis Cockerell, 1916. Canad. Ent. 48: 273. 6.
xantholepis Cockerell. Colo.

Nomada (Xanthidium) xantholepis Cockerell, 1911. U.S. Natl. Mus., Proc. 41: 239. ¢.
xanthophila Cockerell. N. Mex.

Nomada xanthophila Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 414. 6.
xanthura Cockerell. N. Y. to N.S.

Nomada xanthura Cockerell, 1908. Ent. Soc. Wash., Proc. 10: 84. 2.

Genus NOMADA Subgenus HOLONOMADA Robertson

Holonomada Robertson, 1903. Canad. Ent. 35: 174, 177.
Type-species: Nomada superba Cresson. Orig. desig.

Revision: Evans, 1972. Wasmann Jour. Biol. 30: 1-34, 5 figs. (included spp.).

Taxonomy: Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 561-563, 580-588 (part). — Viereck
et al., 1905. Canad. Ent. 37: 283-284, 287. —Swenk, 1913. Nebr. Univ., Studies 12: 8, 77-90.
—Swenk, 1915. Nebr. Univ., Studies 15: 169-172. —Mitchell, 1962. N. C. Agr. Expt. Sta.
Tech. Bul. 152: 361-364, fig. 102 (eastern U. S. spp.).

SPECIES GROUP EDWARDSII

edwardsii edwardsii Cresson. B. C., Wash., Colo., Oreg., Calif.; Mexico (Baja California). Host:
Andrena perimelas Ckll.
Nomada Edwardsii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 72. “36” = @.
Nomada intercepta Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 100. d.
Nomada (Holonomada) edwardsii var. australior Cockerell, 1903. Acad. Nat. Sci. Phila.,
Proc. 55: 562. 6.

Superfamily APOIDEA = 2109

Biology: Linsley and MacSwain, 1955. Wasmann Jour. Biol. 13: 270-274, pl. 1, fig. 6; pls. 4 and
5 (habits, host).
edwardsii vinnula Cresson. Nev.
Nomada vinnula Cresson, 1879. Amer. Ent. Soc., Trans. 7: 202. 2, d.
henningeri Evans. N. Mex. (Kenne), Tex. (Olney).
Nomada (Holonomada) henningeri Evans, 1972. Wasmann Jour. Biol. 30: 21. °.
linsleyi Evans. Calif. (Auburn and Westley).
Nomada (Holonomada) linsleyi Evans, 1972. Wasmann Jour. Biol. 30: 26. 2.
pecosensis Cockerell. N. Mex., Colo.
Nomada xanthophila var. pecosensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 585.
OF

Taxonomy: Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 78. d, °.
rhodotricha Cockerell. Calif.
Nomada (Holonomada) rhodotricha Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 562.
OF

SPECIES GROUP SUPERBA

affabilis affabilis Cresson. II]. to Mass., south to Fla.
Nomada affabilis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 74. ¢.

Taxonomy: Robertson, 1895. Amer. Ent. Soc., Trans. 22: 123. 9, d.
affabilis dallasensis Cockerell. Tex and Kans. to D. C.
Nomada affabilis dallasensis Cockerell, 1911. U. S. Natl. Mus., Proc 39: 654. 2, 6.
grandis Cresson. Colo., Tex.
Nomada grandis Cresson, 1875. Rpt. Geog. Geol. Expl. and Survey West of 100th
Meridian, Ch. 7, p. 725. °.
Nomada (Holonomada) magnifica Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 587. °.
hurdi Evans. Calif. (Joshua Tree Natl. Mon. and Kramer Hills).
Nomada (Holonomada) hurdi Evans, 1972. Wasmann Jour. Biol. 30: 22, figs. 1-4. d.
parkeri Evans. Ariz., N. Mex.
Nomada (Holonomada) parkeri Evans, 1972. Wasmann Jour. Biol. 30: 28. °.
superba malvastri Swenk. Nebr.
Nomada (Holonomada) superba malvastri Swenk, 1913. Nebr. Univ., Studies 12: 80. d.
Nomada (Holonomada) nebrascensis Swenk, 1913. Nebr. Univ., Studies 12: 80. 9.
superba superba Cresson. D. C. and N. C., west to Wis. and Colo.
Nomada superba Cresson, 1863. Ent. Soe. Phila., Proc. 2: 281. ¢.

Taxonomy: Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 340. d, °.
SPECIES GROUP UNASSIGNED
hemphilli Cockerell. Wash., Idaho, Oreg., Nev., Calif.; Mexico (Baja California).
Nomada (Holonomada) hemphilli Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 561. d.

Nomada (Holonomada) excurrens Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 562. d.
Nomada (Nomada?) excellens Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 578. 2.

Genus NOMADA Subgenus LAMINOMADA Rodeck

Nomada subg. Laminomada Rodeck, 1947. Ent. Soc. Amer., Ann. 40: 266.
Type-species: Nomada (Holonomada) hesperia Cockerell. Orig. desig. and
monotypic.
hesperia falconis Rodeck. Calif.
Nomada (Laminomada) hesperia falconis Rodeck, 1947. Ent. Soc. Amer., Ann. 40: 270. d,
OF
hesperia hesperia Cockerell. Calif., Oreg., Wash.
Nomada (Holonomada) hesperia Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 563. d.
Nomada (Holonomada) flavopicta Swenk, 1913. Nebr. Univ., Studies 12: 84. °.

Taxonomy: Rodeck, 1947. Ent. Soc. Amer., Ann. 40: 266. d, °.

2110 Hymenoptera in America North of Mexico

Genus NOMADA Subgenus PACHYNOMADA Rodeck

Nomada subg. Pachynomada Rodeck, 1945. Ent. News 56: 180.
Type-species: Nomada (Holonomada) vincta Say. Orig. desig.

adducta Cresson. Colo., Nebr.

Nomada adducta Cresson, 1878. Amer. Ent. Soe., Trans. 7: 73. 6.

Taxonomy: Swenk, 1913. Nebr. Univ., Studies 12: 21. °.

asteris Swenk. Kans.

Nomada (Holonomada) asteris Swenk, 1913. Nebr. Univ., Studies 12: 89. °.
besseyi Swenk. Kans.

Nomada (Holonomada) besseyi Swenk, 1913. Nebr. Univ., Studies 12: 85. 3.
morrisoni flagellaris Cockerell. Colo.

Nomada (Xanthidium) morrisoni flagellaris Cockerell, 1903. Acad. Nat. Sci. Phila., Proce.

5b 58. 2

morrisoni morrisoni Cresson. Colo. Host: Andrena irana Ckll.

Nomada Morrisoni Cresson, 1878. Amer. Ent. Soc., Trans. 7: 72. °.

Biology: Hicks, 1934. Colo. Univ., Studies 21: 267 (host).
victrix Cockerell. Tex.
Nomada victrix Cockerell, 1911. U.S. Natl. Mus., Proc. 39: 657. 9.
vincta heterochroa Cockerell. Colo.
Nomada vincta var. heterochroa Cockerell, 1921. Amer. Mus. Novitates 24: 1. 6.

vineta vineta Say. Kans. and Nebr., east to N. C.
Nomada vincta Say, 1837. Boston Jour. Nat. Hist. 1: 401. 2 (¢ misdet.).

Taxonomy: Robertson, 1903. Canad. Ent. 35: 177. 2, 6. —Mitchell, 1962. N. C. Agr. Expt.
Sta. Tech. Bul. 152: 379-380 (redescription).

zebrata Cresson. Colo., Kans., Nebr., N. Mex., ?Ariz.
Nomada zebrata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 73. 2, 6.

Genus NOMADA Subgenus CALLINOMADA Rodeck

Nomada subg. Callinomada Rodeck, 1945. Ent. News 56: 181.
Type-species: Nomada (Holonomada) antonita Cockerell. Orig. desig.

Taxonomy: Rodeck, 1949. Ent. Soc. Amer., Ann. 42: 174-186. —Mitchell, 1962. N. C. Agr.

Expt. Sta. Tech. Bul. 152: 372-377 (eastern U.S. spp.).

antonita Cockerell. Colo.
Nomada antonita Cockerell, 1909. Canad. Ent. 41: 35. 6.

aquilarum Cockerell. North. Great Plains, east to Wis. and South. Canada, ?N. Mex.
Nomada aquilarum Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 208. 6.
Nomada cockerelli Graenicher, 1911. Pub. Mus. City Milwaukee, Bul. 1: 240. ¢.
Nomada (Holonomada) dacotensis Swenk, 1913. Nebr. Univ., Studies 12: 88. 9.

mutans Cockerell. Oreg., Wash.
Nomada (Holonomada) mutans Cockerell, 1910. Psyche 17: 91. 2.

placida Cresson. Nebr. to N. Y., south to N. C.
Nomada placida Cresson, 1863. Ent. Soc. Phila., Proc. 2: 291. 2, 6.

Taxonomy: Rodeck, 1945. Ent. News 56: 181 (redescription).

rodecki Mitchell. N. C. (Southern Pines).

Nomada (Callinomada) rodecki Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 376.

é.

snowii Cresson. Colo., Nebr., S. Dak., N. Dak.

Nomada Snowii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 75. 2, 3.

Nomada (Holonomada) omahaensis Swenk, 1915. Nebr. Univ., Studies 15: 171. 6.
verecunda Cresson. Oreg., Nev., Calif.

Nomada verecunda Cresson, 1879. Amer. Ent. Soc., Trans. 7: 208. 2, 6.

Superfamily APOIDEA 2111

Genus NOMADA Subgenus MICRONOMADA Cockerell and Atkins

Nomada subg. Micronomada Cockerell and Atkins, 1902. Ann. and Mag. Nat. Hist. (7) 10:
44.
Type-species: Nomada modesta Cresson. Orig. desig. and monotypic.
Cephen Robertson, 1903. Canad. Ent. 35: 174, 176.
Type-species: Nomada texana Cresson. Orig. desig. and monotypic.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 173. —Cockerell, 1903. Acad. Nat. Sci. Phila.,
Proc. 55: 593 (part), 608-611. —Swenk, 1913. Nebr. Univ., Studies 12: 12, 104-112. —Swenk,
1915. Nebr. Univ., Studies 15: 181-188. — Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul.
152: 354-361, figs. 99-101 (eastern U. S. spp.).
amorphae Swenk. Nebr., Colo.

Nomada (Micronomada) amorphae Swenk, 1913. Nebr. Univ., Studies 12: 108. 9, 6.
arenicola Swenk. Nebr.

Nomada (Micronomada) arenicola Swenk, 1913. Nebr. Univ., Studies 12: 107. 2, 6.
belfragei Cresson. Tex., Kans.

Nomada Belfragei Cresson, 1878. Amer. Ent. Soc., Trans. 7: 72. 9.

Nomada belfragei var. xanthogaster Cockerell, 1911. U. S. Nat. Mus., Proce. 39: 644. 2.

Taxonomy: Swenk, 1915. Nebr. Univ. Studies 15: 184. 2, 6.

crucis Cockerell. Ariz., N. Mex., Tex., Colo., Utah.
Nomada crucis Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 212. 6.
fervida Smith. S. C. to Fla., La.
Nomada fervida Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 247. @.
Nomada (Micronomada) crassula Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 610. “2”
=.
formula Viereck. Calif.
Nomada formula Viereck, 1903. Acad. Nat. Sci. Phila., Proc. 54: 728. 2.

Taxonomy: Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 658. ¢.
garciana Cockerell. N. Mex., Tex.
Nomada (Micronomada) garciana Cockerell, 1907. Entomologist 40: 265. ¢.
gutierreziae Cockerell. N. Mex., Ariz., Colo.
Nomada gutierreziae Cockerell, 1896. Canad. Ent. 28: 284. °.
heiligbrodtii Cresson. Minn. to New England, south to Fla. and Tex.
Nomada Heiligbrodtii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 75. °.
Nomada (Micronomada) tiftonensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 610.
é.
Nomada (Micronomada) modesta rivertonensis Cockerell, 1903. Acad. Nat. Sci. Phila.,
Proc. 55: 610. @.
lamarensis Cockerell. Colo., Tex.
Nomada (Micronomada) lamarensis Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 76. ¢.

Taxonomy: Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 652. d, 2.

limata Cresson. Mexico; Tex.
Nomada limata Cresson, 1878. Amer. Ent. Soc., Trans 7: 76. °.
lippiae lippiae Cockerell. N. Mex.
Nomada lippiae Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 214. ¢.
lippiae sublippiae Cockerell. N. Mex.
Nomada lippiae var. sublippiae Cockerell, 1907. Entomologist 40: 265. 6.
melanoptera Cockerell. Colo.
Nomada (Micronomada) melanoptera Cockerell, 1921. Amer. Mus. Novitates 24: 5. °.

mitchelli Cockerell. Tex.
Nomada (Micronomada) mitchelli Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 654. 2, d.

modesta Cresson. N. Y. to Ala.
Nomada modesta Cresson, 1863. Ent. Soc. Phila., Proc. 2: 286. 2, 6.

2112 Hymenoptera in America North of Mexico

Nomada (Micronomada) modesta var. vegana Cockerell, 1903. Acad. Nat. Sci. Phila., Proc.

55: 610. 2.
Nomada (Micronomada) vegana var. nitescens Cockerell, 1905. Colo. Agr. Expt. Sta., Bul.
947 Oree-

neomexicana Cockerell. N. Mex.
Nomada neomexicana Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 209. ?, d.

pallidelutea Swenk. Ariz.; Mexico (Baja California).
Nomada (Micronomada) pallidelutea Swenk, 1915. Nebr. Univ., Studies 15: 32. 2, 6.
Nomada (Micronomada) penisularis Cockerell, 1925. Pan-Pacific Ent. 1: 180. 2, ¢.

putnami Cresson. Utah, Colo., Nebr., Kans., N. Mex.
Nomada Putnami Cresson, 1876. Davenport Acad. Nat. Sci., Proc. 1: 210. 2, “2” = 6.
Taxonomy: Swenk, 1913. Nebr. Univ., Studies 12: 111. 9.

ridingsii Cresson. Colo.
Nomada Ridingsti Cresson, 1878. Amer. Ent. Soc., Trans. 7: 74. ?.

semisuavis Cockerell. Wash., Calif.
Nomada (Micronomada) semisuavis Cockerell, 1910. Psyche 17: 92. 6.
Taxonomy: Cockerell, 1911. U.S. Natl. Mus., Proc. 39: 658. °.

simplicicoxa Swenk. Ariz.
Nomada (Micronomada) simplicicoxa Swenk, 1915. Nebr. Univ., Studies 15: 182. 9, d.

suavis Cresson. Wash., Idaho, Oreg., Calif. Host: Nomia melanderi Ckll.
Nomada suavis Cresson, 1878. Amer. Ent. Soc., Trans. 7: 74. 2 (6 misdet.).
Nomada flavipes Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 426. 2.

texana Cresson. Tex. and La. north to IIl., west to Colo., Ariz.
Nomada texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 271. 2, 6.
Taxonomy: Robertson, 1903. Canad. Ent. 35: 176. 2, ¢.

uhleri Cockerell. Colo.
Nomada (Micronomada) uhleri Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 77. d.

vierecki Cockerell. N. Mex., Tex., Colo., Nebr.; Mexico.
Nomada vierecki Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 211. °, d.
Nomada (Micronomada) vierecki convolvuli Swenk, 1913. Nebr. Univ., Studies 12: 109. 9,

or
Nomada vierecki race cushmani Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 17: 307.

wheeleri engelmanniae Cockerell. Tex.
Nomada wheeleri engelmanniae Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 649. 2 (6
misdet.).

wheeleri wheeleri Cockerell. Tex.
Nomada (Micronomada) wheeleri Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 604. 2.

wisconsinensis Graenicher. Minn., Wis., Mich., IIl.
Nomada wisconsinensis Graenicher, 1911. Pub. Mus. City Milwaukee, Bul. 1: 239.

Genus NOMADA Subgenus CENTRIAS Robertson
Centrias Robertson, 1903. Canad. Ent. 35: 174, 176.
Type-species: Nomada erigeronis Robertson. Orig. desig.

Taxonomy: Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 588, 591-593, 608-612. — Viereck
et al., 1905. Canad. Ent. 37: 283. —Swenk, 1913. Nebr. Univ., Studies 12: 13, 103-104.
—Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 364-365.

erigeronis Robertson. Kans. and Nebr., east to Mass.
Nomada erigeronis Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 341. 9.
Centrias erigerontis(!) Robertson, 1928. Flowers and Insects, Carlinville, Ill., pp. 9, 66, 69.

Taxonomy: Robertson, 1903. Canad. Ent. 35: 176. 9, d.

Genus NOMADA Subgenus NOMADULA Cockerell
Nomada subg. Nomadula Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 611.

Superfamily APOIDEA 2113

Type-species: Nomada articulata Smith. Orig. desig. (=Nomada americana
“Kirby,” Robertson, Cockerell).

Taxonomy: Viereck et al., 1905. Canad. Ent. 37: 285-287. —Swenk, 1913. Nebr. Univ.,
Studies 12: 13, 101-103. —Swenk, 1915. Nebr. Univ., Studies 15: 25-27. —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 365-371 (eastern U.S. spp.).

articulata Smith. N. Dak. and Nebr., east to New England, south to Fla. and La.

¢Nomada americana Kirby, 1837. Fauna Bor.-Amer., p. 269. 2. Type missing; species
uncertain.

Nomada articulata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 248. 6.

Nomada incerta Cresson, 1863. Ent. Soc. Phila., Proc. 2:-309. 2.

Nomada (Centrias) americana dacotana Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55:
592. od.

Nomada (Nomada) bilobata Swenk, 1913. Nebr. Univ., Studies 12: 25. 3, (2 misdet.).

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 310.

australis Mitchell. Fla. to N. C., N. Y., Ind.
Nomada (Nomadula) australis Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 367. 9,
Ge
crotchii crotchii Cresson. Calif.
Nomada Crotchii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 81. 9.

crotchii nigrior Cockerell. Calif.
Nomada (Xanthidium) crotchii var. nigrior Cockerell, 1903. Acad. Nat. Sci. Phila., Proce.
55: 564. °.
Taxonomy: Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 54. 3.

erythrochroa Cockerell. Wash., Idaho.
Nomada erythrochroa Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 203. 9.

erythrospila Cockerell. Calif.
Nomada (Nomada) erythrospila Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 56. 3.

frankei Cockerell. Colo.
Nomada frankei Cockerell, 1929. N. Y. Ent. Soc., Jour. 37: 443. 3.

frieseana Cockerell. Colo.
Nomada Frieseana Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 28. 9.
Taxonomy: Cockerell, 1911. U. S. Natl. Mus., Proce. 41: 237. 6.

jennei Cockerell. Wash.
Nomada Jennei Cockerell, 1906. Canad. Ent. 38: 281. 6.

martinella Cockerell. N. Mex., Colo.
Nomada (Centrias) martinella Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 590. 9.
Taxonomy: Cockerell, 1905. Colo. Agr. Expt. Sta., Bul. 94: 76. 2, d.

melliventris Cresson. Calif.
Nomada melliventris Cresson, 1878. Amer. Ent. Soc., Trans. 7: 82. ¢.

pascoensis Cockerell. Wash., Calif.
Nomada (Xanthidium) pascoensis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 564. ¢.

rohweri aureopilosa Swenk. Colo.
Nomada (Micronomada) aureopilosa Swenk, 1913. Nebr. Univ., Studies 12: 111. 9.
Taxonomy: Swenk, 1915. Nebr. Univ., Studies 15: 32.
rohweri rohweri Cockerell. Colo.
Nomada robweri Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 488. 2, 6.
rubicunda Olivier. Minn. to N. Y., south to Fla.
Nomada rubicunda Olivier, 1811. Encycl. Meth., v. 8, p. 365.
Nomada torrida Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 250. 9.
scita Cresson. Colo., Nebr., N. Dak.
Nomada scita Cresson, 1878. Amer. Ent. Soc., Trans. 7: 77. 6.

Taxonomy: Swenk, 1915. Nebr. Univ., Studies 15: 181. @, ¢.

2114 Hymenoptera in America North of Mexico

scitiformis Cockerell. Oreg.
Nomada (Centrias) scitiformis Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 591. ¢.

semiscita Cockerell. Colo.
Nomada semiscita Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 28. 6.

seneciophila Mitchell. N. C., Fla.
Nomada (Nomadula) seneciophila Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
370. 2, d.

sophiarum Cockerell. N. Mex.
Nomada sophiarum Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 213. 6.

Genus HEXEPEOLUS Linsley and Michener

Hexepeolus Linsley and Michener, 1937. Pan-Pacifie Ent. 13: 77.
Type-species: Hexepeolus mojavensis Linsley and Michener. Orig. desig.

Revision: Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: 287-289. —Michener, 1944.
Amer. Mus. Nat. Hist., Bul. 82: 274.

mojavensis Linsley and Michener. Calif.
Hexepeolus mojavensis Linsley and Michener, 1937. Pan-Pacific Ent. 13: 78. 2, 3.

rhodogyne Linsley and Michener. Calif. Host: Ancylandrena larreae Timb.?
Hexepeolus rhodogyne Linsley and Michener, 1937. Pan-Pacific Ent. 13: 80. 2, ¢.

Genus PARANOMADA Linsley and Michener

Paranomada Linsley and Michener, 1937. Pan-Pacifie Ent. 13: 82.
Type-species: Paranomada nitida Linsley and Michener. Monotypic and orig. desig.

The species of this genus may be cleptoparasites in the nests of Exomalopsis.

Revision: Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: 289-291 (genera).
—Linsley, 1943. Amer. Ent. Soc., Trans. 69: 103-104 (species keys). —Michener, 1944.
Amer. Mus. Nat. Hist., Bul. 82: 274 (genera). —Linsley, 1945. Ent. News 56: 149-151
(species keys).

californica Linsley. Calif.
Paranomada californica Linsley, 1945. Ent. News 56: 149. 9, 6.

nitida Linsley and Michener. Ariz.
Paranomada nitida Linsley and Michener, 1937. Pan-Pacific Ent. 13: 88. 9.

velutina Linsley. Ariz.; Mexico (Baja California).
Paranomada velutina Linsley, 1939. Pan-Pacific Ent. 15: 10. 3.

Taxonomy: Linsley, 1943. Amer. Ent. Soc., Trans. 69: 105. 2, ¢.
Genus HESPERONOMADA Linsley

Hesperonomada Linsley, 1939. Pan-Pacific Ent. 15: 5.
Type-species: Hesperonomada melanantha Linsley. Monotypic and orig. desig.
Revision: Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: 291-292.

melanantha Linsley. Calif. Host: ?7Exomalopsis spp.
Hesperonomada melanantha Linsley, 1939. Pan-Pacifie Ent. 15: 6. 2, d.

Genus TRIOPASITES Linsley

Triopasites Linsley, 1939. Pan-Pacific Ent. 15: 8.
Type-species: T'riopasites timberlakei Linsley. Monotypic and orig. desig.
Revision: Linsley and Michener, 1939. Amer. Ent. Soc., Trans. 65: 293-294. —Linsley, 1943.
Amer. Ent. Soc., Trans. 69: 99-103.
laguna Linsley. Calif.; Mexico (Baja California).
Triopasites laguna Linsley, 1943. Amer. Ent. Soc., Trans. 69: 102. d.

micheneri Linsley. Ariz.
Triopasites micheneri Linsley, 1943. Amer. Ent. Soc., Trans. 69: 101. °.

Superfamily APOIDEA 2115

pasitura (Cockerell). Tex. Host: Exomalopsis compactula (Ckll.)?

Nomada pasitura Cockerell, 1935. Amer. Mus. Novitates 766: 6. d.
penniger (Cockerell). N. Mex.

Nomada penniger Cockerell, 1894. Ent. News 5: 235. 9.
timberlakei Linsley. Calif.

Triopasites timberlakei Linsley, 1939. Pan-Pacifie Ent. 15: 9. °.

Genus MELANOMADA Cockerell

Melanomada Cockerell, 1903. Acad. Nat. Sci. Phila., Proc. 55: 587.
Type-species: Nomada grindeliae Cockerell. Monotypic and orig. desig.

grindeliae (Cockerell). Nebr., Mont.

Nomada grindeliae Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 210. 3.
heleniella (Cockerell). Tex., Nebr., Kans.

Nomada (Melanomada) heleniella Cockerell, 1911. U. S. Natl. Mus., Proc. 39: 648. ?, 6.
sidaefloris (Cockerell). N. Mex.

Nomada pennigera var. sidaefloris Cockerell, 1898. N. Mex. Univ., Bul. 1: 59. 3.

SUBFAMILY ANTHOPHORINAE

This is a large assemblage of both pollen-collecting and parasitic bees. It is found on all the
continents although it is especially well represented in the Holarctic, Ethiopian and Neotropical
regions. The current classification recognizes about a dozen tribes of which seven are
represented in America north of Mexico. Insofar as known, all of the pollen-collecting species
line their cells with a waxlike substance.

TRIBE EXOMALOPSINI

This tribe consists of several American genera which are chiefly or entirely Neotropical in oc-
currence. Only the genera Ancyloscelis and Exomalopsis extend northward into the United
States. Close relatives of the Exomalopsini include two tribes, the Ancylini (Ancyla and Tar-
salia) which inhabit the more arid areas of the Palaearctic and the Tetrapediini (Tetrapedia)
which live in the tropics of the Neotropical Region. While most of the studied Exomalopsini nest
in the ground and are colonial, if not communal, at least some species of the genus
Paratetrapedia, like the Tetrapediini, make their nests in wood.

Taxonomy: Michener and Moure, 1957. Amer. Mus. Nat. Hist., Bul. 112: 395-452, 91 figs.
(generic classification).

Biology: Torchio, 1974. Kans. Ent. Soc., Jour. 47: 54-63, 21 tables (nest architecture,
comparative behavior).

Genus EXOMALOPSIS Spinola

Although primarily a genus of Neotropical bees, there are several subgenera represented in
America north of Mexico, especially in the more arid areas of the southwestern United States.
The species of Paranomada, Hesperonomada and Triopasites may be cleptoparasitie in the
nests of these bees.

Revision: Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 85-106. —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 235-240, figs. 68-69 (eastern U. S. spp.).

Genus EXOMALOPSIS Subgenus EXOMALOPSIS Spinola

Exomalopsis Spinola, 1853. Accad. Sci. Torino, Mem. (2) 13: 89.
Type-species: Exomalopsis aureopilosa Spinola. Desig. by Taschenberg, 1883.
(=Exomalopsis fulvopilosa Spinola).
?E pimonispractor Holmberg, 1903. Buenos Aires Mus. Nac. de Hist. Nat., An. (3) 2: 426.
Type-species: Epimonispractor gratiosus Holmberg. Orig. desig.
birkmanni Cockerell. Tex. Pollen: Unknown, but visits flowers of Antigonon leptopus.
Exomalopsis birkmanni Cockerell, 1922. Ann. and Mag. Nat. Hist. (9) 10: 265. 9.

2116 Hymenoptera in America North of Mexico

pulchella Cresson. Southern Fla.; Cuba.
Exomalopsis pulchella Cresson, 1865. Ent. Soc. Phila., Proc. 4: 191. 2, 3.
similis Cresson. Southern Fla.; Cuba, Jamaica, Costa Rica. Pollen: Polylectic, visits a wide
variety of flowers for nectar and pollen including Antigonum leptopus, Borreria laevis,
Citrus, Cleome, Cocos nucifera, Comelina, Kallstroemia, Ludwigia, Malpighia
punicifolia, Mimosa pudica, Piriqueta cistoides, Psidium guajava, Rhynchospora
nervosa, Sida, Solanum, Stachytarpheta, Teliostachya alopecuroidea, Trimezia.
Exomalopsis similis Cresson, 1865. Ent. Soc. Phila., Proc. 4: 191. @.
Exomalopsis penelope Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 161. 2, ¢.
Exomalopsis ogilviei Cockerell, 1938. Entomologist 71: 281. 2.

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 109 (synonymy).

Biology: Raw, 1977 (1976). Biotropica 8:270-277, 2 figs., 2 tables (seasonal changes in
numbers and foraging activities).
zexmeniae Cockerell. South. Tex. to Panama. Pollen: Collects pollen from flowers of grasses,
composites and Solanaceae.
Exomalopsis zexmeniae Cockerell, 1912. Ent. News 23: 447. 9.
Exomalopsis azulensis Cockerell, 1949. U.S. Natl. Mus., Proc. 98: 451. 2.

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 108, figs. 62-64 (synonymy,
taxonomic characters, floral relationships).

Genus EXOMALOPSIS Subgenus PHANOMALOPSIS Michener and Moure

Exomalopsis subg. Phanomalopsis Michener and Moure, 1957. Amer. Mus. Nat. Hist., Bul.
112: 430.
Type-species: Exomalopsis jenseni Friese. Orig. desig.
snowi Cockerell. South. Tex.
Exomalopsis snowi Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 18: 73. 6.
solani Cockerell. Tex., N. Mex., Colo (Pueblo), Ariz.; Mexico (Zacatecas). Pollen: Polylege of a
wide variety of flowers including Acacia, Baileya, Cassia, Chrysothamnus, Chilopsis
linearis, Cleome, Croton, Eriogonum, Flaveria, Haplopappus, Kallstroemia, Larrea,
Petalostemon, Solanum, Sphaeralcea, Verbesina.
Exomalopsis solani Cockerell, 1896. Canad. Ent. 28: 25. 9.

Biology: Linsley, MacSwain and Smith, 1954. South. Calif. Acad. Sci., Bul. 55: 83-101.
—Linsley, MacSwain and Smith, 1954. Pan-Pacific Ent. 30: 263-264 (nest). —Michener,
1966. Kans. Ent. Soc., Jour. 39: 315-317 (cooperative nest provisioning). —Hurd and
Linsley, 1975. Smithsn. Contrib. Zool. 193: 40 (nest sites, floral relationships).

solidaginis Cockerell. N. Mex., Ariz., south. Calif., desert. Pollen: Unknown, but visits flowers
of Gutierrezia lucida, Heterotheca, Lippia wrightti, Solidago canadensis arizonica.
Exomalopsis solidaginis Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 452. d.
Exomalopsis verbesinae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 21. @.

Taxonomy: Cockerell, 1907. Ann. and Mag. Nat. Hist. (7) 19: 539. 9, 6.
Genus EXOMALOPSIS Subgenus ANTHOPHORULA Cockerell

Anthophorula Cockerell, 1897. N. Mex. Agr. Expt. Sta., Bul. 24: 44.
Type-species: Anthophorula compactula Cockerell. Monotypic.

Diadasiella Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 64.
Type-species: Synhalonia albicans Provancher. Monotypic and orig. desig.

(=Diadasiella coquilletti Ashmead).

Exomalopsis subg. Pachycerapis Cockerell, 1922. Amer. Mus. Novitates 47: 4.

Type-species: Exomalopsis cornigera Cockerell. Orig. desig.
albata Timberlake. Utah., Ariz., south. Calif., desert. Pollen: Unknown, but visits flowers of
Eriogonum fasciculatum, E. reniforme, E. trichopes, Gutierrezia lucida.
Exomalopsis (Anthophorula) albata Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 92. 2, ¢.

Superfamily APOIDEA 2117

albicans (Provancher). Oreg., Calif. Pollen: Unknown, but visits flowers of Brassica
campestris, B. incana, Cryptantha intermedia, Eriastrum, Eriogonum fasciculatum,
Euphorbia, Eschscholzia californica, Stephanomeria virgata, Trichostema lanceolatum.
Synhalonia albicans Provancher, 1896. Nat. Canad. 23: 27. d.
Diadasiella coquilletti Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 64. 6.
albovestita Timberlake. South. Calif., desert. Pollen: Unknown, but visits flowers of
Cryptantha barbigera, Dalea emoryi, Eriogonum trichopes, Petalonyx thurberi.
Exomalopsis (Anthophorula) albovestita Timberlake. 1947. N. Y. Ent. Soc., Jour. 55: 98. d.
cerei Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Cereus engelmanni.
Exomalopsis (Anthophorula) cerei Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 100. 2.
compactula (Cockerell). Tex., N. Mex., Ariz. Parasite: Triopasites pasitura (Ckll.)? Pollen:
Unknown, but visits flowers of Aplopappus spinulosus, Convolvulus hermannioides,
Dithyraea wislizenii, Grindelia, Phacelia congesta, Verbesina.
Anthophorula compactula Cockerell, 1897. N. Mex. Agr. Expt. Sta., Bul. 24: 44. 9.

Taxonomy: Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 415. 2, 6.
completa Cockerell. Tex., N. Mex. Pollen: Unknown, but visits flowers of Convolvulus
hermannioides, Phacelia congesta.
Exomalopsis (Anthophorula) compactula var. completa Cockerell, 1935. Amer. Mus.
Novitates 766: 5. 2, 6.
cornigera Cockerell. Ariz.
Exomalopsis (Pachycerapis) cornigera Cockerell, 1922. Amer. Mus. Novitates 47: 5. do.
deserticola Timberlake. South. Calif., desert. Pollen: Unknown, but visits flowers of Asclepias
erosa, Chilopsis linearis, Cleomella obtusifolia, Eriogonum reniforme, Heliotropium
spathulatum.
Exomalopsis (Anthophorula) deserticola Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 95. 2,
3.

eriogoni Timberlake. South. Calif., deserts. Pollen: Unknown, but visits flowers of Cleomella
obtusifolia, Eriogonum reniforme, E. trichopes, Heliotropium spathulatum, Hugelia
virgata, Pectis papposa.
Exomalopsis (Anthophorula) eriogoni Timberlake, 1947. N. Y. Ent. Soe., Jour. 55: 94. °,
é.
euphorbiae Timberlake. Ariz., Calif. Pollen: Unknown, but visits flowers of Boerhaavia,
Euphorbia.
Exomalopsis (Anthophorula) euphorbiae Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 102.
noe
gutierreziae Timberlake. Ariz. Pollen: Unknown, but visits flowers of Gutierrezia.
Exomalopsis (Anthophorula) gutierreziae Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 97.
OF

palmarum Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Eriogonwm
inflatum.
Exomalopsis (Anthophorula) palmarum Timberlake, 1947. N. Y. Ent. Soce., Jour. 55: 99. 3.
rufiventris Timberlake. West. Tex., Ariz. Pollen: Unknown, but visits flowers of Boerhaavia,
Tidestromia lanuginosa.
Exomalopsis (Anthophorula) rufiventris Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 101.
Oral
torticornis Cockerell. South. Calif. Pollen: Collects pollen from flowers of Hugelia virgata, but
also visits flowers of Calochortus kennedyi, Centaurea melitensis, Chaenactis
glabriuscula, Chorizanthe parryi, Convolvulus arvense, Cryptantha intermedia, Encelia
farinosa, Eriogonum fasciculatum polifolii, Eschscholzia californica, Gutierrezia
californica, Hemizonia tenella, Lotus scoparius, L. strigosus, Marrubium vulgare,
Mentzelia lindleyi, Navarretia intertexta, Opuntia vaseyi, Sphaeralcea ambigua.
Exomalopsis torticornis Cockerell, 1927. Ent. Soc. Amer., Ann. 20: 399. 6.

Biology: Hicks, 1936. Canad. Ent. 68: 47 (nest).

varleyi Timberlake. South. Calif., desert (Warren’s Well).
Exomalopsis (Anthophorula) varleyi Timberlake, 1947. N. Y. Ent. Soce., Jour. 55: 96. 3.

2118 Hymenoptera in America North of Mexico

Genus EXOMALOPSIS Subgenus ANTHOPHORISCA Michener and Moure

Exomalopsis subg. Anthophorisca Michener and Moure, 1957. Amer. Mus. Nat. Hist., Bul.
112: 433.
Type-species: Melissodes pygmaea Cresson. Orig. desig.
asteris Mitchell. Tex., Ind. Pollen: Unknown, but visits flowers of Aster.
Exomalopsis (Anthophorisca) asteris Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
236, fig. 68. 2, d.
chionura Cockerell. Cent. Calif. Pollen: Stores pollen of Grindelia camporum.
Exomalopsis (Anthophorula) chionura Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 203.
ORIG

Taxonomy: Rozen, 1957. Ent. Soc. Amer., Ann. 50: 469-475 (larva).

Biology: Rozen and MaeNeill, 1957. Ent. Soc. Amer., Ann. 50: 522-529 (nest, life history).
chlorina Cockerell. N. Mex. Pollen: Unknown, but visits flowers of Sphaeralcea.
Exomalopsis chlorina Cockerell, 1918. Ann. and Mag. Nat. Hist. (9) 2: 477. @.
micheneri Timberlake. Miss. Pollen: Unknown, but visits flowers of Gerardia.
Exomalopsis (Anthophorula) micheneri Timberlake, 1947. N. Y. Ent. Soc., Jour. 55: 105.
OPaor
morgani (Cockerell). Tex. Pollen: Unknown, but visits flowers of Helianthus.
Anthophorula morgani Cockerell, 1914. Entomologist 47: 114. 9.
nitens Cockerell. South. Calif. Pollen: Unknown, but visits flowers of Aster, Calochortus
splendens, Calycadenia multiglandulosa, Chlorogalum parviflorum, Brodiaea crocea, B.
isioides, Grindelia camporum, G. rubriflorum, Hemizonia fasciculata, Lessingia
germanorum, Lotus scoparius, Malacothamnus arcuatus, Malvastrum fasciculatum,
Navarretia heterodoxa, Opuntia littoralis, O. parryi.
Exomalopsis nitens Cockerell, 1915. Pomona Col. Jour. Ent. Zool. 7: 231. 9.
pygmaea (Cresson). Nebr., Colo., Tex. Pollen: Unknown, but visits flowers of Helianthus
annuus.
Melissodes pygmaea Cresson, 1872. Amer. Ent. Soc., Trans. 4: 279. °.
Exomalopsis Bruneri Crawford, 1902. Canad. Ent. 34: 238. 2, ¢.

Biology: Schwarz, 1896. Ent. Soc. Wash., Proc. 4: 24-26 (sleeping habits).
sidae Cockerell. Tex., N. Mex., Ariz., South. Calif., Mexico (Baja California). Pollen: Apparently
an oligolege of the Malvaceae, visiting flowers of Sida hederacea, Sphaeralcea.
Predator: Philanthus arizonicus Bohart.
Exomalopsis sidae Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 160. 2, d.
texana Friese. Tex. Pollen: Unknown, but visits flowers of Heleniwm tenuifolium.
Exomalopsis texana Friese, 1899. K. K. Naturhist. Hofmus., Ann. 14: 264. @, 6.

Genus ANCYLOSCELIS Latreille

Ancyloscelis Latreille, 1829. In Cuvier, Le Regne Animal, ed. 2, v. 5, p. 355.
Type-species: Ancylosceles ursinus Haliday. First included species.
Ancylosceles(!) Haliday, 1837. Linn. Soc. London, Trans. 17: 320.
Dipedia Friese, 1906. Flora og Fauna, v. 8, p. 92.
Type-species: Ancyloscelis armatus Smith. Desig. by Lutz and Cockerell, 1920
(=Chalcis apiformis Fabricius).

All species visit flowers of Ipomoea for pollen and nectar, but also visit flowers of other plants
for nectar.

Revision: Michener, 1942. Pan-Pacific Ent. 18: 108-113.
apiformis (Fabricius). Brazil to Tex. Parasite: Anthrax limatulus Say, Monodontomerus
mexicanus Gahan, Trophocleptria schraderi Mich.?
Chalcis apiformis Fabricius, 1793. Ent. System., p. 195. 3.
Ancyloscelis armatus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 367. 6.
Nomia tarsalis Westwood, 1875. Ent. Soc. London, Trans. p. 221, pl. 5, fig. 5.
Melissodes toluca Cresson, 1878. Acad. Nat. Sci. Phila., Proc., p. 219. 2 (6 misdet.).

Superfamily APOIDEA 2119

Exomalopsis chionocincta Cockerell, 1949. U. S. Natl. Mus., Proc. 98: 453. 9.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 325. —Michener, 1954. Amer. Mus.
Nat. Hist., Bul. 104: 110, figs. 50-52 (synonymy, characters). —Moure, 1960. Studia Ent. 3:
114-115 (synonymy, notes on type).

Biology: Michener, 1974. Kans. Ent. Soce., Jour. 47: 19-22, 4 figs. (nest architecture, nests
intermingled with nests of Melitoma ?euglossoides; reported as ?armatus). —Torchio,
1974. Kans. Ent. Soc., Jour. 47: 54-63, 2 tables (nest architecture, nests intermingled with
nests of Melitoma ?euglossoides, life history, associates; reported as ?armatus).

sejunctus Cockerell. Tex., Colo., Ariz.
Ancylosceles sejuncta Cockerell, 1933. Amer. Mus. Novitates 595: 1. 3.
Ancylosceles maculifera Cockerell, 1934. Amer. Mus. Novitates 697: 8. 3.

Taxonomy: Cockerell, 1935. Amer. Mus. Novitates 766: 4. ?.
TRIBE MELITOMINI

This tribe is found only in the New World and is represented in America north of Mexico by
the genera Diadasia, Melitoma and Ptilothrix. Although species of these genera occupy both
continents, only the genus Diadasia contains numerous species in the more arid parts of North
and South America. As a group the Melitomini have established a specialized dependency upon
the pollens of a limited number of plant families including notably only certain genera in the
Cactaceae, Compositae, Convolvulaceae, Malvaceae and Onagraceae.

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 128 (generic classification).

Biology: Linsley, MacSwain and Smith, 1956. South. Calif. Acad. Sci., Bul. 55: 83-101, pl. 26
(biological characters). —Linsley and MacSwain, 1952. Calif. Univ. Pubs. Ent. 9: 267-290,
pls. 1-6 (biological relationships with Anthophorini).

Genus PTILOTHRIX Smith

Ptilothrix Smith, 1853. Cat. Hym. Brit. Mus., v. 1, p. 131.
Type-species: Ptilothrix plumatus Smith. Monotypic.
Ptilothryx Marschall, 1873. Nomenclator Zoologicus, p. 269. Emend.
Emphor Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 476.
Type-species: Melissodes bombiformis Cresson. Monotypic and orig. desig.
Energoponus Holmberg, 1903. Mus. Nac. Buenos Aires, An. (8) 2: 406.
Type-species: Ptilothrix plumatus Smith. Desig. by Sandhouse, 1943.
(=Energoponus strenuus Holmberg).

Taxonomy: Moure, 1947. Soc. Ent. Argentina, Rev. 13: 24 (synonymy).
bombiformis (Cresson). Kans. and IIl. to N. J., south to Tex. and Fla.; Mexico. Pollen: Stores
pollen of Hibiscus, but also visits flowers of Althaea, Cephalanthus, Cirsium, Ipomoea,
Verbena, Vernonia for nectar. 7
Melissodes bombiformis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 219. 2, 6.
Emphor fuscojubatus Cockerell, 1913. Psyche 20: 107. @.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1075, figs. 212-218 (larva). —Mitchell,
1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 240-242, fig. 67 (synonymy, redescription).

Biology: Robertson, 1890. Canad. Ent. 22: 217. —Knab, 1911. Ent. Soc. Wash., Proc. 13: 170.
—Grossbeck, 1911. N. Y. Ent. Soc., Jour. 19: 238. —Nichols, 1913. Psyche 20: 107 (as
fuscojubatus). —Robertson, 1914. Ent. News 25: 70. —Robertson, 1918. Canad. Ent. 50:
320. —Robertson, 1925. Psyche 32: 278. —Davis, 1926. Brooklyn Ent. Soc., Bul. 21: 127.
—Rau, 1930. Brooklyn Ent. Soc., Bul. 25: 28. —Rau, 1934. Acad. Sci. St. Louis, Trans. 28:
222. —Michener, 1947. Amer. Midland Nat. 38: 451 (habits).

sumichrasti (Cresson). Ariz. (Marana and Sahuarita); Mexico. Parasite: Dasymutilla
connectens (Cameron)?, D. eminentia Mickel?, D. foxi (Ckll.)?, D. phoenix (Fox)?,
Lytta variabilis Duges, Nemognatha chrysomeloides L. , Pseudomethoca praeclara
(Blake)?, Pollen: Stores pollen of Gossypium (cultivated), Ipomoea longifolia, I. pringlei,

2120 Hymenoptera in America North of Mexico

Kallstroemia grandiflora, but visits other flowers like Tribulus terrestis presumably for

nectar.
Melissodes sumichrasti Cresson, 1878. Acad. Nat. Sci. Phila., Proc., p. 218. 2, ¢.

Biology: Linsley, MacSwain and Smith, 1956. South. Calif. Acad. Sci., Bul. 55: 83-101, pl. 26,
figs. 5, 10 (nest, larva, life history). —Butler, 1967. Pan-Pacific Ent. 43: 8-14 (nest, life
history). —Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 13 (nest site, floral
relationships).

Genus DIADASIA Patton

Diadasia Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 475.
Type-species: Melissodes enavata Cresson. Orig. desig.
Dasiapis Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 450.
Type-species: Dasiapis ochracea Cockerell. Monotypic.
Leptometria Holmberg, 1903. Mus. Nac. Buenos Aires, An. (3) 2: 409.
Type-species: Leptometria pereyrae Holmberg. Desig. by Brethes, 1910.
Coquillettapis Viereck, 1909. Ent. Soc. Wash., Proce. 11: 47.
Type-species: Melissodes nigrifrons Cresson. Monotypic and orig. desig.
(=Coquillettapis melittoides Viereck).
Diadasina Moure, 1950. Dusenia 1: 392.
Type-species: Melitoma paraensis Ducke. Orig. desig.
Diadasiana(!) Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 129.

Bees of this genus occur only in the Americas and are distributed chiefly in the warm tem-
perate areas of North and South America, although a few species are present in the moist trop-
ics. The species are oligolectic, obtaining pollen primarily from the Malvaceae (Callirhoe, Sida,
Sidalcea, Sphaeralcea), Convolvulaceae (Calystegia, Convolvulus), Compositae (Helianthus),
Cactaceae (Opuntia) and the Onagraceae (Clarkia). The known parasites include species of the
family Bombyliidae (Anthrax, Villa), Meloidae (Lytta, Tetraonyx), Mutillidae (Dasymutilla,
Sphaeropthalma), and Rhipiphoridae (Rhipiphorus). The bee genus Protepeolus is a clep-
toparasite of at least one species of Diadasia.

Revision: Cockerell, 1905. Amer. Nat. 39: 741-743. —Timberlake, 1941. Brooklyn Ent. Soc.,
Bul. 36: 2-11 (key).

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 129-131, table 2 (synonymy,
subgeneric characters).

Biology: Lutz and Cockerell, 1920. Amer. Mus. Nat. Hist., Bul. 42:583-587 (floral and seasonal
records). —Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 199-235, 2 pls., 2 tables
(nesting habits, nest architecture, floral relationships, parasites). —Linsley and MacSwain,
1958. Evolution 12: 219-223 (floral constancy). —Schlissing, 1970. Ecology 51: 1061-1067
(sequence and timing of foraging). —Cazier and Linsley, 1974. Amer. Mus. Novitates 2546:
14 (floral relationships). —EHickwort, Eickwort and Linsley, 1977. Kans. Ent. Soc., Jour.
50:1-17, 9 figs., 3 tables (nest aggregations).

afflicta afflicta (Cresson). N. Mex., Tex. Pollen: Collects pollen from flowers of Callirhoe.
Melissodes afflicta Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 217. 2, 6.

Biology: Snyder, Barrows and Chabot, 1976. Kans. Ent. Soc., Jour. 49: 200-208, 2 figs. (nest
architecture, pollen mass, insertion of egg).
afflicta perafflicta Cockerell. Kans. Pollen: Collects pollen from flowers of Callirhoe.
Diadasia afflicta perafflicta Cockerell, 1905. Amer. Nat. 39: 744. d, 9.
afflictula Cockerell. N. Mex.
Diadasia afflictula Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 366. d.
albovestita Provancher. Calif.
Diadasia albovestita Provancher, 1896. Nat. Canad. 23: 27. 9.
Didasia(!) alboresta(!) Fowler, 1899. Canad. Ent. 31: 285.
angusticeps Timberlake. Calif. Pollen: Collects pollen from flowers of Clarkia.
Diadasia angusticeps Timberlake, 1939. Brooklyn Ent. Soc., Bul. 34: 15. 6, @.

Superfamily APOIDEA 2121

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 201 (flower relationships).
—MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 26-29 (flower
relationships).

australis australis (Cresson). Colo., N. Mex., Tex., Ariz. Pollen: Collects pollen from flowers of
Opuntia.
Melisiodes australis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 214. °, d.
australis californica Timberlake. Calif., Colo., N. Mex., Calif.; Mexico (Zacatecas). Pollen:
Collects pollen from flowers of Cactaceae, principally Opuntia. Other subspecies of

australis are known from Mexico. :
Diadasia australis californica Timberlake, 1940. Brooklyn Ent. Soc., Bul. 35: 28. 2, d.

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 201-203.
bituberculata (Cresson). Calif. Parasite: Lytta melaena (LeC.), Rhipiphorus diadasiae Linsley
and MacSwain, Sphaeropthalma auraria (Blake), S. unicolor (Cress.), Villa apicola Cole,
V. tricellula Cole. Pollen: Collects pollen from flowers of the introduced Convolvulus
arvensis as well as from the native species of Calystegia and Convolvulus.
Melissodes bituberculata Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 218. d.
Didasia(!) cinerea Fowler, 1899. Canad. Ent. 31: 285. ¢.

Biology: Linsley and MacSwain, 1952. Pan-Pacific Ent. 28: 131-135. —Linsley and MacSwain,
1957. Wasmann Jour. Biol. 15: 203. —Schlising, 1972. Pan-Pacifie Ent. 48: 175-188, 1 fig., 3
tables (foraging and nest provisioning behavior).

consociata Timberlake. Calif., Nev., Ariz. Parasite: Anthrax nidicola Cole, Rhipiphorus
dammersi Barber, R. smithi Linsley and MacSwain, Villa apicola Cole. Pollen: Collects
pollen principally from flowers of Sida and may also collect pollen from flowers of

Sphaeralcea.
Diadasia consociata Timberlake, 1939. Brooklyn Ent. Soc., Bul. 34: 11. d, 9.

Biology: Linsley, MacSwain and Smith, 1952. Calif. Univ. Pubs. Ent. 9: 267-290, 6 pls.
(bionomics). —Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 203, 205, pl. 1, fig. 1
(nesting habits, flower relationships, parasites).

diminuta (Cresson). B. C. to Calif., east to Nebr., Kans. and Tex.; Mexico. Ecology: Nests are
sometimes usurped by Osmia seclusa Sandhouse. Parasite: Dasymutilla foxi (Ckll.),
Rhipiphorus sexdens Linsley and MacSwain, Villa perplexa (Cogq.). Pollen: Collects
pollen from flowers of Sphaeralcea and Kallstroemia grandiflora.
Melissodes diminuta Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 215. ¢.
Melissodes apacha Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 217. 2.

Biology: Bohart, 1955. Ent. Soc. Wash., Proc. 57:235-236 (nest usurpation). —Linsley and
MacSwain, 1957. Wasmann Jour. Biol. 15: 205-211, pl. 2. (nesting habits, flower
relationships, parasites). —Eickwort, Eickwort and Linsley, 1977. Kans. Ent. Soc., Jour.
50:1-17, 9 figs., 3 tables (nest aggregation, life history).

enavata (Cresson). Kans., Colo., N. Mex., Tex., Utah, Ariz., Calif; Mexico. Parasite: Anthrax

nidicola Cole, Dasymutilla fulvohirta (Cress.). Pollen: Collects pollen from flowers of
Compositae, principally those of Helianthus. Diadasia tricincta listed in synonymy
below is almost certainly a synonym of Synhalonia actuosa (Cress.), see Timberlake,
1969. Calif. Univ. Pubs. Ent. 57: 13.

Melissodes enavata Cresson, 1872. Amer. Ent. Soc., Trans. 4: 280. 9.

Melissodes ?ursina Cresson, 1872. Amer. Ent. Soc., Trans. 4: 281. d.

Melissodes densa Cresson, 1872. Amer. Ent. Soe., Trans. 4: 282. ¢.

Eucera arctos Dalla Torre, 1896. Cat. Hym., v. 10, p. 225. Proposed to replace ursina,
preoccupied.in Eucera in Dalla Torre’s sense.

?Diadasia 3-cincta Provancher, 1896. Nat. Canad. 23: 28. 2.

?Diadasia tricincta Fowler, 1899. Canad. Ent. 31: 286. Emend.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1075, figs. 206-211 (larva).
Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 212, pl. 1, fig. 5 (nesting
habits, flower relationships, parasites).

friesei Cockerell. ?Calif., Ariz.
Diadasia friesei Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 192. 2.

2122 Hymenoptera in America North of Mexico

laticauda Cockerell. Calif. Pollen: Collects pollen from flowers of Malvastrum and
Sphaeralcea.
Diadasia laticauda Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 103. 9.
Diadasia crassicauda Cockerell, 1915. Pomona Jour. Ent. Zool. 7: 231. 6.

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 219 (floral relationships).
lutzi deserticola Timberlake. Calif. (Chocolate Mts.). Pollen: Collects pollen from flowers of
Sphaeralcea.
Diadasia lutzi deserticola Timberlake, 1940. Brooklyn Ent. Soc., Bul. 35: 23. 3, 2.
lutzi difficilis Timberlake. Ariz., Calif.; Mexico (Baja California). Pollen: Collects pollen from
flowers of Sphaeralcea.
Diadasia lutzi difficilis Timberlake, 1940. Brooklyn Ent. Soc., Bul. 35: 22. 6, @.
lutzi lutzi Cockerell. Wyo., N. Mex., Ariz., Calif.; Mexico (Baja California). Pollen: Collects
pollen from flowers of Sphaeralcea.
Diadasia lutzi Cockerell, 1924. Amer. Mus. Novitates 113: 2. 6, 2.

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 212-213, pl. 1, fig. 3 (nesting
habits, flower relationships).
martialis Timberlake. South. Calif., Ariz.; Mexico (Baja California). Pollen: Collects pollen from
flowers of Sphaeralcea.
Diadasia martialis Timberlake, 1940. Brooklyn Ent. Soe., Bul. 35: 25. 3, @.
megamorpha Cockerell. N. Mex. Pollen: Collects pollen from flowers of Sphaeralcea and
Kallstroemia grandiflora.
Diadasia megamorpha Cockerell, 1898. Amer. Ent. Soc., Trans. 25: 192. 6, @.
nigrifrons (Cresson). Idaho, Calif. Pollen: Collects pollen from flowers of Sidalcea.
Melissodes nigrifrons Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 195. 9.
Didasia(!) nerea Fowler, 1899. Canad. Ent. 31: 285. °, 6.
Coquillettapis melittoides Viereck, 1909. Ent. Soc. Wash., Proc. 11: 48. 2.
Diadasia nigrifrons var. epileuca Cockerell, 1925. Calif. Acad. Sci., Proc. (4) 14: 203. 9°.

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 215.
nitidifrons Cockerell. Wash., Calif., Utah. Pollen: Collects pollen from flowers of Sphaeralcea.
Diadasia nitidifrons Cockerell, 1905. South. Calif. Acad. Sci., Proc. 4: 104. ¢.
Diadasia skinneri Cockerell, 1909. Ent. News 20: 206. ¢.

Taxonomy: Michener, 1947. Ann. and Mag. Nat. Hist. (10) 19: 403. 9.

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 215.
ochracea (Cockerell). Wash., Calif., Ariz., Colo., N. Mex., Tex.; Mexico (Zacatecas). Parasite:

Protepeolus singularis Linsley and Michener, Rhipiphorus sexdens Linsley and
MacSwain, Sparnoplius brevicornis Loew. Pollen: Collects pollen from flowers of
Sphaeralcea and Kallstroemia grandiflora. Diadasia olivacea (Cresson, 1878. Acad.
Nat. Sci., Phila., Proc. 30: 216), described from Mexico, may be a senior synonym
according to Timberlake (1957. Jn Linsley and MacSwain, Wasmann Jour. Biol. 15:215);
however, Krombein (1967. U. S. Dept. Agr., Agr. Monog. 2, suppl. 2: 502) deletes the
species from the fauna of America north of Mexico. The type of D. olivacea was
collected by Sumichrast who made his collections of Hymenoptera in the environs of
Orizaba and Cordoba (see Cresson, 1868. Amer. Ent. Soc., Trans. 2: 1). This is well to
the south of the known southernmost occurrence (Zacatecas) of D. ochracea. The types
of D. olivacea and D. ochracea, both males, must be reexamined in order to determine
the validity of the proposed synonymy.

Dasiapis ochracea Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 450. d.

Diadasia blaisdelli Cockerell, 1924. Pan-Pacific Ent. 1: 54. 9.

Biology: Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 215-217, pl. 1, fig. 4.
—Kickwort, Eickwort and Linsley, 1977. Kans. Ent. Soc., Jour. 50:1-17, 9 figs., 1 table (nest
aggregations, nest architecture, life history, parasites, as olivacea).

opuntiae Cockerell. Coastal south. Calif. Pollen: Collects pollen from flowers of Opuntia.
Diadasia rinconis opuntiae Cockerell, 1901. Canad. Ent. 33: 286. 9.

Superfamily APOIDEA 2123

palmarum Timberlake. South. Calif. desert. Pollen: Collects pollen from flowers of
Sphaeralcea.
Diadasia palmarum Timberlake, 1940. Brooklyn Ent. Soe., Bul. 35: 27. d, °.
piercei Cockerell. Tex., Ariz.
Diadasia piercei Cockerell, 1911. Canad. Ent. 43: 132. d, 2.
rinconis mimetica Cockerell. Calif. (isl. off coast). Pollen: Collects pollen from flowers of
Opuntia. Another subspecies (petrina) is known from Baja California.
Diadasia australis mimetica Cockerell, 1924. Pan-Pacifie Ent. 1: 53. 9.
rinconis rinconis Cockerell. Tex., N. Mex., Ariz., Calif.; Mexico (Baja California). Pollen:
Collects pollen from flowers of Opuntia.
Diadasia rinconis Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 154. 2.
Diadasia rinconi(!) Snow, 1906. Kans. Acad. Sci., Trans. 20: 137.
sphaeralcearum affinis Timberlake. Tex. to south. Calif. and Nev. Pollen: Collects pollen from
flowers of Sphaeralcea.
Diadasia sphaeralcearum affinis Timberlake, 1939. Brooklyn Ent. Soc., Bul. 34: 14. d, °.
sphaeralcearum sphaeralcearum Cockerell. N. Mex., Ariz. Pollen: Collects pollen from flowers
of Sphaeralcea.
Diadasia sphaeralcearum Cockerell, 1905. Amer. Nat. 39: 744. d.

Taxonomy: Michener, 1937. Ann. and Mag. Nat. Hist. (10) 19: 403. °.
tuberculifrons Timberlake. Ariz., Calif., Nev. Pollen: Collects pollen from flowers of
Sphaeralcea.
Diadasia tuberculifrons Timberlake, 1939. Brooklyn Ent. Soc., Bul. 34: 13. d, °.
vallicola Timberlake. Ariz., Calif.; Mexico (Baja California). Parasite: Rhipiphorus diadasiae
Linsley and MacSwain, Sphaeropthalma blakei Fox, Villa sp. near flavipilosa Cole.
Pollen: Collects pollen from flowers of Sphaeralcea.
Diadasia vallicola Timberlake, 1940. Brooklyn Ent. Soc., Bul. 35: 24. ¢.

Biology: Hall and Hurd In Linsley and MacSwain, 1957. Wasmann Jour. Biol. 15: 217-218
(nest, parasites).
vestita Timberlake. N. Mex. (Rio Puerco in Bernalillo Co.).
Diadasia vestita Timberlake, 1956. Pan-Pacifie Ent. 32: 91. d.

Genus MELITOMA Lepeletier and Serville

Melitoma Lepeletier and Serville, 1828. Encycl. Meth., Hist. Nat. Ins., v. 10, p. 529.
Type-species: Melitoma euglossoides Lepeletier and Serville. Monotypic.
Entechnia Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 476.
Type-species: Anthophora taurea Say. Monotypic and orig. desig.
Meliphila Schrottky, 1902. Buenos Aires Mus. Nac. de Hist. Nat. An. 7: 310.
Type-species: Apis segmentaria Fabricius. Monotypic and orig. desig. (=Meliphila
ipomoeae Schrottky and Melitoma euglossoides Lepeletier and Serville).

All species visit flowers of Ipomoea for pollen and nectar, but also visit a wide variety of
other flowers for nectar.

grisella (Cockerell and Porter). N. Dak., Nebr., Kans., Colo., N. Mex.
Entechnia grisella Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 409. d.
Entechnia dakotensis Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 409. °.

Biology: Hungerford and Williams, 1912. Ent. News 23: 258. —Cockerell, 1934. Amer. Mus.
Novitates 697: 10. —Linsley, 1960. N. Y. Ent. Soc., Jour. 68: 17-18 (floral relationships).
segmentaria (Fabricius). Tex. to Argentina. Parasite: Anthrax cintalapa Cole,

Monodontomerus mandibularis Gahan, Trophocleptria schraderi Michener?

Apis segmentaria Fabricius, 1804. Systema Piezatorum, p. 371. ?.

Melitoma euglossoides Lepeletier and Serville, 1828. Encyl. Meth., Hist. Nat. Ins. v. 10, p.
529.

Anthophora fulvifrons Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 341.

Melissodes? marginella Cresson, 1872. Amer. Ent. Soc., Trans. 4: 282. d, °.

Meliphilia ipomoeae Schrottky, 1902. Mus. Nac. Buenos Aires, An. 7: 311.

2124 Hymenoptera in America North of Mexico

Taxonomy: Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 131-132, figs. 89-91 (genitalia).
—Moure, 1960. Studia Ent. 3: 115-116 (synonymy, notes on types).

Biology: Linsley, MacSwain and Smith, 1956. South Calif. Acad. Sci. Bul. 55: 94, figs. 6, 11
(larva). —Michener, 1974. Kans. Ent. Soc., Jour. 47: 19-22 (intermingled nests with
Ancyloscelis ?apiformis, reported as ?armatus). —Torchio, 1974: Kans. Ent. Soc., Jour.
47: 54-63, 2 tables (nest association with Ancyloscelis ?armatus).

taurea (Say). N. J. to Fla., west to Ill, Kans. and Mo. Parasite: Monodontomerus mandibularis
Gahan, Triepeolus donatus (Sm.).
Anthophora taurea Say, 1837. Boston Jour. Nat. Hist. 1: 410. 6, 2.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 242-244, figs. 7-8
(redescript.).

Biology: Walsh, 1868. Amer. Ent. 1: 10. —Ashmead, 1894. Psyche 7: 25. —Robertson, 1914.
Ent. News 25: 68. —Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 175. —Rau, 1929. Psyche 36:
155. —Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 222. —Michener, 1975. Kans. Ent. Soc.,
Jour. 48: 194-200, 3 figs. (nesting site, supersedure by Paranthidium jugatorium).

TRIBE EUCERINI

This is a very large tribe of pollen-collecting bees which is found on all the continents except
Australia. It is especially diverse in the New World, particularly in the Neotropical Region. In
America north of Mexico, the Eucerini are represented by 15 of the 18 genera known to occur on
the North American continent. Most of the species are solitary, but some live in colonies and all
make their nests in the ground.

Taxonomy: Moure and Michener, 1955. Dusenia 6: 239-331, 35 figs. (generic classification of
Neotropical genera). —LaBerge, 1957. Amer. Mus. Novitates 1837: 1-44, 42 figs. (generic
classification of North and Central American genera). —LaBerge, 1962. Parana Univ., Bol.,
Zool. 11: 1-12 (types of Eucerine bees in Brit. Mus. Nat. Hist.). —Rozen, 1965. Amer. Mus.
Novitates 2233: 3-13, figs. 1-34 (larvae).

Genus SYNHALONIA Patton

Synhalonia Patton, 1879. U. S. Geol. Geog. Survey, Bul. 5: 473.

Type-species: Melissodes fulvitarsis Cresson. Orig. desig.
Eusynhalonia Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 63.

Type-species: Melissodes edwardsiti Cresson. Monotypic and orig. desig.
Synalonia Robertson, 1905. Amer. Ent. Soc., Trans. 31: 365. Emend.

The bees of this North American genus fly primarily during the spring months and are ob-
served only exceptionally during the summer. A second generation may occur in some species
since specimens have been collected as late as August. Insofar as known, the females do not col-
lect pollen from the flowers of Compositae, but visit the flowers of a wide range of plant fami-
lies, including especially the Ranunculaceae, Leguminosae, Hydrophyllaceae, Boraginaceae, and
Saxifragaceae. At least some of the species are apparently oligolectic.

Revision: Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 1-76, 80 figs. (spp. of western U. S.).

Taxonomy: Robertson, 1905. Amer. Ent. Soc., Trans. 31: 366-367 (Illinois species).
—Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 74-97. —Cockerell, 1914. Ann. and Mag. Nat.
Hist. (8) 8: 284-286 (partial key). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
312-323, figs. 87-89 (eastern U. S. spp.).

Biology: Rust and Clement, 1977. Kans. Ent. Soc., Jour. 50:41-43, table 3 (spp. visiting
flowers of Collinsia sparsiflora).
acerba (Cresson). Calif., Oreg., Nev., Utah, Mont. Pollen: Apparently an oligolege of
Arctostaphylos, but also visits flowers of Ribes awreum, Taraxacum vulgare.
Melissodes acerba Cresson, 1879. Amer. Ent. Soc., Trans. 7: 210. 9.
Melissodes nevadensis Cresson, 1879 (not 1874). Amer. Ent. Soc., Trans. 7: 209. ¢. Preoce.
Melissodes intrudens Cresson, 1879. Amer. Ent. Soc., Trans. 7: 225. N. name.

Taxonomy: Fowler, 1899. Canad. Ent. 10: 138. ¢.

Superfamily APOIDEA 2125

actuosa (Cresson). Wash. and Idaho, south to Calif., Utah, Ariz., and N. Mex. Pollen: Polylege,

visits a wide variety of flowers including Agoseris heterophylla, Amsinckia
douglasiana, A. intermedia, Antirrhinum, Aplopappus, Astragalus, Brassica
campestris, Brodiaea laxa, Calochortus, Chaenactis, Cirsium, Collinsia, Coreopsis
californica, Cryptantha intermedia, Dodecatheon, Eriodictyon californicum, Erodium
cicutarium, Eschscholzia californica, Haplopappus linearifolius, Helianthus nuttallii,
Hesperochiron californica, Lasthenia chrysostoma, L. gracilis, Layia glandulosa, L.
platyglossa, Lesquerella gordonii, Linanthus, Lotus, Lupinus bicolor, L. densiflorus, L.
nanus, L. succulentus, Medicago hispida, Melilotus, Oenothera pallida, Oreocarya,
Orthocarpus purpurascens, Phacelia ciliata, P. distans, P. humilis, Plagiobothrys
californicus, Prunus, Ranunculus californicus, Raphanus sativus, Rhamnus,
Rhododendron, Salix, Salvia columbariae, Sambucus, Sisyrinchium bellum, Tamarix
gallica, Trifolium repens, T. tridentatum, Wyethia. Diadasia tricincta Provancher,
listed in this catalog as possibly a synonym of Diadasia enavata (Cresson), may be a
synonym of this species (see Timberlake, 1969. Calif. Univ. Pubs. Ent. 57:13).

Melissodes actuosa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 208. °.

Synhalonia californica Fowler, 1899. Canad. Ent. 31: 137. 2. Preoce.

Synhalonia fowleri Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 28. °.

Synhalonia territella Cockerell, 1909. Entomologist 38: 146. ¢.

albescens Timberlake. Calif., Ariz., Nev. Pollen: Polylege, visits flowers of Amsinckia, Aster

abatus, Dalea saundersti, Delphinium parishii, Larrea tridentata, Lycium torreyi,
Oenothera clavaeformis, Prosopis, Salvia carduacea, Sphaeralcea ambigua, S. orcuttii,
Tamarix.

Synhalonia albescens Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 14, figs. 5, 6. 2, ¢.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 42 (floral information).
amsinckiae Timberlake. Wash. (Pullman and Wawawai), Calif.; Mexico (Baja California).
Pollen: Apparently an oligolege of Amsinckia including A. douglasiana, A. intermedia,
A. eastwoodae, A. tessellata, but also visits flowers of Arctostaphylos glauca, Encelia
californica, Lantana sellowiana, Phacelia, Raphanus sativus, Salvia columbariae, S.
mellifera.

Synhalonia amsinckiae Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 15, figs. 7, 8. 2, ¢.
angustifrons Timberlake. Oreg., Calif., Ariz., Utah; Mexico (Baja California). Pollen: Polylege,
visits a wide variety of flowers including Amsinckia intermedia, Arctostaphylos

mariposa, Astragalus pachypus, Brassica incana, Cercidium, Cryptantha,
Cynoglossum, Dudleya, Encelia californica, Eriogonum, Lantana sellowiana, Larrea
tridentata, Lotus scoparius, Lupinus excubitus, L. formosus, L. paynei, Parkinsonia
aculeata, Penstemon antirrhinoides, P. palmeri, P. spectabilis, Phacelia distans,
Prunus fasciculata, Rhus trilobata, Salvia apiana, S. columbariae, S. dorrii, S.
mellifera, Sambucus, Scrophularia californica.
Synhalonia angustifrons Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 16, figs. 9, 10. 2, 6.
aragalli Cockerell. Colo., Tex. Pollen: Unknown, but visits flowers of Astragalus, Oxytropis
lambertii.
Synhalonia frater aragalli Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 25. 9.
argyrophila (Cockerell). Tex. Pollen: Unknown, but visits flowers of Astragalus, Dalea
formosa.
Tetralonia argyrophila Cockerell, 1909. Entomologist 42: 147. ¢.
atriventris (Smith). Minn. to Mass., south to Ga. Pollen: Unknown, but visits flowers of
Aesculus, Astragalus, Lathyrus, Lupinus, Mertensia, Penstemon hirsutus, Robinia,
Vicia.
Melissodes atriventris Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 310. d.
Synhalonia atriventris fuscipes Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 54. °.

Preoce.
Tetralonia robertsoni Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 13: 288. 2. N. name.

Taxonomy: Robertson, 1897. Acad. Sci. St. Louis, Trans. 7: 353. d, °.
Biology: Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 211 (nesting habits).

2126 Hymenoptera in America North of Mexico

bakeri Timberlake. N. Mex. (Aztec).
Synhalonia bakeri Timberlake, 1973. Ent. Soc. Wash., Proc. 75: 317. d.
belfragii (Cresson). Mich., Ill., Ind., Tex. Pollen: Unknown, but visits flowers of Aesculus,
Arabis, Camassia, Cercis canadensis, Claytonia, Collinsia, Dentaria, Dicentra,
Erythronium, Geranium, Hydrophyllum, Isopyrum, Lithospermum, Mertensia,
Penstemon ambiguus, Phacelia, Polemonium, Ranunculus, Viola.
Melissodes belfragii Cresson, 1872. Amer. Ent. Soc., Trans. 4: 278. °.
Melissodes honesta Cresson, 1872. Amer. Ent. Soe., Trans. 4: 279. 6.
birkmanniella (Cockerell). Tex.
Tetralonia birkmanniella Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 93. 3.
californica californica (Cresson). Calif. Pollen: Unknown, but visits flowers of Raphanus
sativus.
Melissodes Californica Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 196. 3.
californica deserticola Timberlake. South Calif., desert. Pollen: Apparently polylectic, visits
flowers of Amsinckia, Astragalus, Coreopsis californica, Gilia davyi, G. latifolia,
Isomeris, Oenothera, Phacelia, Salvia carduacea, Sphaeralcea, Tamarix gallica.
Synhalonia californica deserticola Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 24. 9, ¢.
carolinensis (Dalla Torre). N. C.
Tetralonia atrifrons Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 308. 2. Preoce.
Eucera carolinensis Dalla Torre, 1896. Cat. Hym., v. 10, p. 228. 2. N. name.

Taxonomy: LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 950.
cercidis Timberlake. Okla. (Alabaster Caverns State Park). Pollen: Unknown, but visits
flowers of Cercis canadensis.
Synhalonia cercidis Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 24, figs. 21, 22. 6.
chrysobotryae (Cockerell). Colo. to Okla., south to Tex. Pollen: Unknown, but visits flowers of
Astragalus including A. lindheimeri, Cercis canadensis, Ribes longifolium.
Tetralonia chrysobotryae Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 332. 9, 6.
chrysophila (Cockerell). N. Mex., Colo., ?7Idaho. Pollen: Unknown, but visits flowers of Ribes
aureum.
Tetralonia chrysophila Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 13: 384. 9.
conformis Timberlake. Okla. (Waurika), Tex. (Midland). Pollen: Unknown, but visits flowers of
Aster tanacetifolius, Astragalus lindheimeri.
Synhalonia conformis Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 27, figs. 25, 26. d, 2.
cordleyi Viereck. B. C. to Calif., Idaho, Colo., Utah. Pollen: Apparently polylectic, visits
flowers of Agoseris, Amsinckia, Asclepias, Astragalus, Balsamorrhiza, Brodiaea,
Camissonia, Cirsium, Convolvulus, Cryptantha, Encelia, Erigeron, Eriogonum,
Erysimum, Lupinus, Marrubium, Melilotus, Mesembryanthemum, Mirabilis,
Oenothera, Phacelia, Raphanus, Salvia, Sidalcea, Sphaeralcea, Statice, Taraxacum,
Thelypodium, Trifolium, Vicia.
Synhalonia Cordleyi Viereck, 1905. Canad. Ent. 16: 316. 2, ¢.
Tetralonia poetica Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 13: 424. d.
Tetralonia cordleyi postica(!) Cockerell, 1935. Pan-Pacifie Ent. 11: 53.
Tetralonia cordleyi orophila Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 13: 284. 2.
crenulaticornis (Cockerell). Colo., N. Mex.
Melissodes crenulaticornis Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 454. 6.
Melissodes crenulaticornis form maculata Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2:
455. 6.
Taxonomy: Cockerell, 1899. Entomologist 32: 157. °.
cressoniana Cockerell. Tex., Kans., Nebr., Ariz., Utah. Pollen: Unknown, but visits flowers of

Larrea tridentata.
Synhalonia cressoniana Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 177. 9.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 42, tables 14, 15 (floral
relationships).

Superfamily APOIDEA 2127

delphinii Timberlake. Wash. to Calif., Utah. Pollen: Unknown, but visits flowers of Bloomeria,
Brodiaea, Delphinium, Platystemon, Trifolium.
Synhalonia delphinii Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 30, figs. 29, 30. 2, d.
dorsata Timberlake. Calif. Pollen: Apparently polylectic, visits flowers of Amsinckia
intermedia, Brodiaea crocea, Collinsia bicolor, Convolvulus occidentalis, Coreopsis
lanceolata, Eriodictyon californicum, E. crassifoliwm, Eschscholzia californica,
Grindelia, Lathyrus, Lotus scoparius, Lupinus bicolor, L. formosus, L. nanus,
Penstemon antirrhinoides, Salvia columbariae, S. mellifera, Trichostema parishii.
Synhalonia dorsata Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 32. 6, 2.
douglasiana (Cockerell). Wash. (Steamboat Rock).
Tetralonia Douglasiana Cockerell, 1906. Canad. Ent. 38: 278. °.

dubitata (Cresson). Minn. to Pa., south to S. C. and Tex. Pollen: Unknown, but visits flowers of
Aesculus, Astragalus, Dicentra, Melilotus, Mertensia, Robinia, Salvia.
Melissodes dubitata Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 194. 2, 6.

edwardsii (Cresson). B. C. to Calif., Idaho, Mont., Utah, Nev., Colo., Wyo., N. Mex.; Mexico

(Baja California). Pollen: Polylectic, visits a wide variety of flowers including Allium,
Amsinckia, Astragalus, Abronia, Arnica, Balsamorrhiza, Brassica, Brodiaea, Besseya,
Cirsium, Collinsia sparsiflora, Convolvulus, Conium, Dalea, Daucus, Erysimum,
Eschscholzia, Encelia, Erigeron, Helianthus, Hydrophyllum, Ligustrum, Lithosperum,
Lotus, Lupinus, Malvastrum, Medicago, Melilotus, Mentha, Mesembryanthemum,
Mirabilis, Nemophila, Oenothera, Penstemon, Physocarpus, Phacelia, Phoradendorn,
Potentilla, Raphanus Rhamnus, Rubus, Sidalcea, Sphaeralcea, Stachys, Taraxacum,
Trifolium, Thelypodium, Vicia, Viola, Zygadenus.

Melissodes edwardsti Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 195. 3.

Synhalonia edwardsii race angustior Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 347.
3 (2 misdet.).

Tetralonia edwardsti vagabunda Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 95. d.

Biology: Rust and Clement, 1977. Kans. Ent. Soc., Jour. 50:42, table 3 (floral visitation).
frater albopilosa Fowler. Oreg., Calif. Pollen: Apparently polylectic, visits a wide variety of
flowers including Amsinckia, Astragalus, Brodiaea, Camissonia tanacetifolia,

Ceanothus, Chamaebatia, Cirsium, Convolvulus, Lathyrus, Layia, Lotus, Lupinus,
Medicago, Penstemon, Ranunculus, Stachys, Thermopsis, Trifolium, Vicia.
Synhalonia albopilosa Fowler, 1899. Canad. Ent. 10: 138. 6.

Taxonomy: Cockerell, 1924. Pan-Pacific Ent. 1: 55.
frater frater (Cresson). Colo., Wyo., Utah, Idaho, Oreg., Nev., N. Mex. (Barela Mesa). Pollen:
Apparently polylectic, visits flowers of Astragalus, Hedysarum, Lithospermum,
Lupinus, Medicago, Mertensia, Ribes, Vicia. An unnamed variety of this occurs in
Utah, Colorado and Washington in which the metasoma of the female is entirely black
beyond tergum I.
Melissodes frater Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 197. 6.
Tetralonia lata lautipes Cockerell, 1924. Amer. Mus. Novitates 113: 2. 9.
frater lata (Provancher). B. C., Wash., Oreg. Pollen: Unknown, but visits flowers of
Hydrophyllum.
Melissodes lata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 302. 2.
Melissodes nigricornis Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 302. 3.
Synhalonia edwardsii race latior Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 347. 6.
fulvitarsis annae (Cockerell). Colo., Wyo., Utah, N. Mex.
Tetralonia annae Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 114. 2.
Tetralonia annae patruelis Cockerell, 1924. Amer. Mus. Novitates 113: 2. 9.
Tetralonia rotgeri Cockerell, 1937. Ent. News 48: 256. °.
fulvitarsis fulvitarsis (Cresson). Alta. and B. C. to Calif., Nev., Idaho, Colo., Wyo., Kans.
(Lawrence). Pollen: Unknown, but visits flowers of Astragalus, Cryptantha, Phlox,
Taraxacum, Thelypodium.
Melissodes fulvitarsis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 196. ¢.
Tetralonia Yakimensis Cockerell, 1906. Canad. Ent. 38: 278. ¢.
Tetralonia medicata Cockerell, 1911. Canad. Ent. 48: 34. 2.

2128 Hymenoptera in America North of Mexico

Taxonomy: Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 474. 9.

fulvohirta (Cresson). N. C. to Ga. Pollen: Unknown, but visits flowers of Elaeagnus, Vicia.
Melissodes fulvohirta Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 213. ¢.

fuscotincta Cockerell. Ariz., Colo.
Synhalonia fuscotincta Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 178. d.
hamata (Bradley). Eastern U.S., west to S. Dak., Colo., Kans., Nebr. and Mo. Pollen:
Apparently polylectic, visits a wide variety of flowers including Aesculus, Asclepias,
Astragalus, Baptisia, Blephilia, Brassica, Camassia, Capsella, Cardamine, Cercis,
Claytonia, Collinsia, Commandra, Convolvulus, Cornus, Crataegus, Delphinium,
Dianthera, Dodacatheon, Ellisia, Erigeron, Fragaria, Geranium, Gleditsia,
Gymnocladus, Heracleum, Hydrophyllum, Iris, Krigia, Lithospermum, Lythrum,
Lobelia, Medicago, Melilotus, Mertensia, Monarda, Nepeta, Oenothera, Oxalis,
Penstemon, Petalostemon, Phlox, Podophyllum, Polemonium, Prunus, Ptelea, Pyrus,
Ranunculus, Ribes, Robinia, Rosa, Rubus, Ruellia, Salix, Salvia, Scutellaria, Senecio,
Stellaria, Tradescantia, Trifolium, Verbena, Verbesina, Viburnum, Vicia, Viola, Vitis,
Zizia.
Tetralonia hamata Bradley, 1942. Ent. News 53: 189. 6, 9.
hirsutissima (Cockerell). B. C.
Tetralonia hirsutissima Cockerell, 1916. Ann. and Mag. Nat. Hist. (8) 17: 428. @.
hurdi Timberlake. B. C. to Calif. Pollen: Apparently polylectic, collects pollen from flowers of
Collinsia sparsiflora, but also visits flowers of Allium, Brodiaea, Cryptantha, Lupinus,
Phacelia, Taraxacum.
Synhalonia hurdi Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 42, figs. 43, 44. d, 2.

Biology: Rust and Clement, 1977. Kans. Ent. Soce., Jour. 50:41, 43, table 3 (nest, cell
provisions, floral relationships).
illinoensis Robertson. IIl., Mo., Okla. Pollen: Unknown, but visits flowers of Astragalus
lindheimeri, Lithospermum canescens.
Synhalonia illinoensis Robertson, 1902. Canad. Ent. 34: 49. 3.
lepida (Cresson). Colo., to Okla. and Tex. Pollen: Unknown, but visits flowers of Actinia,
Astragalus, Cercis, Gaillardia, Lesquerella, Marrubium, Sphaeralcea.
Melissodes lepida Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 198. 6.
lunata Timberlake. Oreg., Calif. Utah, Ariz.; Mexico (Baja California). Pollen: Apparently
polylectic, visits flowers of Amsinckia intermedia, Arbutus menziesii, Arctostaphylos
glauca, A. mariposa, A. pungens, Brassica, Calendula, Collinsia sparsiflora,
Cryptantha, Isomeris arborea, Encelia californica, Lasthenia chrysostoma, Lupinus
albifrons, L. densiflorus, L. latifolius, L. micranthus, L. nanus, L. succulentus,
Nemophila menziesii, Phacelia, Phlox, Ranunculus occidentalis, Ribes indecorum,
Salix, Salvia sonomensis, Sidalcea malvaeflora, Solanum umbelliferum, Trichostema
parishu, Trifolium tridentatum, Wyethia angustifolia.
Synhalonia lunata Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 46. d, °.

Biology: Rust and Clement, 1977. Kans. Ent. Soc., Jour. 50:42, table 3 (floral visitation).
lutziana (Cockerell). Colo. Pollen: Unknown, but a male visited flowers of Sphaeralcea
concinna.
Tetralonia lutziana Cockerell, 1933. Amer. Mus. Novitates 595: 2. ¢.
lycii Cockerell. N. Mex. Pollen: Unknown, but visits flowers of Lycium, plum, lilac.
Synhalonia lycti Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 348. 2.
mohavensis Timberlake. Calif., Nev., Ariz.; Mexico (Baja California). Pollen: Unknown, but
visits flowers of Sphaeralcea ambigua, S. emoryi, S. orcuttii.
Synhalonia mohavensis Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 50, figs. 55, 56. 3.
monozona Timberlake. Oreg. (Klamath Co.), Calif. (Sierra Nevada Mts.).
Synhalonia monozona Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 51. 9.
pagosana (Cockerell). Colo. (Pagoso Springs).
Tetralonia pagosana Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 228. °.
pallidihirta Timberlake. Tex., Mont. Pollen: Unknown, but visits flowers of Dalea formosa.
Synhalonia pallidihirta Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 53. °@.

Superfamily APOIDEA —=_2129

phaceliae (Cockerell). N. Mex., Utah., Ariz. Pollen: Unknown, but visits flowers of Descurainia
sophia, Lesquerella gordonii, Phacelia corrungata, Sphaeralcea.
Tetralonia phaceliae Cockerell, 1911. Amer. Ent. Soc., Trans. 37: 238. 2.
primaveris Timberlake. Calif. (Colorado and Mojave Deserts), Nev., Utah, Ariz. (Mohave Co.).
Pollen: Apparently polylectic; visits flowers of Amsinckia, Astragalus, Aster, Brassica,
Delphinium, Encelia, Erysimum, Gilia, Hyptis, Isomeris, Larrea, Lotus, Lupinus,
Oenothera, Phacelia, Prunus, Salvia, Sphaeralcea, Tamarix, Thelypodium.
Synhalonia primaveris Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 55, figs. 59, 60. d, °.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 43 (floral relationships).
quadricincta Timberlake. Calif., Nev., Utah. Pollen: Unknown, but visits flowers of
Astragalus, Berberis fremontii, Dalea, Melilotus officinalis, Oenothera clavaeformis var.

auriantiaca, Sphaeralcea ambigua, Stanleya, Tamarix, Thelypodium laciniatum.
Synhalonia quadricincta Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 57, figs. 61, 62. °,
é.

rosae Robertson. Va. to Ohio, Mo. and Kans., south to Fla. and Tex. Pollen: Apparently
polylectic; visits flowers of Asclepias, Batodendron, Blephilia, Cornus, Croton,
Dianthera, Diospyros, Elaeagnus, Geranium, Houstonia, Hydrophyllum, Iris, Linaria,
Melilotus, Monarda, Nemophila, Oenothera, Penstemon, Pontederia, Rosa, Rubus,
Stachys, Trifolium, Verbena, Xanthisma.
Synhalonia rosae Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 54. 2.
Tetralonia fedoris Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 93. d.
speciosa (Cresson). S. Dak. to Texas, west to Utah and Idaho. Pollen: Collects pollen from
flowers of Oenothera pallida, but also visits flowers of Balsamorrhiza, Brazoria,
Caragana, Gaillardia, Monarda, Penstemon.
Melissodes speciosa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 198. °.
Melissodes dilecta Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 199. 3.
Synhalonia gillettei Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 203. 6.
Synhalonia gillettei snoviana Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 179. 6.
Synhalonia astragalina Cockerell, 1905. Ent. News 16: 271. 6.
Tetralonia astragalina clarissima Cockerell, 1933. Ann. and Mag. Nat. Hist. (10) 11: 372.
é.

Taxonomy: Cockerell, 1905. Entomologist 38: 148. ¢. —Bradley, 1942. Ent. News 53: 190.

Biology: Folsom, 1922. Ent. Soc. Amer., Ann. 15: 182 (nesting habits). —Linsley, MacSwain
and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 45-46 (floral relationships).
stretchii (Cresson). Oreg., Calif. Pollen: Unknown, but visits flowers of Cirsium, Coreopsis,
Eriodictyon, Fremontodendron, Lupinus, Vicia.
Melissodes stretchii Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 207. °.
Synhalonia idiotes Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 105. 2.
suavis (Cresson). Colo.
Melissodes suavis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 210. 2.
texana Timberlake. Tex. (Dallas). Pollen: Unknown, but visits flowers of Cercis canadensis.
Synhalonia texana Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 62, figs. 69, 70. ¢.
tricinctella Timberlake. Calif., Utah. Pollen: Apparently polylectic; visits flowers of
Amsinckia intermedia, Aster abatus, Astragalus tener, Chaenactis fremontii, Coreopsis
californica, Cryptantha intermedia, Dalea saundersii, Encelia farinosa, Erysimum,
Gilia multicaulis, Haplopappus cooperi, H. linearifolius, Lotus scoparius, L. strigosus,
Malacothrix glabrata, Plagiobothrys californicus, Rhus trilobata, Ribes indecorum,
Salvia columbariae, Sambucus, Sphaeralcea, Tamarix gallica.
Synhalonia,tricinctella Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 63, figs. 71, 72. d, °.
truttae Cockerell. N. Mex., Ariz. Pollen: Unknown, but visits flowers of Iris missouriensis.
Synhalonia truttae Cockerell, 1905. Entomologist 38: 147. d, °.
venusta carinata (Timberlake). Calif.; Mexico (Baja California). Pollen: Apparently strictly

oligolectic on Clarkia unguiculata.
Tetralonia venusta carinata Timberlake, 1961. Pan-Pacific Ent. 37: 212. d.

2130 Hymenoptera in America North of Mexico

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 31-34, 45, fig. 8 (floral
relationships).
venusta venusta (Timberlake). Calif., Ariz., Nev.; Mexico (Baja California). Pollen: Apparently
oligolectic on Camissonia clavaeformis, but visits other flowers evidently for nectar
including Larrea tridentata, Sphaeralcea orcuttii.
Tetralonia venusta Timberlake, 1961. Pan-Pacific Ent. 37: 209. @, 6.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 20, 46 (behavior,
floral relationships). —Linsley, MacSwain and Raven, 1964. Calif. Univ. Pubs. Ent. 33: 77
(behavior, floral relationships). —MacSwain, Raven and Thorp, 1978. Calif. Univ. Pubs.
Ent. 70: 30, fig. 8 (floral relationships). —Hurd and Linsley, 1975. Smithsn. Contrib. Zool.
193: 43 (floral relationships).

virgata Cockerell. Oreg., Calif. Pollen: Unknown, but visits flowers of Astragalus pomonensis,
Brodiaea capitata, Cirsium, Phacelia, Pogogyne parviflora, Rhododendron, Trichostema
lanata.
Synhalonia belfragei virgata Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 100. 9.
zonata Timberlake. Oreg., Calif. Pollen: Unknown, but visits flowers of Amsinckia intermedia,
Cryptantha, Lupinus nanus, Nemophila menziesii, Plagiobothrys.
Synhalonia zonata Timberlake, 1969. Calif. Univ. Pubs. Ent. 57: 68, figs. 79, 80. 2, é.

Genus SYNTRICHALONIA LaBerge

Syntrichalonia LaBerge, 1957. Amer. Mus. Novitates 1837: 10.
Type-species: Melissodes exquisita Cresson. Monotypic and orig. desig.
exquisita (Cresson). Tex., N. Mex., Ariz.; Mexico (D. F., Durango, Jalisco and Zacatecas).
Pollen: Oligolege, principally composite tribes Helenieae and Heliantheae including
Encelia, Helenium hoopesii, Helianthus annuus, Heliopsis parviflora, Verbesina
encelioides, V. oreophila, Viguiera dentata, but visits other flowers for nectar.
Melissodes exquisita Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 213. @.
Melissodes herricki Cockerell, 1905. Psyche 12: 98. 6, °.

Biology: Zavortink, 1975. Pan-Pacifie Ent. 51: 240-242, table 3 (host plants, behavior,
distribution).

Genus EUCERA Scopoli

Eucera Scopoli, 1770. Annus Hist. Nat., v. 4, p. 8.
Type-species: Apis longicornis Linnaeus. Desig. by Latreille, 1810.
maculata Lepeletier. America. No doubt exotic or incorrectly assigned to Eucera.
Eucera maculata Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 129. @.

Genus XENOGLOSSA Smith

The bees of this North and Central American genus are dependent solely upon the pollen and
to a large extent upon the nectar of both cultivated and uncultivated Cucurbita (squashes,
gourds and pumpkins). Like the species of the genus Peponapis, the males commonly spend
most of the day and a good part of the night in the wilted and closed flowers of these plants. At
the beginning of the season and before nesting activities are commenced it is also not unusual to
encounter females in the wilted and closed flowers. The pollen collecting devices of the species
are species-specific and apparently this has influenced the ability of the different species to col-
lect and utilize pollens of various Cucurbita, both wild and domestic. These bees are excep-
tionally valuable pollinators of the squashes, gourds and pumpkins.

Revision: Hurd and Linsley, 1964. Hilgardia 35: 384-425, figs. 1-11 (U. S. spp.). —Hurd and
Linsley, 1967. Ent. Soc. Amer., Ann. 60: 988-1007, 19 figs., 2 tables, 5 maps (North and
Centr. Amer. spp.). —Hurd and Linsley, 1970. Calif. Univ. Pubs. Ent. 62: 1-39, 11 figs. 4
tables, 3 maps (classification).

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 246-249, figs. 73-74 (eastern
U.S. spp.). —Rozen, 1965. Amer. Mus. Novitates 2233: 6-11, figs. 1-19 (larva).

Superfamily APOIDEA 213]

Biology: Michelbacher, Smith and Hurd, 1964. Calif. Agr. 18: 2-4, figs. (pollination of
squashes, gourds and pumpkins). —Hurd and Linsley, 1964. Hilgardia 35: 376-382, 2 tables
(biol. summary). —Michelbacher, Hurd and Linsley, 1968. Bee World 49: 159-167, 6 figs.
(feasibility of introducing squash bees into the Old World). —Hurd, Linsley and Whitaker,
1971. Evolution 25: 218-234, 4 figs., 3 tables (squash bees and origin of cultivated
Cucurbita).

Genus XENOGLOSSA Subgenus EOXENOGLOSSA Hurd and Linsley

Xenoglossa subg. Eoxenoglossa Hurd and Linsley, 1970. Calif. Univ. Pubs. Ent. 62: 34.
Type-species: Melissodes strenuwa Cresson. Orig. desig.
kansensis Cockerell. N. C. and Ga., west to Colo., N. Mex. and Tex. Pollen: Oligolectie on
Cucurbita foetidissima and domestic species of Cucurbita.
Xenoglossa strenua var. Kansensis Cockerell, 1905. Psyche 12: 104. 6.
strenua (Cresson). Transcont., Md. to Fla., west to south. Calif., south to San Luis Potosi,
Durango, Baja California and Sinaloa, Mexico. Parasite: Triepeolus remigatus (Fabr.).
Pollen: Oligolectic on Cucurbita foetidissima and domestic species of Cucurbita. Pollen
collecting females have been found occasionally at the flowers of Cucurbita digitata and
C. palmata, but these plants do not appear to be preferred pollen sources.
Melissodes strenua Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 213. 2, 6.
Xenoglossa cucurbitarum Cockerell, 1896. Canad. Ent. 28: 192. d, °.

Taxonomy: Bohart, 1964. Pan-Pacifie Ent. 40: 174-182, 17 figs. (larva). —Rozen, 1965. Amer.
Mus. Novitates 2233: 10-11, figs. 15-19 (larva).

Biology: Bohart, 1964. Pan-Pacific Ent. 40: 174-182, 17 figs. (nesting habits, nest architecture,
parasite). —Bohart, 1966. Pan-Pacific Ent. 42: 255-262, 20 figs. (parasite).

Genus XENOGLOSSA Subgenus XENOGLOSSA Smith

Xenoglossa Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 315.
Type-species: Xenoglossa fulva Smith. Monotypic.

SPECIES GROUP PATRICIA

angustior Cockerell. Calif., Ariz., N. Mex.; Mexico (Baja California and Sonora). Pollen:
Oligolectic on Cucurbita digitata, C. foetidissima, C. palmata and most, if not all,
domestic Cucurbita grown within its geographic range.
Xenoglossa patricia angustior Cockerell, 1899. Entomologist 33: 64. ¢.

Taxonomy: Rozen, 1965. Amer. Mus. Novitates 2233: 8-10, figs. 3-10 (larva).
patricia Cockerell. N. Mex., Tex.; Mexico (Chihuahua, Coahuila, and Durango). Pollen:
Oligolectic on Cucurbita foetidissima and most, if not all, domestic Cucurbita grown
within its geographic range.
Xenoglossa patricia Cockerell, 1896. Canad. Ent. 28: 191. d, 2.
SPECIES GROUP FULVA

gabbii crawfordi Cockerell. South. Ariz.; Mexico (south to Michoacan and Morelos). Pollen:
Oligolectic on Cucurbita martinezti, C. sororia and most, if not all, domestic Cucurbita
grown within its geographic range. The typical subspecies occurs in southern Mexico
(Oaxaca and Chiapas) and Central America (Costa Rica, E] Salvador, Guatemala,
Nicaragua, and possibly Panama).
Xenoglossa crawfordi Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 367. d.
Xenoglossa dugesi Cockerell, 1919. U. S. Natl. Mus., Proc. 55: 194. °.

Biology: Michelbacher and Hurd, 1968. Pan-Pacific Ent. 44: 58-68, 1 fig., 3 tables (nest site,
late season foraging behavior). —Michelbacher and Hurd, 1968. Folia Ent. Mexicana 18-19:

110-111 (ecology and distribution).
Genus CEMOLOBUS Robertson

Cemolobus Robertson, 1902. Canad. Ent. 34: 324.
Type-species: Xenoglossa ipomoeae Robertson. Monotypic and orig. desig.

2132 Hymenoptera in America North of Mexico

ipomoeae (Robertson). IIll., Pa., N. C., Ga. Pollen: Oligolectic on species of Ipomoea.
Xenoglossa ipomoeae Robertson, 1891. Amer. Ent. Soc., Trans. 18: 65. 2, d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 250-251, figs. 75, 76
(redescription).

Genus ANTHEDONIA Michener

Anthedon Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 53. Preoce.
Type-species: Melissodes compta Cresson. Monotypic.

Anthedonia Michener, 1942. N. Y. Ent. Soc., Jour. 50: 282. N. name.

Abda Sandhouse, 1943. U. S. Natl. Mus., Proc. 92: 521. N. name.

Taxonomy: LaBerge, 1955. Kans. Ent. Soc., Jour. 28: 132-135, 2 figs. (synopsis). —LaBerge,
1957. Amer. Mus. Novitates 1837: 19-22, figs. 19-21 (generic diagnosis).
compta (Cresson). N. J. and Ga. west to Utah and N. Mex.; Mexico (Durango). Pollen:
Oligolectic on Oenothera biennis, but visits other flowers for nectar.
Melissodes compta Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 199. 2, 6.

Taxonomy: LaBerge, 1955. Kans. Ent. Soc. Jour. 28: 182, fig. 1. 6, 2. —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 152: 252-253, fig. 77 (redescription).
Biology: Robertson, 1914. Ent. News 25: 72.
nevadensis (Cresson). Tex. west to Calif.; Mexico (Durango). Pollen: Oligolectic on Oenothera,
but visits other flowers for nectar.
Melissodes nevadensis Cresson, 1874. Amer. Ent. Soc., Trans. 5: 102. 6.
Melissodes californica Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 114. 2, ¢
Preocc.
Eucera smithii Dalla Torre, 1896. Cat. Hym., v. 10, p. 247. 6, 2. N. name.
Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 328. —LaBerge, 1955. Kans. Ent.
Soc., Jour. 28: 134, fig. 2. d, 9.

Genus SVASTRA Holmberg

Revision: LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 926-1027 (as subg. Brachymelissodes and
E pimelissodes of the genus Melissodes).

Taxonomy: Michener, LaBerge and Moure, 1955. Dusenia 6: 219-220. —Moure and Michener,
1955. Dusenia 6: 294-298. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 255-263,
figs. 79-80 (spp. of eastern U. S. ).

Genus SVASTRA Subgenus SVASTRA Holmberg

Svastra Holmberg, 1884. Acad. Nac. Cienc. Cordoba, Actas 5: 127.
Type-species: Svastra bombilans Holmberg. Orig. desig.

The typical subgenus does not occur in North America.
Genus SVASTRA Subgenus BRACHYMELISSODES LaBerge

Melissodes subg. Brachymelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 926.
Type-species: Hucera cressonii Dalla Torre. Orig. desig.
cressonii (Dalla Torre). Iowa, Nebr. and east. Colo. south to Tex.; Mexico (Durango). Pollen:

Collects pollen from flowers of Euphorbia marginata and possibly Vernonia, but also
visits other flowers including Cardia, Helianthus, Polygonum.

Melissodes brevicornis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 281. ¢. Preoce.

Eucera cressonii Dalla Torre, 1896. Cat. Hym., v. 10, p. 229. N. name.

Melissodes petulciformis Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 364. °.

Taxonomy: Crawford, 1903. Canad. Ent. 35: 334. 2. —Cockerell, 1906. Ann. and Mag. Nat.
Hist. (7) 17: 367.

Biology: Cockerell, 1915. Ent. News 26: 364 (resting cluster).

Superfamily APOIDEA 2133

Genus SVASTRA Subgenus EPIMELISSODES Ashmead

Epimelissodes Ashmead, 1899. Amer. Ent. Soe., Trans. 26: 63.
Type-species: Melissodes atripes Cresson. Monotypic and orig. desig.

SPECIES GROUP ATRIPES

atripes atrimitra (LaBerge). N. J. south to Fla., Ala., Miss. Pollen: Unknown, but visits
flowers of Afzelia, Bradburya virginiana, Buddleia, Chamaecrista, Gerardia,
Gossypium herbaceum, Hibiscus, Koellia hysopifolia, Kuhnistera, Monarda punctata,
Passiflora incarnata, Primula vulgaris, Rhus.
Melissodes (Epimelissodes) atripes atrimitra LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 949.
yess

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 259 (floral records).
atripes atripes (Cresson). IIl., Mo., and La., west to N. Mex. and Colo. Pollen: Unknown, but
visits flowers of Asclepias incarnata, Aster, Bidens aristosa, Cassia fasciculata,
Chrysopsis, Cleome serrulata, Dalea lanata, Gaillardia, Gonolobus laevis, Gossypiwm
herbaceum, Helenium, Helianthus annuus, Ipomoea pandurata, Kuhnistera, Lythrum
alatum, Melilotus alba, Pycnanthemum virginianum, Thelesperma megapotamicum,
Verbena hastata, V. stricta, Vernonia fasciculata.
Melissodes atripes Cresson, 1872. Amer. Ent. Soc., Trans. 4: 275. 2.

Taxonomy: Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 54. 2, ¢. —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 152: 257, fig. 80 (redescription, floral records).
atripes georgica (Cresson). East coast of Ga. and Fla. Pollen: Unknown, but visits flowers of
Bidens, Rhus.
Melissodes georgica Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 200. d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 259 (tax. characters, floral
records).

SPECIES GROUP OBLIQUA

aegis (LaBerge). Tex. to Fla., north to N. C. Pollen: Unknown, but visits flowers of Aster
pruinosa, Chrysopsis, Helenium tenuifolium, Helianthus annuus, H. radula,
Kuhnistera, Vernonia glauca.
Melissodes (Epimelissodes) aegis LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 959. 6, °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 256-257, fig. 79
(redescription, floral records).
comanche (Cresson). Colo., N. Mex., Tex. Pollen: Unknown, but males visit flowers of
Ximensia californica.
Melissodes comanche Cresson, 1872. Amer. Ent. Soc., Trans. 4: 276. 2, 6.
grandissima (Cockerell). Southeast. Tex. Pollen: Unknown, but the sexes have been collected
at flowers of Ximensia encelioides.
Melissodes grandissima Cockerell, 1905. Canad. Ent. 37: 334. 2.
Melissodes atripes var. acomanche Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 109. d.

Taxonomy: Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 361. ¢. —LaBerge, 1963. Kans.
Ent. Soc., Jour. 36: 56 (geogr. records).
helianthelli (Cockerell). Tex., and Kans. west to South. Calif.; Mexico (Baja California). Pollen:
Unknown, but has been collected at flowers of Gossypium herbaceum, Helianthus
annuus, Verbesina encelioides.
Melissodes helianthelli Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 525. °.
machaerantherae (Cockerell). Tex. (El Paso) to South. Calif.; Mexico (Chihuahua, Coahuila,
Sonora). Pollen: Unknown, but visits flowers of Asclepias subverticillata, Aster,
Baccharis, Cryptantha oblata, Cucurbita palmata, Haplopappus, Heleniwm hoopesii,
Helianthus, Kallstroemia, Machaeranthera, Melilotus alba, Verbesina oreophila,

Wislizenia refracta.
Melissodes machaerantherae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 21. d, °.

Taxonomy: LaBerge, 1963. Kans. Ent. Soc., Jour. 36: 55 (geogr. and floral records).

2134 Hymenoptera in America North of Mexico

obliqua caliginosa (Cresson). N. J. to Ga. Pollen: Unknown, but visits flowers of Borrichia
frutescens, Cirsium, Cucurbita, Helenium tenuifolium, Helianthus atrorubens, H.
microcephalum, H. zonatus, Vernonia glauca, V. noveboracensis.
Melissodes caliginosa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 192. 2, d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 261 (redescript.).

obliqua expurgata (Cockerell). Wash. and Idaho south to Calif., Colo. and Ariz.; Mexico (Baja
California). Pollen: Unknown, but visits flowers of Achillea, Artemisia, Asclepias,
Brassica adpressa, Centaurea, Centromadia pungens, Chrysopsis grandiflora, C.
lanceolata, Coreopsis lanceolata, Engelmannia pinnatidifa, Gaillardia, Geranium
atropurpureum, Gilia capitata, Grindelia camporum, G. nana, Haplopappus
vernonioides, Helianthus annuus, H. bolanderi, H. ciliaris, H. gracilentus, H. petiolaris,
Heliotropium curassavicum, Hemizonia pungens, Lepidospartum squamatum,
Medicago sativa, Melilotus alba, Petalostemon flavescens, Pluchea sericea, Ratibida
columnaris, Scabiosa, Senecio douglas, Solidago, Verbesina encelioides, Wislizenia
refracta.

Melissodes obliqua var. expurgata Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 230. 6.

obliqua obliqua (Say). Southern Ont. and eastern Mont., south to N. Mex. and N. J.; west of
the Appalachians to Fla.; Mexico (Tamaulipas, Coahuila). Parasite: Triepeolus concavus
(Cress.), T. rufithorax Graenicher, Timulla vagans rufinota Mickel? Pollen: Presumably
oligolectic on the Compositae, but visits a wide variety of flowers including Abutilon
theophrasti, Ambrosia, Asclepias incarnatus, A. syriaca, Aster ericoides, Bidens
involucrata, B. aristosa, Blephilia hirsuta, Boltonia asteroides, Carduus crispus,
Cassia, Cephalanthus occidentalis, Chrysopsis, Cirsium discolor, C. lanceolatum, C.
undulatus, Cleome serrulata, Coreopsis tinctoria, C. tripteris, Echinacea pallida,
Erigeron philadelphus, Euphorbia, Gaillardia, Gossypium herbaceum, Grindelia
squarrosa, Helenium altissimum, H. autumnale, Helianthus annuus, H. divaricatus, H.
grosse-serratus, H. laetiflorus, H. maximillianus, H. mollis, H. petiolaris, H.
scaberimus, H. strumosus, H. tuberosus, Heliopsis helianthioides, H. laevis, Ipomoea
pandurata, Kuhnistera purpurea, Lacinaria pycnostachys, Lactuca floridana, Lobelia
leptostachys, L. siphillitica, Lythrum alatum, L. salicaria, Medicago sativa, Melilotus
alba, Monarda fistulosa, Nepeta cataria, Oenothera biennis, Petalostemon candidens, P.
oligophyllum, Prionopsis ciliata, Psoralea tenuiflora, Pycnanthemum flexuosum, P.,
pilosum, Ratibida columnaris, R. pinnata, Rudbeckia hirta, R. laciniata, R.
subtomentosa, R. triloba, Schrankia uncinata, Silphium integrifolium, S. laciniatum, S.
perfoliatum, S. speciosum, S. terebinthinaceum, Solidago missouriensis, S. rigida, S.
serotina, Teucrium canadense, Trifolium pratense, Verbena stricta, Vernonia baldwinia,
V. baldwinia interior, V. fasciculata, Verbesina helianthoides, Zinnia.

Macrocera obliqua Say, 1837. Boston Jour. Nat. Hist. 1: 403. ¢.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 259-261, fig. 80
(redescription). —Rozen, 1964. Amer. Mus. Novitates 2170: 9-12, figs. 6, 7, 9-13 (larva).

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24: 35. —Custer, 1928. Canad. Ent. 60: 28.
—Custer, 1929. Psyche 36: 293 (nesting habits, cocoon construction). —Rozen, 1964. Amer.
Mus. Novitates 2170:1-13, 15 figs. (nesting habits, parasites, ecology).

texana eluta (LaBerge). Deserts of south. Calif., Ariz.; Mexico (Chihuahua, Coahuila). Pollen:
Unknown, but visits flowers of Asparagus, Asteraginus, Helianthus annuus,
Heterotheca subaxillaris.
Melissodes (Epimelissodes) texana eluta LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1000. ¢,
OF

texana texana (Cresson). Kans. and Okla., south to Tex., west to N. Mex. and Colo. Pollen:
Unknown, but visits flowers of Chrysothamnus, Flaveria angustifolia, Helianthus,
Prionopsis ciliatus.
Melissodes texana Cresson, 1878. Amer. Ent. Soc., Trans. 4: 276. 2, do.
Melissodes texana flaveriae Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 108. @.

Superfamily APOIDEA 2135

SPECIES GROUP SILA

pallidior LaBerge. Calif. (Riverside Co.). Pollen: Unknown, but visits flowers of
Hoffmanseggia.
Svastra (Epimelissodes) pallidior LaBerge, 1963. Kans. Ent. Soc., Jour. 36: 52. 2, d.
sila (LaBerge). Ariz., Tex.; Mexico (Baja California and Chihuahua). Pollen: Oligolege of
Compositae especially Baileya pleniradiata and Heterotheca psammophila, but also
visits flowers of Heterotheca subaxillaris, Kallstroemia grandiflora, Psilostrophe
cooperi.

Melissodes (Epimelissodes) sila LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1002. d, 9.
Taxonomy: LaBerge, 1963. Kans. Ent. Soc., Jour. 36: 54 (geogr. and floral records).
Biology: Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 14-15 (floral relationships).

SPECIES GROUP PETULCA

petulca petulca (Cresson). Eastern Tex. north through Kans. to IIl., east to Fla., and north

along coast to N. J. Pollen: Unknown, but visits flowers of Borrichia frutescens,
Coreopsis cardaminefolia, Gaillardia pulchella, Gossypium herbaceum, Helenium
tenuifolium, Helianthus, Lepachys pinnata, Lithospermum canescens, Monarda
citriodora, M. punctata, Pycnanthemum flexuosum, Pyrrhopappus geiseri, Ratibida
columnaris, R. columnifera pulcherrima, Rudbeckia amplexicaulis, R. bicolor, R.
submentosa, Verbesina helianthoides.

Melissodes petulca Cresson, 1878. Acad. Nat. Sci. Phila., Proce. 30: 201. °@.

Melissodes illinoensis Robertson, 1895. Amer. Ent. Soe., Trans 22: 126. 9, 6.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 261-262, fig. 80
(redescription).
petulea suffusa (Cresson). Tex. and Kans. west to Colo. and Ariz.; Mexico (Chihuahua, Nuevo
Leon, Tamaulipas). Pollen: Unknown, but visits flowers of Baccharis, Borrichia
frutescens, Chrysopsis hispida, Coreopsis, Gaillardia pulchella, G. suavis, Helenium
autumnale, H. microcephalum, Helianthus annuus, Medicago sativa, Monarda
citriodora, M. punctata, Nepeta cataria, Opuntia, Phacelia, Ratibida columnaris, R.
tagetes, Rudbeckia hirta, Sphaeralcea, Tetragonothera ludoviciana, Thelespermo
megapotamicum.
Melissodes suffusa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 203. 2, ¢.
Melissodes townsendi Cockerell, 1896. Entomologist 29: 304. ¢.
sabinensis laterufa (Cockerell). Tex., N. Mex., ?Ariz. (Baboquivari Mts.); Mexico (Coahuila,
Durango).
Melissodes suffusa var. laterwfa Cockerell, 1934. Amer. Mus. Novitates 697: 10. 2.
sabinensis nubila (LaBerge). South. Calif.; Mexico (Baja California). Pollen: Unknown, but
visits flowers of Bebbia juncea, Chrysanthemum segetum, Chrysothamnus, Coreopsis
lanceolata, Corethrogyne filaginifolia, Encelia farinosa, Gutierrezia californica, G.
sarothrae, Haplopappus vernonioides, Helianthus annuus, Hemizonia paniculata,
Heterotheca grandiflora, Senecio douglasii, Stephanomeria exigua.
Melissodes (Epimelissodes) sabinensis nubila LaBerge, 1956. Kans. Univ. Sci. Bul. 37:
LO ZA SGr er
sabinensis sabinensis (Cockerell). South Calif., Ariz., N. Mex. Pollen: Unknown, but visits
flowers of Acacia, Dalea albiflora, Haplopappus, Heterotheca, Kallstroemia grandiflora,
Petalostemon, Psilostrophe cooperi, Sida diffusa, Sphaeralcea, Verbesina, Viguiera.
Melissodes sabinensis Cockerell, 1924. Amer. Mus. Novitates 113: 1. 6.

Genus SVASTRA Subgenus IDIOMELISSODES LaBerge

Melissodes subg. Idiomelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1027.
Type-species: Melissodes duplocincta Cockerell. Monotypic and orig. desig.

Taxonomy: LaBerge, 1957. Amer. Mus. Novitates 1837: 5, 7, 23-25 (generic status).
—LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 284 (as subgenus of Svastra).

2136 Hymenoptera in America North of Mexico

duplocincta (Cockerell). N. Mex. to Calif. (Colorado Desert); Mexico (Baja California,
Chihuahua and Coahuila). Pollen: Collects pollen from flowers of Ferrocactus wislizent
and is probably an oligolege of Ferrocactus and possibly other cacti, but visits other
flowers for nectar including Asclepias, Cevallia sinuata, Eriogonum, Gossypium
herbaceum, Hoffmanseggia densiflora, Lippia wrightii, Melochia tomentosa,
Sphaeralcea, Wislizenia refracta var. mamillata.
Melissodes duplocincta Cockerell, 1905. Psyche 12: 108. d, °.

Biology: Zavortink, 1975. Pan-Pacific Ent. 51: 236-240, tables 1, 2 (host plants, behavior,
distribution).

Genus XENOGLOSSODES Ashmead

?Tetraloniella Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 61.

Type-species: Macrocera graia Eversmann. Monotypic and orig. desig.
Xenoglossodes Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 63.

Type-species: Melissodes albata Cresson. Monotypic and orig. desig.

Revision: Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 74-92.

albata (Cresson). Ill., S. Dak., Kans., Colo., Okla., Tex. Pollen: Unknown, but visits flowers of
Petalostemon purpureum.
Melissodes albata Cresson, 1872. Amer. Ent. Soc., Trans. 4: 281. 9, 6.

Taxonomy: Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 474. d, °.
arizonica Cockerell. Ariz., Calif. (Imperial Co.).
Xenoglossodes arizonica Cockerell, 1937. Amer. Mus. Novitates 948: 7. d.
bishoppi (Cockerell). Tex.
Melissodes bishoppi Cockerell, 1914. Canad. Ent. 46: 414. d.
davidsoni (Cockerell). Calif.; Mexico (Baja California). Pollen: Unknown, but visits flowers of
Convolvulus occidentalis, Coreopsis maritima, Encelia californica.
Xenoglossa davidsoni Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 28. 2.
Synhalonia hirsutior Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 29. 6.

Taxonomy: Cockerell, 1935. Pan-Pacific Ent. 11: 58. d.

eriocarpi (Cockerell). Tex., N. Mex., Ariz., south. Calif. Pollen: Unknown, but visits flowers of
Larrea tridentata, Sideranthus gracilis.
Exomalopsis eriocarpi Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 453. ?.

Biology: Hurd and Linsley, 1975. Smithsn. Contr. Zool. 193: 42 (floral relationships).

excurrens Cockerell. Colo., N. Mex. Pollen: Unknown, but visits flowers of Chrysothamnus
wrightii, Verbesina encelioides.
Xenoglossodes excurrens Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 448. °.

Taxonomy: Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 259. —Cockerell, 1923. Ent. News
34: 50.
gutierreziae Cockerell. N. Mex. Pollen: Unknown, but visits flowers of Gutierrezia.
Xenoglossides gutierreziae Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 16: 218. °.
habrocoma Cockerell. Tex.
Xenoglossodes habrocoma Cockerell, 1935. Amer. Mus. Novitates 766: 7. GarOR
helianthorum Cockerell. Tex. Pollen: Unknown, but visits flowers of Helianthus.
Xenoglossodes helianthorum Cockerell, 1914. Canad. Ent. 46: 415. d.
imitatrix Cockerell and Porter. N. Mex. Pollen: Unknown, but visits flowers of Sphaeralcea

lobata.
Xenoglossodes imitatrix Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 407.
Ghee
lippiae lippiae (Cockerell). N. Mex., Ariz. Pollen: Unknown, but visits flowers of Lippia
wrightii, Phacelia congesta.
Synhalonia crenulaticornis lippiae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 25. d.

Taxonomy: Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 16: 224. °.

Superfamily APOIDEA 2137

lippiae semilippiae (Cockerell). Ariz.
Synhalonia lippiae semilippiae Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 179. 9.
neotomae Cockerell. N. Mex.
Xenoglossodes neotomae Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 314. 2, 6.
pallidicauda Cockerell. Tex.
Xenoglossodes pallidicauda Cockerell, 1934. Amer. Mus. Novitates 697: 11. 2.
parksi (Cockerell). Tex. (vicinity of San Antonio). Pollen: Unknown, but visits flowers of
Convolvulus hermannioides.
Melissodes agilis var. parksi Cockerell, 1935. Amer. Mus. Novitates 766: 5. 3.
pimella (Cockerell). Ariz. ;
Melissodes pimella Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 363. 3.
pomonae (Cockerell). Calif. (cismontane). Pollen: Possibly an oligolege of Hemizonia including
H. fasciculata, H. parryi, H. virgata.
Tetralonia pomonae Cockerell, 1915. Pomona Jour. Ent. Zool. 7: 230. é.
Tetralonia pomona(!) Cockerell, 1930. Ann. and Mag. Nat. Hist. (10) 5: 410.
spissa (Cresson). Colo., Tex.
Melissodes spissa Cresson, 1872. Amer. Ent. Soe., Trans. 4: 280. ?.
wilmattae Cockerell. Tex.
Xenoglossodes wilmattae Cockerell, 1917. In W. P. Cockerell, N. Y. Ent. Soc., Jour. 25: 191.
OF

Genus PEPONAPIS Robertson

Although the genus Peponapis is present in both North and South America, different species
occupy each continent, and while these faunas are in near geographic contact in the Panamani-
an-Colombian region they are evidently isolated from one another. Like the bees of the genus
Xenoglossa, they are solely dependent upon the pollen and to a large extent upon the nectar of
both cultivated and uncultivated Cucurbita (squashes, gourds and pumpkins). Also as in
Xenoglossa, the males commonly spend most of the day and a good part of the night in the
wilted and closed flowers of these plants. It is not unusual to encounter females at the beginning
of the nesting season in the wilted and closed flowers. One species, Peponapis pruinosa (Say)
has been introduced into Hawaii (Oahu and Hawaii), but it is not known whether the species has
become successfully established. The pollen collecting devices of these bees are species-specific
and apparently this has influenced the ability of the different species to collect and utilize pol-
lens of the various Cucurbita, both wild and domestic. At least several of the species have ex-
tended their ranges well beyond their original distribution following the spread and develop-
ment of cultivated Cucurbita. Unquestionably these bees are the most important pollinators of
the squashes, gourds and pumpkins.

Revision: Hurd and Linsley, 1964. Hilgardia 35: 425-472, figs. 1, 2, 12-18 (U. S. spp.). —Hurd
and Linsley, 1966. Ent. Soc. Amer., Ann. 59: 835-851, 12 figs., 1 table, 8 maps (U. S. and
Mexican spp.). —Hurd and Linsley, 1970. Calif. Univ. Pubs. Ent. 62: 1-39, 11 figs., 4 tables,
3 maps (classification).

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 244-246, fig. 72 (eastern U. S.
sp.). —Rozen, 1965. Amer. Mus. Novitates 2233: 6, 11, figs. 20-27 (larva).

Biology: Michelbacher, Smith and Hurd, 1964. Calif. Agr. 18: 2-4, figs. (pollination of
squashes, gourds and pumpkins). —Hurd and Linsley, 1964. Hilgardia 35: 376-382, 2 tables
(biol. summary). —Michelbacher, Hurd and Linsley, 1968. Bee World 49: 159-167, 6 figs.
(feasibility of introducing squash bees into the Old World). —Hurd, Linsley and Whitaker,
1971. Evolution 25: 218-234, 4 figs., 3 tables (squash bees and origin of cultivated
Cucurbita): —Michelbacher, Hurd and Linsley, 1971. Bee World 52: 156-166, 4 figs.
(experimental introduction of squash bees to improve yields of squashes, gourds and
pumpkins).

Genus PEPONAPIS Subgenus EOPEPONAPIS Hurd and Linsley

Peponapis subg. Eopeponapis Hurd and Linsley, 1970. Calif. Univ. Pubs. Ent. 62: 20.
Type-species: Xenoglossa utahensis Cockerell. Orig. desig.

2138 Hymenoptera in America North of Mexico

michelbacherorum Hurd and Linsley. South Ariz.; Mexico (Nayarit, Sinaloa and Sonora).
Pollen: Oligolectic on uncultivated mesophytic Cucurbita, viz., Lundelliana and Sororia
groups, as well as most, if not all, domestic Cucurbita grown within its range.
Peponapis michelbacherorum Hurd and Linsley, 1964. Hilgardia 35: 437, fig. 14. d, @.
utahensis (Cockerell). Ariz., N. Mex., Tex., 7?Utah; Mexico to Costa Rica. Pollen: Oligolectic on
uncultivated, mesophytic Cucurbita, viz., Lundelliana and Sororia groups, as well as
most, if not all, domestic Cucurbita grown within its range. The species was described
from a specimen collected prior to the admission of Arizona and New Mexico into the
Union and there is a possibility that it had been collected even before Utah gained
statehood. Thus the northernmost contemporary record of the species from Tempe,
Arizona may be as far north as the species occurs.
Xenoglossa utahensis Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 182. 9.

Genus PEPONAPIS Subgenus PEPONAPIS Robertson

Peponapis Robertson, 1902. Canad. Ent. 34: 324.
Type-species: Macrocera pruinosa Say. Orig. desig.

PRUINOSA SPECIES GROUP

limitaris (Cockerell). South. Tex.; Mexico to Panama. Pollen: Oligolectie on uncultivated,
mesophytie Cucurbita including C. fraterna and C. martinezii, as well as most, if not all,
domestic Cucurbita.
Xenoglossa pruinosa limitaris Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 18: 73. 3.
Tetralonia tenuifasciata Friese, 1916. Stettin. Ent. Ztg. 77: 295, 332. 2, d. Preoce.
Tetralonia tenuimarginata Friese, 1921. Stettin. Ent. Ztg. 82: 75, 78. N. name.
pruinosa (Say). Maine to Ga., west to Idaho and Calif.; Mexico (Central Plateau and west
coast, south to Oaxaca). Pollen: Oligolectic on uncultivated xerophytic Cucurbita
including C. foetidissima and C. galleotti as well as most, if not all, domestic Cucurbita.
The geographic range of the species has been significantly expanded following the
development and widespread cultivation of domestic Cucurbita. The species has been
introduced into Hawaii (Oahu and Hawaii), but it is not known whether it has become
established. Attempts have also been made to introduce this species into New Zealand,
but so far these have been unsuccessful.
Macrocera pruinosa Say, 1837. Boston Jour. Nat. Hist. 1: 405. ¢, 2.
Xenoglossa spriuna Howard, 1901. Insect Book, PI. Viii, fig. 2. 6. Lapsus calami.
Xenoglossa angelica Cockerell, 1902. Ent. News 13: 103. 6, @.
Xenoglossa (Peponapis) howardi Cockerell, 1918. Ann. and Mag. Nat. Hist. (9) 2: 420. 3,
Q

Xx enoglossa pruinosa var. lutzi Cockerell, 1923. Canad. Ent. 55: 205. 2, d.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7): 34. —Mathewson, 1968. Kansas Ent.
Soc., Jour. 41: 255-261, 1 fig. (nest construction and life history). —Michelbacher, Hurd and
Linsley, 1971. Bee World 52: 156-166, 4 figs. (experimental introductions). —Hurd, Linsley
and Michelbacher, 1974. Smithsn. Contrib. Zool. 168: 1-17, 4 figs., 8 tables (nat. history).

Morphology: Mathewson, 1965. Kans. Ent. Soc., Jour. 28: 209-233, 25 figs. (internal).

Genus PEPONAPIS Subgenus XEROPEPONAPIS Hurd and Linsley

Peponapis subg. Xeropeponapis Hurd and Linsley, 1970. Calif. Univ. Pubs. Ent. 62: 28.
Type-species: Peponapis timberlakei Hurd and Linsley. Monotypic and orig. desig.
timberlakei Hurd and Linsley. Deserts, N. Mex. to Calif., Nev.; Mexico (Baja California and
Sonora). Pollen: Oligolectie on uncultivated, xerophytic Cucurbita including C. digitata
and C. palmata of the Digitata group. There is no evidence that the species collects
pollen from either the xerophytic C. foetidissima or domestic Cucurbita even though
some specimens including females have been collected in these flowers.
Peponapis timberlakei Hurd and Linsley, 1964. Hilgardia 35: 428, figs. 2, 12. gd, 9.

Genus PEPONAPIS Subgenus XENOPEPONAPIS Hurd and Linsley
Peponapis subg. Xenopeponapis Hurd and Linsley, 1970. Calif. Univ. Pubs. Ent. 62: 29.

Superfamily APOIDEA =. 21389

Type-species: Melissodes crassidentata Cockerell. Monotypic and orig. desig.

crassidentata (Cockerell). Southern Tex.; Mexico to Costa Rica. Pollen: Oligolectic on
uncultivated Cucurbita including C. gracilior, C. kellyana, C. lundelliana, C. martinezii,
C. radicans, and C. sororia as well as most, if not all, domestic Cucurbita.
Melissodes crassidentata Cockerell, 1949. U. S. Natl. Mus., Proc. 98: 466. 6.

Genus MELISSODES Latreille

This large genus, which although present in both North and South America as well as the
West Indies, is especially well represented by numerous species in North America. Of the eight
currently recognized subgenera only the subgenus Ecplectica does not extend into America
north of Mexico.

Revision: LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1032-1194, 129 figs. (Pt. I of a gen. rev. of
North and Centr. Amer. spp.). —LaBerge, 1956. Kans. Univ. Sci. Bul. 38: 533-578, 13 figs.
(Pt. II of a gen. rev. treating the subgenera Tachymelissodes, Apomelissodes,
Psilomelissodes, Heliomelissodes). —LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 283-663, 125
figs. (Pt. III (final) of a gen. rev. treating the subgenera Callimelissodes and
Eumelissodes).

Taxonomy: Robertson, 1905. Amer. Ent. Soc., Trans. 31: 367-371 (Illinois species).
—Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 521-522 (large species). —Cockerell,
1906. Amer. Ent. Soc., Trans. 32: 74-92. —Michener, 1953. Kans. Univ. Sci. Bul. 35: 1073,
figs. 199-205 (larva). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 263-310, figs.
67-85, 1 table (eastern U.S. spp.). —LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 227-242
(geogr. records, n. spp., floral information).

Genus MELISSODES Subgenus MELISSODES Latreille

Melissodes Latreille, 1829. In Cuvier, Regne Animal, ed. 2, v. 5, p. 354.
Type-species: Melissodes fonscolombei Romand. First included species, 1841.
Melissoda Drapiez, 1841. Dict. Classique Sci. Nat., v. 4, p. 241. Emend. Preoce.
bimaculata bimaculata (Lepeletier). Maine west to east. N. Dak. and Colo., south to north.

Fla. and east. Tex. Parasite: Triepeolus lunatus concolor Robt. Pollen: Unknown, but
visits flowers of Abutilon theophrasti, Agastache nepetoides, Althaea rosea, Arctium
minus, Asclepias tuberosa, A. verticillata, Asparagus, Aster novae-angliae, A.
paniculata, Astragalus canadensis, Baptisia tinctoria, Bidens aristosa, Blephilia
hirsuta, Brauneria purpurea, Cacalia reniformis, Campanula americana, C.
rotundifolia, Cassia fasciculata, Cephalanthus occidentalis, Cicuta maculata, Cirsiwm
lanceolatum, Convolvulus sepium, Cuphea peliolata, Cucurbita pepo, Dalea onobrychis,
Desmodium bracteosum, D. canadense, D. dillenii, D. paniculatum, Dianthera
americana, Dipsacus sylvestris, Echinocystis lobata, Eupatorium coelestinum, Gaura
biennis, Gerardia grandiflora, Gladiolus, Gossypium herbaceum, Grindelia, Helenium
autumnale, Helianthus annuus, H. divaricatus, H. grosse-serratus, H. tuberosus,
Hibiscus lasiocarpus, H. militaris, Impatiens biflora, Ipomoea pandurata, I. purpurea,
Jacquemontia temnifolia, Lepachys pinnata, Lespedeza procumbens, Lobelia
leptostachys, L. siphillitica, Lythrum alatum, Malva rotundifolia, M. sylvestris,
Medicago sativa, Melilotus alba, M. officinale, Mentha, Monarda fistulosa, M. mollis,
M. punctata, Nepeta cataria, Oenothera biennis, Oxalis stricta, Petalostemon
purpureum, Petunia, Phystostegia virginiana, Platycodon grandiflorum, Polygonum
pennsylvanicum, Prunella vulgaris, Pycnanthemum flexuosum, Ratibida, Rhus
copallina, Rudbeckia triloba, Sagittaria, Scrophularia marilandica, Scutellaria
lateriflora, Seymeria macrophylla, Sicyos angulatus, Silphium laciniatum, S.
perfoliatum, Siscanna, Solidago, Stachys palustris, Strophostylis, Symphoricarpos
occidentalis, S. orbiculata, Teucrium canadense, Trifolium pratense, Verbena hastata,
V. stricta, V. urticaefolia, V. baldwini interior, Vernonia fasciculata, V. spicata,
Veronica virginica, Vitex agnus-castus.

Macrocera bimaculata Lepeletier, 1825. Encycl. Method. Ins., v. 10, p. 528. °.

Macrocera binotata Say, 1837. Boston Jour. Nat. Hist. 1: 404. 2, ¢.

Macrocera nigra Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 112. ¢.

2140 Hymenoptera in America North of Mexico

Melissodes melanosoma Cockerell, 1905. Canad. Ent. 37: 266. 6.

Biology: Ashmead, 1894. Psyche 7: 25 (nest). —Banks, 1902. N. Y. Ent. Soc., Jour. 10: 209
(nest). —Folsom, 1922. Ent. Soc. Amer., Ann. 15: 183 (nest). —Rau, 1922. Acad. Sci. St.
Louis, Trans. 24: 34 (nest).

bimaculata nulla LaBerge. South. Fla. and Keys. Pollen: Unknown, but visits flowers of
Lythrum lineare.
Melissodes (Melissodes) bimaculata nulla LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1109.
Cra o
blanda LaBerge. Tex., Okla. Pollen: Unknown, but visits flowers of Borrichia frutescens,
Gaillardia, Opuntia, Sphaeralcea.
Melissodes (Melissodes) blanda LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1084. d, 2.
colliciata Cockerell. Ariz., Mexico (Chihuahua, Durango, Jalisco, Zacatecas, Puebla, Michoacan,
Oaxaca). Pollen: Unknown, but visits flowers of Asclepias, Baccharis glutinosa,
Gutierrezia.
Melissodes colliciata Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 257. 6.
Melissodes (Melissodes) elusa LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1061. 6, 2.

Taxonomy: LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 655 (synonymy). — LaBerge, 1963. Nebr.
Univ. State Mus., Bul. 4: 234-235 (geogr. records).
communis alopex Cockerell. South. Calif. to B. C., east to Alta. and Utah. Pollen: Unknown,
but visits flowers of Asclepias erosa, Cirsium, Clarkia amoena, C. elegans, Cryptantha
intermedia, Datura meteloides, Durantia plumiert, Encelia californica, Eriodictyon
trichocalyx, Eriogonum fasciculatum polifolium, Gilia capitata, Grindelia camporum,
Hugelia virgata, Lotus scoparius, Malvastrum fasciculatum, Medicago sativa,
Melilotus, Mesembryanthemum crystallinum, Monardella lanceolata, Opuntia
littoralis, Penstemon, Phacelia ramosissima, Salvia apiana, S. carnosa, S. clevelandit,
Scabiosa, Sphaeralcea fasciculata, Stachys apargioides, Stephanomeria exigua,
Trifolium involucrata.
Melissodes alopex Cockerell, 1928. Psyche 35: 238. ¢.
communis communis Cresson. Southeastern Ariz., north through eastern Colo. and Wyo. to
N. Dak. and east through Ill. and Ind. in the north and through Tex. in the south to the
Atlantic. Pollen: Polylectic, appears to prefer pollen from flowers of Leguminosae and
Labiatae, but visits a wide variety of flowers including Abutilon theophrasti, Althaea
rosea, Amorpha, Asclepias syriaca, A. tuberosa, Baptisia, Befaria racemosa, Blephilia
hirsuta, Brazoria truncata, Campanula, Cassia fasciculata, Chrysopsis, angustifolia,
Cirsium discolor, C. lanceolatum, Cleome serrulata, Convolvulus, Croton, Cucurbita,
Cyrilla parviflora, Dalea multiflora, Dianthera americana, Echium vulgare,
Gossypium herbaceum, Grindelia, Helenium tenuifolium, Helianthus annuus, H.
lenticularis, Heliotropium, Hyrta, Ipomoea, Lactuca, Lythrum alatum, L. lineare,
Malva sylvestris, Medicago sativa, Melilotus alba, Mentha, Monarda citriodora, M.
fistulosa, M. pectinata, M. punctata, Nepeta cataria, Oenothera elliptica, O. laciniata,
Opuntia, Passiflora incarnata, Petalostemon occidentale, P. purpureum, P. violaceum,
Phaseolus, Platycodon grandiflorium, Prionopsis, Proboscidea louisianica,
Pycnanthemum flexuosum, Ratibida columnifera, Rudbeckia, Rhus glabra, Salvia,
Sidalcea reticulata, Solanum elaeagnifolium, Solidago serotina, Teucrium canadense,
Thelesperma megapotamicum, Verbena stricta, Vernonia baldwini interior, V. glauca.
Melissodes communis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 204. 2, 6.
Melissodes hortivagans Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 180. 3, 2.
Melissodes Martini Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 526. 9.
Melissodes variabilis Robertson, 1905. Amer. Ent. Soc., Trans. 31: 368. 2, 6.
Melissodes xanthopteralis Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 362. d.
Melissodes manni Cockerell, 1924. Amer. Mus. Novitates 113: 1. ¢.
Melissodes hortivagans melanotica Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 231.
oF
comptoides Robertson. N. J. to Fla., west to S. Dak., Colo., Ariz. Pollen: Polylectic, appears to
prefer pollen of Euphorbia marginata, Medicago sativa and Melilotus alba, but visits
other flowers including Amphiachyris dracunculoides, Asclepias incarnatus, Aster,

Superfamily APOIDEA 2141

Blephilia hirsuta, Campanula americana, Cassia fasciculata, Cleome serrulata, Cicuta
maculata, Cirsium lanceolatum, Diodea teres, Gossypium herbaceum, Helenium
tenuifolium, Helianthus annuus, H. petiolaris, Lepachys pinnata, Lespedeza virginica,
Ludwigia alternifolia, Lycopus americanus, Lythrum alatum, L. lineare, L. salicaria,
Monarda citriodora, M. fistulosa, M. punctata, Petalostemon purpureum, Prunella
vulgaris, Pcynanthemum flecuosum, P. pilosum, P. virgianum, Ratibida, Solidago
serotina, Symphoricarpos, Taxaracum officinale, Teucrium canadense, Verbena hastata,
V. stricta, Vernonia fasciculata, V. longifolia, V. noveboracensis.

Melissodes comptoides Robertson, 1898. Acad. Sci. St. Louis, Trans. 8: 52. 2, d.

Melissodes martini hitei Cockerell, 1908. Ann. and Mag. Nat. Hist. (8) 2: 331. °.

gilensis gilensis Cockerell. Ariz., N. Mex., Colo. Pollen: Polylectic, visits flowers of Arabis,

Asclepias tuberosa, Cercidium torreyanum, Cirsium, Lippia lyciddea, Lotus brightii,
Lupinus, Malva, Melilotus alba, M. officinalis, Monarda menthaefolia, M. stricta,
Nolina, Robinia neomexicana. Another subspecies, Melissodes gilensis crenata
LaBerge, occurs in Mexico.

Melissodes gilensis Cockerell, 1896. Entomologist 29: 306. 9, d.

Melissodes epicharina Cockerell, 1905. Psyche 12: 99. °.

maesta LaBerge. Tex.; Mexico (Coahuila, San Luis Potosi and Zacatecas). Pollen: Unknown,

but visits flowers of Helenium, Monarda citriodora, Phacelia.

Melissodes (Melissodes) maesta LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1095. 6, 9.

paroselae Cockerell. South. Calif. to southwest. Tex.; Mexico (Baja Calif., Sonora, Chihuahua,

Nayarit). Pollen: Polylectic, visits flowers of Acacia greggti, Argemone, Asclepias
subulata, Aster canescens, Cucurbita, Cleome luteum, Dalea albifrons, Dasylirion
wheeleri, Eriogonum trichopes, Franseria eriocentra, Gaillardia, Gossypium herbaceum,
Haplopappus heterophylla, Helianthus annuus, Heterotheca subaxillaris, Kallstroemia
grandiflora, Larrea tridentata, Leucophyllum frutescens, Lippia, Lygodesmia juncea,
Medicago sativa, Opuntia, Parosela scoparia, Pectis papposa, Petalostemon, Prosopis
glandulosa, Psilostrophe cooperi, Pyrrhopappus multicaulis, Rhus, Salix, Senecio
longilobus, Thurberia thespesioides, Verbesina encelioides, V. exauriculata, Wedelia
incarnata, Wislizenia palmeri, W. refracta, Zinnia grandiflora.

Melissodes parosetae Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 528. ¢.

Melissodes paroselae Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 16: 477. Justified
emendation of parosetae.

Melissodes Helenae Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 365. 2.

Melissodes communis gratior Cockerell, 1923. Calif. Acad. Sci., Proc. 12: 85. 9.

Biology: Mathewson and Daly, 1955. Kans. Ent. Soc., Jour. 28: 120. —Linsley, 1962, Ent. Soc.
Amer., Ann. 55: 161, fig. 8. —Butler, Todd, McGregor and Werner, 1962. Ariz. Univ. Agr.
Expt. Sta. Tech. Bul. 139: 1-11. —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 41
(floral relationships with Larrea).

tepaneca Cresson. Kans., Ill. and N. C. south through Tex. and Mexico to Panama. Pollen:

Polylectic, visits flowers of Abutilon avicennes, A. theophrasti, Agastache breviflora,
Asclepias syriaca, A. tuberosa, Aster tanacetifolium, Baccharis, Borrichia frutescens,
Brazoria truncata, Callirhoe involucrata, Cephalanthus occidentalis, Cercidium
texanum, Coreopsis palmata, Cornuta grandifloria, Cuphea balsamona, Dalea grisea,
Davilla knuthii, Donnellsmithia hintonti, Eryngium leavenworthu, Eysenhardtia
polystacha, Gaillardia suavis, Gossypium herbaceum, Grindelia, Helenium
microcephalum, Hibiscus tiliaceus, Ipomoea longifolia, I. murucoides, I. triloba,
Lactuca pulchella, Lindheimeria texana, Lippia, Lythrum alatum, L. lanceolatum,
Marrubium vulgare, Medicago, Monarda punctata, Opuntia lindheimeri, Parkinsonia,
Petalostemon multiflorium, P. purpureum, P. violacewm, Phacelia, Phlox, Prosopis,
Ratibida columnifera pulcherrima, Rubus, Rudbeckia, Salvia, Sida acuta, Sphaeralcea,
Sisyrinchium campestre, Teucrium canadense, Verbesina encelioides, Verbena
officinalis, V. stricta, Vernonia aschenborniana.

Melissodes tepaneca Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 210. 2 in part (d
misdet.).

Melissodes petalostemonis Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 58. °.

Melissodes galvestonensis Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 181. 2, d.

2142 Hymenoptera in America North of Mexico

Melissodes bruesi Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 110. @.

Melissodes loena Cockerell, 1909. Entomologist 42: 148. ¢.

Melissodes masuca Cockerell, 1909. Entomologist 42: 148. 3.

Melissodes tepaneca aschenborniana Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 10: 28.
é.

Melissodes aurescens Cockerell, 1949. U. S. Natl. Mus., Proc. 98: 462. °.

tepida tepida Cresson. North. Utah to Calif. (Sierra Nevada Mts.), north to south. Idaho and

northwest. Oreg. Pollen: Presumably polylectic, visits flowers of Asclepias, Astragalus
bolanderi, Glycyrrhiza lepidota, Helianthus annuus, Medicago sativa, Melilotus,
Mentha, Trifolium pratense.

Melissodes tepida Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 210. ?.

tepida timberlakei Cockerell. South. Calif. northeast to Sierra Mts., north to southern Oreg.
west of Sierras. Parasite: Triepeolus lineatulus Ckll. and Sandh., T. timberlakei Ckll.
Pollen: Polylectic, visits the flowers of many species of plants for pollen and or nectar
evidently preferring the pollen of Leguminosae, Labiatae, Euphorbiaceae, but also visits
Althaea rosea, Artemisia, Asclepias eriocarpa, Aster, Bellis, Brassica adpressa, B.
geniculata, B. incana, Centaurea cyanus, Centromadia pungens, Chrysanthemum,
Clarkia amoena, Cleomella, Coreopsis lanceolata, Croton californicus, Cryptantha
intermedia, Cucurbita, Daucus carota, Durantia plumieri, Eremocarpus setigerus,
Eriastrum virgatum, Eriogonum fasciculatum, E. gracile, E. involucratum,
Eschscholzia californica, Geranium, Gilia capitata, Gossypium, Gutierrezia californica,
G. sarothrae, Helianthus annuus, H. bolanderi, H. petiolaris, Heliotropiwm
curassavicum, Hemizonia pungens, Hugelia virgata, Ipomoea, Lippia filiformis, L.
lanceolata, Lotus americanus, L. scoparius, Madia, Marrubium vulgare, Medicago
sativa, Melilotus alba, M. indica, Mesembryanthemum crystallinum, Papaver
heterophyllum, Phacelia ramosissima, Phaseolus, Phyla lanceolata, P. nodiflora rosea,
Plantago bigelovii, Pluchea borealis, P. camphorata, P. sericea, Raphanus, Rosa,
Salsola kali, Scabiosa, Senecio douglasti, Sicyos, Sida hederacea, Stachys ajugoides, S.
bullata, Stephanomeria exigua, S. virgata, Trichostema lanceolatum, Trifolium
mvolucratum, T. repens, Wislizenia refracta.

Melissodes timberlakei Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 18: 624. 2.

Biology: Linsley, 1946. Econ. Ent., Jour. 39: 20-23, 25 (nesting habits).

tepida yumensis LaBerge. Southeast. Calif., south. Nev., southwest. Utah, Ariz.; Mexico (Baja
Calif.). Pollen: Polylectic, visiting flowers of Aster spinosa, Cercidium torreyanum,
Chilopsis linearis, Citrullus, Cynodon dactylon, Gossypium, Lippia brevipes, Medicago
sativa, Melilotus alba, Phyla nodiflora, Pluchea, Salix, Sida hederacea, Sphaeralcea
orcutti, Tamarix gallica.

Melissodes (Melissodes) tepida yumensis LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1128. 6,

OF

Biology: Butler, Todd, McGregor and Werner, 1962. Ariz. Univ. Agr. Expt. Sta. Tech. Bul.
139: 1-11 (floral relationships).
tessellata LaBerge. South. coastal Calif.; Mexico (Baja Calif., Jalisco). Pollen: Polylectic, visits
a wide variety of flowers for pollen and or nectar including Acacia greggti, Asclepias
erosa, Baccharis glutinosa, Bebbia juncea, Brassica geniculata, Centaurea melitensis,
Chrysanthemum segetum, Chrysothamnus veganus, Clarkia amoena, C. elegans, -
Corethrogyne bernardina, C. filaginifolia, Croton californicus, Cucurbita foetidissima,
Durantia plumieri, Encelia farinosa, Eriogonum fasciculatum, Gutierrezia bernardina,
G. californica, G. sarothrae, Haplopappus palmeri, H. vernonioides, Helianthus annuus,
Hemizonia fasciculatum, H. paniculata, Hugelia virgata, Lippia filiformis, Lotus
scoparius, Malvastrum fasciculatum, Marrubium vulgare, Medicago sativa, Melilotus
alba, Monardella villosa, Opuntia parryi, Parkinsonia aculeata, Pluchea camphorata,
Polygonum lapathifolium, Salvia apiana, Senecio douglasii, Stephanomeria virgata,
Trichostema lanceolatum, Wislizenia refracta.
Melissodes (Melissodes) tessellata LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1157, figs.
101-103. 3, 9.

Superfamily APOIDEA 2143

thelypodii thelypodii Cockerell. West Tex. to southeast. Calif.; Mexico (Chihuahua, Coahuila,

Durango). Pollen: Polylectic, takes pollen from a wide variety of plants including
Carthamnus, Gossypium, Kallstroemia grandiflora, but also visits the flowers of
Asclepias, Astragalus cottonii, Baccharis glutinosa, Chamaesaracha coronopus,
Cirsium, Convolvulus, Gaillardia, Helianthus annuus, Hoffmanseggia jamesii, Ipomoea
mexicana, Larrea tridentata, Lippia cuneifolia, Lygodesmia juncea, Medicago sativa,
Melilotus, Solanum elaeagnifolium, Sphaeralcea, Thelypodium linearifolium, Thurberia
thespesioides, Wedeliella incarnata. Another subspecies, Melissodes thelypodii stulta
LaBerge, occurs in Mexico (Colima, Durango, Jalisco, Michoacan, Oaxaca and San Luis
Potosi), Guatemala and Honduras.

Melissodes thelypodii Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 527. 9.

Melissodes kallstroemiae Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 16: 216. d.

Melissodes kallstroemiae var. phenacoides Cockerell, 1905. Ann. and Mag. Nat. Hist. (7)
16221725

Melissodes thurberiae Cockerell, 1914. Ent. Soc. Wash., Proc. 16: 31. 2.

Biology: Butler, Todd, McGregor and Werner, 1962. Ariz. Univ. Agr. Expt. Sta. Tech. Bul.
139: 1-11. —Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 14 (floral relationships).

Genus MELISSODES Subgenus APOMELISSODES LaBerge

Melissodes subg. Apomelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1175.
Type-species: Melissodes fimbriata Cresson. Orig. desig.
apicata Lovell and Cockerell. Maine to Fla. Pollen: Oligolectic, obtains both pollen and nectar
from flowers of Pontederia, but visits other flowers for nectar including Cirsium,
Hydrocotyle umbellata, Melilotus alba, Oenothera speciosa, Stachys floridana,
Trifolium hybridum.
Melissodes apicata Lovell and Cockerell, 1906. Psyche 13: 111. 2.

Taxonomy: LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 233-234 (geogr. and floral
records).
baileyi Cockerell. Tex. Pollen: Unknown, but visits flowers of Callirhoe and Gaillardia.
Melissodes Baileyi Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 361. 2.

Taxonomy: LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 231-233. ¢ (geogr. and floral
records).
fimbriata Cresson. Va. to Ga., west to Tex., Kans. Pollen: Oligolectic, obtains both pollen and
nectar from flowers of Oenothera, but visits other flowers for nectar including
Helianthus petiolaris, Kneiffia.
Melissodes fimbriata Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 203. 2, d.
mitchelli LaBerge. N. C., Ga., Fla. Pollen: Apparently oligolectic, visits flowers of Opuntia.
Melissodes (Apomelissodes) mitchelli LaBerge, 1956. Kans. Univ. Sci. Bul 38: 556, fig. 7.
Sp Se

Genus MELISSODES Subgenus HELIOMELISSODES LaBerge

Melissodes subg. Heliomelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1172.
Type-species: Melissodes desponsa Smith. Orig. desig.
desponsa Smith. N.S. to N. C., Ala., west to N. Dak. and Okla. Pollen: Oligolectic, obtains
pollen from flowers of Cirsium, but visits other flowers for nectar including Aster
novaeangliae, Brauneria purpurea, Carduus crispus, Cassia, Centaurea jacea, Coreops
aristosa, Helianthus annuus, H. grosse-serratus, Inula helenium, Monarda fistulosa,
Pontederia cordata, Pycnanthemum, Rudbeckia laciniata, Silphiwm, perfoliatum,
Solidago canadensis, S. puberula, Verbena stricta.
Melissodes desponsa Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 310. 9.
Melissodes daponsa(!) Bridwell, 1899. Kans. Acad. Sci., Trans. 16: 211.
Melissodes nigripes Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 311. 2 (¢ misdet.).
Melissodes cnici Robertson, 1901. Canad. Ent. 33: 230. 2, d.

2144 Hymenoptera in America North of Mexico

rivalis Cresson. B. C. and Calif. east to Man., Minn., south to Ariz., N. Mex. and Tex. Parasite:

Nemognatha dichroa LeC.? Pollen: Apparently oligolectic, obtains pollen from
composites of the tribe Cynareae, especially the genus Cirsium, but visits other flowers
presumably for nectar including Apocynum, Asclepias, Carduus caulescens, Centaurea
americana, Helianthus annuus, Penstemon cyananthus, Rudbeckia, Solidago,
Teucrium, Trifolium repens, T. pratense, Verbena.

Melissodes rivalis Cresson, 1872. Amer. Ent. Soe., Trans. 4: 277. 6.

Melissodes desponsiformis Cockerell, 1905. In Viereck, Canad. Ent. 37: 320. 2.

Melissodes mysops Cockerell, 1905. Entomologist 38: 146. 6, @.

Melissodes hexacantha Cockerell, 1905. Psyche 12: 100. 3.

Melissodes nigrosignata Cockerell, 1905. Psyche 12: 101. 9.

Melissodes habilis Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 229. ¢.

Biology: Scullen, 1928. Pan-Pacific Ent. 4: 176 (as mysops, nesting habits). —LaBerge, 1956.
Kans. Univ. Sci. Bul. 38: 571-572 (additional information on nesting habits based on notes
supplied by H. A. Scullen).

Genus MELISSODES Subgenus TACHYMELISSODES LaBerge

Melissodes subg. Tachymelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1170.
Type-species: Melissodes dagosa Cockerell. Orig. desig.

dagosa Cockerell. Idaho and Wash. to Calif. and Colo. Pollen: Unknown, but visits flowers of
Asclepias, Cirsium, Medicago sativa, Melilotus alba, Norta altissima, Sphaeralcea,
Thelypodium brachycarpa.
Melissodes dagosa Cockerell, 1909. Canad. Ent. 41: 128. 6.
Melissodes calloleuca Cockerell, 1924. Pan-Pacifie Ent. 1: 55. 2.
opuntiella Cockerell. Utah, Calif., Ariz., Tex.; Mexico (Coahuila Mexico and Zacatecas). Pollen:
Polylectic, obtains pollen from such flowers as Cirsium, Lindheimera texana,
Sphaeralcea pedatifolia, Verbesina encelioides, but visits other flowers including
Cercidium texanum, Erigeron compositus, Helenium microcephalum, Helianthus
annuus, Opuntia lindheimeri, Parkinsonia, Prosopis, Tetradymia canescens.
Melissodes opuntiella Cockerell, 1911. Canad. Ent. 43: 131. 6, °.
Melissodes albocincta Cockerell, 1919. Ann. and Mag. Nat. Hist. (9) 2: 119. °.
sphaeralceae Cockerell. Ariz., N. Mex., Tex.; Mexico (Aguascalientes, Distrito Federal, Mexico,
San Luis Potosi, Zacatecas). Pollen: Unknown, but visits flowers of Melilotus alba,
Monarda, Sphaeralcea angustifolia, S. fendleri.
Melissodes sphaeralceae Cockerell, 1896. Entomologist 29: 304. ¢.

Genus MELISSODES Subgenus EUMELISSODES LaBerge

Melissodes subg. Eumelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1177.
Type-species: Melissodes agilis Cresson. Orig. desig.

agilis Cresson. South. Canada, U. S., except Fla.; Mexico (Chihuahua, Coahuila, Durango).
Pollen: Apparently an oligolege of Helianthus including H. annuus, H. atrorubens, H.
bolanderi, H. ciliaris, H. coronarius, H. divaricatus, H. grosse-serratus, H. laetiflorus,
H. lenticularis, H. maximillianus, H. mollis, H. petiolaris, H. pumilis, H. radulus, H.
rigidus, H. salicifolius, H. scaberrimus, H. subrhomboideus, H. tuberosus, but visits a
wide variety of flowers: Abutilon theophrasti, Althaea rosea, Aplopappus spinulosus,
Arctium, Argemone platyceras, Bidens aristosa, B. laevis, Blephilia hirsuta, Brassica
juncea, Brauneria pallida, Carduus crispus, Carya pecan, Cassia chamaecrista, C.
fasciculata, Centromadia pungens, Chrysopsis hispidus, Chrysothamnus, Cirsium
altissimum, C. discolor, C. lanceolatum C. undulatum, Clematis, Cleome lutea, C.
serrulata, Convolvulus, Coreopsis lanceolata, C. palmata, C. tripteris, Cosmos, Datura
meteloides, Enceliopsis, Engelmannia pinnatifida, Ericameria palmeri, Eschscholzia
californica, Eupatorium purpureum, Eustoma artemifolium, Gaillardia cristata, Gilia,
Grindelia squarrosa, Gutierrezia sarothrae, Haplopappus palmeri, Helenium
autumnale, H. laciniatum, Heliopsis, Heliotropium, Hemizonia pungens, Hibiscus,
Ipomoea, Lactuca pulchella, Lepachys pinnata, Liatris pycnostachya, Medicago sativa,
Melilotus alba, Mentha canadensis, Monarda fistulosa, Penstemon, Pepo, Petalostemon

Superfamily APOIDEA 2145

occidentale, P. purpureum, Phacelia, Physostegia parviflora, Platycodon grandiflorum,
Pluchea camphorata, Prionopsis, Pycnanthemum flexuosum, P. pilosum,
Pyrrhopappus multicaulis, Rudbeckia hirta, R. laciniata, R. triloba, Schrankia
uncinata, Senecio, Silphium integrifolium, S. laciniatum, S. perfoliatum, S. speciosum,
Sium cicutaefolium, Solidago canadensis, S. serotina, S. trinervata, Teucrium
canadense, Verbena hastata, V. stricta, Verbesina encelioides, V. exauriculata, V.
occidentalis, Vernonia baldwini interior, V. baldwini occidentalis, V. fasciculata,
Veronica, Vitex agnus-castus, Wislizenia refracta.

Melissodes agilis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 204. 6.

Melissodes aurigenia Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 212. 2, 6.

Biology: Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7): 34 (nest site, burrow). —Custer, 1928.
Canad. Ent. 60: 28-31, 2 figs. (possibly this species using same burrow entrance as Svastra
obliqua Say).

appressa LaBerge. Calif. Pollen: Oligolege of autumnal flowering Compositae, especially
Gutierrezia, Haplopappus, Heterotheca, but visits other flowers including Alyssum
maritinum, Aster exilis, Centromadia pungens, Croton californicus, Grindelia
californica, G. camporum, Helianthus annuus, Heliotropium curassavicum, H.
oculatum, Hemizonia pungens, Lessingia glandulifera, Melilotus alba, Phacelia
crenulata, Pluchea camphorata, Senecio, Solidago confinis, S. occidentalis.
Melissodes (Eumelissodes) appressa LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 616. 9, ¢.
bicolorata LaBerge. Northern Calif., Nev., Utah. Pollen: Unknown, but visits flowers of
Chrysothamnus, Haplopappus arborescens, Melilotus, Penstemon palmeri.
Melissodes (Eumelissodes) bicolorata LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 441. 2, 6.
bidentis Cockerell. N. Dak. and Minn., east to N. Y., south to Tex. Pollen: Apparently an
oligolege of Compositae, visits flowers of Bidens, Echinacea pallida, Gossypium
herbaceum, Helianthus annuus, H. maximillianus, H. tuberosus, Physostegia
parviflora, Rudbeckia laciniata, Sonchus terrestris.
Melissodes bidentis Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 362. 9.
bimatris LaBerge. B. C. to Calif., east to Colo. and N. Mex. Pollen: Probably oligolectic,
obtaining pollen principally from flowers of Chrysothamnus including C. nauseosus, C.
n. consimilis, C. n. gnaphalodes, C. n. hololeucus, C. n. mohavensis, C. n. occidentalis,
C. n. speciosus, C. paniculatus, C. parryi, C. viridulus, C. viscidiflorus viscidiflorus, and
Haplopappus acradenius, H. arborescens H. palmeri, but also visits flowers of
Artemisia, Aster, Bigelovia, Centromadia pungens, Chaematoris, Eriogonum, Grindelia
camporum, Gutierrezia californica, G. lucida, G. sarothrae, Helianthus, Rhamnus
californica, Senecio.
Melissodes (Eumelissodes) bimatris LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 431. 2, 6.
boltoniae Robertson. Pa. to Minn. south to Fla. and Tex. Pollen: Oligolege of Compositae,
visiting flowers of Abutilon avicennae, Amphiachyris dracunculoides, Aster ericoides
villosus, A. multiflorus, A. novaeangliae, A. praeatus, Bidens involucrata, Boltonia
asteroides, Cassia, Chrysopsis, Cirsium, Coreopsis aristosa, Helenium altissimum, H.
autumnale, H. tenuifolium, Helianthus annuus, H. grosse-serratus, H. tuberosus,
Heterotheca latifolia, Lythrum, Ratibida pinnata, Rudbeckia triloba, Silphium, Solidago
canadensis, S. missouriensis, Verbena stricta, Vernonia baldwini interior. In the
vicinity of Lawrence, Kansas females have been observed almost invariably collecting
pollen from flowers of Amphiachyris dracunculoides.
Melissodes boltoniae Robertson, 1905. Amer. Ent. Soc., Trans. 31: 368. °.
Melissodes melandri Cockerell, 1906. Amer. Ent. Soe., Trans. 32: 109. @.
brevipyga LaBerge. Ariz., Calif., Idaho. Pollen: Oligolege of Compositae, visits flowers of Aster
spinulosus, A. tephrodes, Baccharis emoryi, Chrysothamnus, Gutierrezia californica, G.
sarothrae, Haplopappus acradenius, Palafoxia linearis, Pectis papposa.
Melissodes (Eumelissodes) brevipyga LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 606, figs.
111-112. 2, d.
cerussata LaBerge. Calif. (Blythe, Hopkins Well and Ludlow). Pollen: Unknown, but visits
flowers of Geraea canescens and Hoffmanseggia in the fall.
Melissodes (Eumelissodes) cerussata LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 436. 2.

2146 Hymenoptera in America North of Mexico

Taxonomy: LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 238-239. ¢.
confusa Cresson. Alta., Man. and Sask. south through Calif., Ariz. and N. Mex. to southern

Mexico (Veracruz and Oaxaca). Pollen: Probably oligolectic on a wide range of
Compositae, visits flowers of Agoseris glauca, Aplopappus, Argemone platyceras, Aster
canescens, Bidens triplinervia var. macrantha, Ceanothus, Cirsium undulatum,
Cleome, Cosmos parviflorus, Erigeron uniflorus, Eryngium asperum, Gaillardia,
Geranium atropurpureum, Grindelia squarrosa, Helenium bigelovii, H. hoopesii,
Heliopsis scabra, Lactuca pulchella, Lotus, Medicago sativa, Monarda pectinata,
Penstemon, Petalostemon purpureum, Phacelia, Polymentha, Ratibida columnaris,
Rudbeckia hirta, R. laciniata, Senecio bigelovii, S. purchianus, Solidago trinervata,
Teucrium occidentale, Verbena stricta, Verbesina encelioides, Vicia pulchella.

Melissodes confusa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 205. 2, 3.

Melissodes ruidosensis Cockerell, 1896. Entomologist 29: 305. 3.

Melissodes tenuitarsis Cockerell, 1905. Psyche 12: 99. 3.

Melissodes civica Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 258. 9.

Melissodes atraticornis Cockerell, 1934. Amer. Mus. Novitates 697: 9. 6.

coreopsis Robertson. Alta. east to Minn., south to IIl., Tex. and Ariz. Pollen: Oligolectic,
principally Helianthus and related genera, but visits a wide variety of flowers for pollen
and or nectar including Amphiachyris dracunculoides, Amorpha canescens, A. fruticosa,
Aster ericoides villosus, A. multiflora, A. novaeangliae, A. paniculatus, A. praeatus,
Bidens involucrata, Boltonia asteroides, Chrysopsis angustifolia, Chrysothamnus
graveolus, Cirsium, Clematis, Cleome serrulata, Cooperia pedunculata, Convolvulus,
Coreopsis grandiflorum, C. palmata, C. tinctoria, Cosmos, Echinacea angustifolia, E.
pallida, E. purpurea, Erucastrum pollichii, Eryngium leavenworthii, Eupatorium
altissimum, Euphorbia, Eustoma russellianum, Gaillardia pulchella, Geranium,
Gossypium herbaceum, Grindelia squarrosa, Gutierrezia sarothrae, Haplopappus,
Helenium autumnale, H. laciniatum, H. latifolia, H. nudiflorum, H. tenuifolium,
Heterotheca subaxillaris, Helianthus annuus, H. grosse-serratus, H. maximillianus, H.
petiolaris, H. salicifolius, H. tuberosus, Heliopsis helianthoides, Marrubium vulgare,
Medicago sativa, Melilotus alba, M. officinalis, Monarda citriodora, M. pectinata, M.
punctata, Nepeta cataria, Opuntia lindheimeri, O. macrorhiza, Parosela, Petalostemon
candidum, P. oligophyllum, P. purpureum, Prionopsis ciliata, Psoralea floribunda,
Ratibida columnaris, R. pinnata, Rudbeckia amplexicaulis, R. bicolor, R. hirta, R.
laciniata, R. triloba, Salsola pestifer, Silphium perfoliatum, S. speciosum, Solidago
canadensis, S. rigida, S. serotina, Tetragonotheca ludoviciana, Tetraneuris linearifolia,
Trifolium repens, Verbena officinalis, V. stricta, Verbesina encelioides, Vernonia.
Melissodes coreopsis Robertson, 1905. Amer. Ent. Soc., Trans. 31: 368. 2.
Melissodes agilis semiagilis Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 364. 3.
Melissodes confusiformis Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 366. 9.
Melissodes helianthophila Cockerell, 1914. Ann. and Mag. Nat. Hist. (8) 14: 361. 6.
denticulata Smith. Que. west to N. Dak., south to Fla. and Tex. Pollen: Apparently an
oligolege of Vernonia including V. altissima, V. baldwini, V. b. interior, V. fasciculata,
V. glauca, V. noveboracensis, V. texana, but visits other flowers for nectar including
Amphiachyris dracunculoides, Carduus crispus, Convolvulus sepium, Eupatorium
purpurea, Ipomoea pandurata, Pycnanthemum, Silphium laciniatum, Solidago juncea,
Symphoricarpos, Verbena hastata, V. stricta.
Melissodes denticulata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 311. 6.
Melissodes senilis Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 311. 2.
Melissodes perplexa Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 206. 9, 6.
Melissodes vernoniana Robertson, 1905. Amer. Ent. Soc., Trans. 31: 368. 2, 6.
dentiventris Smith. Southeastern Canada, Mass. west to Kans., south to Ga. and Tex. Pollen:
Oligolectic, principally Aster including A. anomalus, A. dumosus, A. ericoides, A. e.
villosus, A. novaeangliae, A. paniculatus, A. sagitifolius, A. turbinellus, but visits
flowers of other Compositae as well as other families including Bidens aristosa, B.
polylepis, Boltonia asteroides, Coreopsis tripterus, Chrysopsis mariana, C.
microcephala, Eupatorium perfoliatum, E. serotinum, Helianthus annuus, H.
divaricatus, H. grosse-serratus, H. radula, Isopappus divaricatus, Lespedeza virginica,

Superfamily APOIDEA 2147

Lippia lanceolata, Polygonum pennsylvanicum, Solidago canadensis, S. rigida, S.
ulmifolia, Verbena hastata, Vernonia, Veronica.
Melissodes dentiventris Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 312. 6.
Melissodes autumnalis Robertson, 1905. Amer. Ent. Soc., Trans. 31: 369. 2, d.
Melissodes megacerata Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 362. ¢.
elegans LaBerge. Tex., Kans. Pollen: Probably an oligolege of Compositae, visits flowers of
Aplopappus divaricatus, Aster, Boltonia asteroides, Cleome, Croton monanthrogynus,
Eupatorium serotinum, Fagopyrum, Grindelia, Helenium tenuifolium, Heterotheca
latifolia, H. subaxillaris, Polygonum, Prionopsis ciliata, Solidago serotina, Verbesina
encelioides.
Melissodes (Eumelissodes) elegans LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 490. 2, d.
exilis LaBerge. Ariz. (Tucson). Pollen: Unknown, but visits flowers of Aster, Viguiera.
Melissodes (Eumelissodes) exilis LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 648. 2, ¢.
expolita LaBerge. Ariz., south. Calif.; Mexico (Sonora). Pollen: Unknown, but visits flowers of
Baileya, Chrysothamnus, Encelia farinosa, Sphaeralcea emoryi.
Melissodes (Eumelissodes) expolita LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 505. 2, d.
fasciatella LaBerge. Ariz. Pollen: Presumably an oligolege of fall flowering Compositae, visits
flowers of Chrysothamnus, Erigeron, Gutierrezia, Haplopappus gracilis, Heterotheca
subaxillaris, Viguiera.
Melissodes (Ewmelissodes) fasciatella LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 445. °.
fumosa LaBerge. Md. to Nebr. south to S. C., Ala., Miss. Pollen: Presumably an oligolege of
Compositae ahd seems to prefer flowers of the genus Solidago including S. canadensis,
S. glaberrima, S. rupestris, S. serotina, but also visits flowers of Asclepias tuberosa,
Aster dumosus, Boltonia asteroides, Chrysopsis microcephala, Eupatorium
linearifolium, Grindelia, Helenium, Koellia.
Melissodes (Eumelissodes) fumosa LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 486, figs.
90-9 1eeoniGr
gelida LaBerge. Mont. and N. Dak. south to N. Mex. and Tex. Pollen: Oligolege of Compositae
with some preference for flowers of the genus Helianthus including H. annuus, H.
petiolaris, but also visits flowers of Argemone, Chrysopsis, Cosmos, Echinacea,
Engelmannia pinnatifida, Gaillardia, Lacinaria squarrosa, Monarda pectinata,
Ratibida columnaris, Sphaeralcea, Thelesperma gracile, Verbena stricta, Vernonia.
Melissodes (Eumelissodes) gelida LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 575. 2, d.

grindeliae Cockerell. Minn. to Wash., south to southern Calif., Ariz., N. Mex. Pollen: Oligolege
of Compositae, visits flowers of Aster commutatus, Chrysopsis hispida, Chrysothamnus
nauseosus, Cleome serrulata, Croton, Grindelia squarrosa, Haplopappus, Helianthus,
Hymenoxys floribunda, H. richardsonii, Liatris punctata, Lupinus, Melilotus alba, M.
officinalis, Petalostemon occidentale, Phacelia glandulosa, Polymentha, Psilostrophe
gnaphaloides, Ratibida columnaris, R. tagetes, Rudbeckia laciniata, Verbena stricta.
Melissodes grindeliae Cockerell, 1898. Denison Univ. Sci. Labs., Bul. 11: 66. 6, °.
humilior Cockerell. Ariz., N. Mex., west Tex.; Mexico (Chihuahua). Pollen: Presumably an
oligolege of Compositae, visits flowers of Aster crassulus, A. spinosus, A. tanacetifolius,
Grindelia, Haplopappus acradenius, Helenium autumnale, Heterotheca, Lygodesmia
juncea, Solidago occidentalis, Sphaeralcea emoryi, Verbesina encelioides.
Melissodes humilior Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 447, 450. °.
Melissodes intermediella Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 102. 2.
hurdi LaBerge. Calif. (Central Valley). Pollen: Presumably an oligolege of late summer and
fall flowering Compositae including Artemisia, Centromadia pungens, Grindelia
camporum, Gutierrezia californica, Hemizonia pungens, Heterotheca grandiflora,
Lessingia glandulifera, Solidago.
Melissodes (Eumelissodes) hurdi LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 526. 2, d.
hymenoxidis Cockerell. Colo., Wyo., Utah, Mont., Calif. Pollen: Presumably an oligolege of late
summer and fall flowering Compositae including Aster foliaceus, Chrysopsis, Grindelia,
Haplopappus suffruticosus, Hymenoxys lingulaefolia, but also visits Lupinus and an
unspecified mint.
Melissodes hymenoxidis Cockerell, 1906. Amer. Mus. Nat. Hist., Bul. 22: 443. 2.

2148 Hymenoptera in America North of Mexico

Taxonomy: Lanham, 1977. Kans. Ent. Soc., Jour. 50: 312 (holotype).
Melissodes fremontii Cockerell, 1907. Entomologist 40: 268. ¢.
Melissodes kelloggi Cockerell, 1919. Ent. News 30: 293. 6.
illata Lovell and Cockerell. P. E. I. to Alta., south to N. C. in the mts. and IIl. Pollen: Oligolege
of Compositae, especially Solidago and Aster, visits flowers of Aster azuwreus, Cirsium
arvense, Chrysanthemum leucanthemum pinnatifidum, Grindelia, Helianthus
strumosus, Hieracium auriantiacum, Leontodon, Rudbeckia serotina, Solidago
canadensis, S. gigantea leiophylla, S. juncea, Sonchus, Tanacetum vulgare.
Melissodes illata Lovell and Cockerell, 1906. Psyche 13: 110. 2, 6.
limbus LaBerge. Ariz., N. Mex. and Tex., to central Mexico (Jalisco and Hidalgo). Pollen:
Oligolege of Compositae, especially Verbesina, Aplopappus and Baileya, visits flowers
of Aplopappus gracilis, A. spinulosus, A. tenuisectus, Argemone, Aster, Baccharis,
Baileya multiradiata, Chrysothamnus, Encelia farinosa, Eriogonum, Gossypium
herbaceum, Grindelia, Gutierrezia, Hymenothrix wislizenii, Kallstroemia grandiflora,
Marrubium vulgare, Melilotus alba, Pectis papposa, Sphaeralcea, Verbesina encelioides,
V. exauriculata, Wedeliella incarnata.
Melissodes (Eumelissodes) limbus LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 585. 2, d.
lutulenta LaBerge. B. C. to Sask., south to Calif. Ariz. and Mexico (Guanajuato). Pollen:
Oligolege of Compositae, especially Chrysothamnus, Grindelia and Solidago, visits
Achillea millefolia, Anthemis cotulla, Aster spinosus, Centromadia pungens,
Chaemataxis, Chrysothamnus, Cleome, Grindelia squarrosa, Gutierrezia, Helianthus,
Melilotus alba, Mentzelia veatchiana, Solidago occidentalis.
Melissodes (Eumelissodes) lutulenta LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 599. 2, ¢.
manipularis Smith. Fla. to N. C., ?Va. Pollen: Unknown, but visits flowers of Chrysopsis,
Kuhnistera, Petalostemon. The Virginia record pertains to a collection made at the
flowers of Eupatorium in Falls Church and has not been verified since it was published
in 1907 by Cockerell.
Melissodes manipularis Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 312. 6.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 329.
melanura (Cockerell). Calif., Nev. Pollen: Presumably an oligolege of summer and fall
flowering Compositae including Aster, Corethrogyne bernardina, Erigeron, Gutierrezia
californica, G. sarothrae, Haplopappus linearifolius, Lessingia glandulifera, Solidago
elongata, but also visits flowers of Eriogonum virgatum.
Exomalopsis melanurus Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 59. @.
menuachus Cresson. B. C., Alta. and N. Dak., south to Calif., Tex., and III.; Mexico (Chihuahua,
Zacatecas). Parasite: Triepeolus occidentalis Cress.? Pollen: Oligolege of Compositae,
especially Grindelia and Solidago, visits flowers of Argemone intermedia, A. platyceras,
Aster laevis, A. multiflora, Cassia chamaecrista, Chrysopsis, Chrysothamnus graveolus
glabrata, Cleome serrulata, Eustoma russellianum, Gaillardia, Grindelia inornata, G.
squarrosa, Gutierrezia californicum, G. sarothrae, Helianthus annuus, H. petiolaris,
Hymenothrix wislizenia, Medicago sativa, Melilotus alba, Petalostemon oligocephalum,
Polygonum, Rudbeckia laciniata, Sidalcea neomexicana, Solidago canadensis, S. rigida,
Verbena, Viguiera, Xanthocephalum gymnospermoides.
Melissodes Menuachus Cresson, 1868. Amer. Ent. Soc., Trans. 1: 388. ¢.
Melissodes mennacus(!) Uhler, 1877. U. S. Geol. Geog. Survey, Bul. 3: 783.
Melissodes pallida Robertson, 1895. Amer. Ent. Soc., Trans. 22: 127. °.
Melissodes Mizeae Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 522. °.
Melissodes blakei Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 523. 9.
Melissodes lavata Cockerell, 1924. Pan-Pacific Ent. 1: 56. 2.
Melissodes octobris Cockerell, 1934. Ent. News 45: 30. 2.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 225 (nest site).
micheneri LaBerge. Calif. (Sierra Nevada Mts. and Peninsular Mts. of south. Calif.). Pollen:
Oligolege of Compositae, especially Chrysothamnus and Chrysopsis, visits flowers of
Bigelovia, Chrysopsis fastigiata, C. villosa, Chrysothamnus nauseosus, Haplopappus
arborescens, H. bloomeri angustatus, Solidago.
Melissodes (Eumelissodes) micheneri LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 642. 2, ¢.

Superfamily APOIDEA = 2149

microsticta Cockerell. B. C. to Sask. south to southern Calif., Nev., Utah., Colo., and Tex.
Pollen: Oligolege of Compositae, visits flowers of Artemisia, Aster adscendens, A.
canescens, A. delectabilis, A. foliaceus, Calyptridium umbellatum, Carduus, Cichorium
intybus, Chrysopsis fastigiata, C. villosus, Chrysothamnus nauseosus, C. n. consimilis,
Corethrogyne, Cryptantha intermedia, Erigeron stenophyllus, Eriogonum subscaposum,
Grindelia squarrosa, Gutierrezia californica, G. sarothrae, Haplopappus, Helenium
bigelovii, Helianthus, Hemizonia wheeleri, H. wrightii, Malvastrum paniculata,
Penstemon, Phacelia humilis, Senecio ionophyllus, Solidago elongata, S. occidentalis,
Sphaeralcea fasciculata.
Melissodes microsticta Cockerell, 1905. In Viereck, Canad. Ent. 37: 321. 6.
monoensis LaBerge. Calif. (Mono Co.). Pollen: Unknown, but visits flowers of Chrysothamnus.
Melissodes (Eumelissodes) monoensis LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 623. 2, 6.
montana Cresson. Colo. and Utah, south through Tex., N. Mex., and Ariz. to south. Mexico
(Oaxaca). Pollen: Oligolege of Compositae, visits flowers of Aplopappus gracilis,
Asclepias, Aster commutatus, Croton, Erigeron, Eriogonum, Grindelia, Gutierrezia,
Helianthus annuus, Heterotheca psammophila, H. subaxillaris, Polymentha, Senecio
longilobus, Verbesina exauriculata, but also is known to collect pollen from flowers of
Kallstroemia grandiflora.
Melissodes montana Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 202. 2, ¢.
Melissodes hirsuta Smith, 1879. Descr. n. spp. Hym. Brit. Mus., p. 116. d.
Eucera oajacana Dalla Torre, 1896. Cat. Hym. v. 10, p. 242. N. name for hirsuta Smith, a
secondary homonym in Eucera.
Melissodes atrifera Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 256. d.
Melissodes atrifera sandiarum Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 257. 6.

Biology: Cazier and Linsley, 1974. Amer. Mus. Novitates 2546: 14 (floral relationships).
moorei Cockerell. Calif. Pollen: Oligolege of Compositae, visits flowers of Corethrogyne,
Erigeron glaucus, Grindelia platyphylla, Hemizonia paniculata, Heterotheca
grandiflora, Solidago.
Melissodes moorei Cockerell, 1926. Ann. and Mag. Nat. Hist. (9) 18: 624. 2.
nivea Robertson. N. Y. to Minn. and Kans., south to N. C., Ala., and Miss. Pollen: Oligolege of
Compositae, especially Solidago, Aster and Liatris, visits flowers of Aster ericoides, A.
sagittifolius, Bidens laevis, Boltonia asterioides, Chrysopsis mariana, Gerardia,
Helenium, Helianthus annuus, H. atrorubens, Lacinaria, Lepachys pinnata, Liatris
graminifolia, Prionopsis ciliata, Solidago canadensis, S. lanceolata, S. rigida, S.
serotina, Vernonia glauca.
Melissodes nivea Robertson, 1895. Amer. Ent. Soc., Trans. 22: 127. °.
ochraea LaBerge. South. Calif., Ariz. Pollen: Presumably an oligolege of Compositae, visits
flowers of Baccharis, Chrysothamnus, Gutierrezia, Haplopappus acradenius.
Melissodes (Eumelissodes) ochraea LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 428. 9, d.
pallidisignata Cockerell. B. C. to Man., south through south. Calif. and N. Mex. to central
Mexico (Aguascalientes). Pollen: Oligolege of Compositae especially Haplopappus,
Chrysothamnus, Layia, Erigeron and Grindelia, visits flowers of A plopappus gracilis,
Aster adscendens delectabilis, Centromadia pungens, Chrysopsis hispida,
Chrysothamnus nauseosus, C. n. mojavensis, C. n. occidentalis, C. viscidiflorus typicus,
Cirsium, Cleome serrulata, Ericameria palmeri, Grindelia camporum, G. latifolia, G.
platyhylla, G. squarrosa, Gutierrezia californica, G. sarothrae, Haplopappus acradenius,
H. vernonioides, Helianthus, Heliotropium, Hemizonia pungens, Heterotheca
grandiflora, Lepachys, Lessingia glandulifera, Medicago sativa, Melilotus alba,
Petalostemon occidentalis, Pluchea persica, Psilostrophe gnaphalodes, Senecio
ionophyllus, Solidago canadensis, S. confinis, S. occidentalis, Wislizenia refracta.
Melissodes nigrosignata pallidisignata Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 180. 2.
Melissodes menuacha vernonensis Viereck, 1905. Canad. Ent. 37: 320. 2, d.
Melissodes praelauta Cockerell, 1905. Psyche 12: 102. 6.

Biology: Thorp and Chemsak, 1964. Pan-Pacific Ent. 40: 75-83, 2 figs. (life history).

2150 Hymenoptera in America North of Mexico

paucipuncta LaBerge. Ariz. Pollen: Possibly oligolectic on Opuntia, a male has been taken at
flowers of Bebbia.
Melissodes (Eumelissodes) paucipuncta LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 650. ?, d.
paulula LaBerge. Wash. to southern Calif. Pollen: Oligolege of Compositae, especially
Haplopappus, Gutierrezia, and Solidago, visits flowers of Baccharis emoryi, Croton
californicus, Ericameria palmeri, Erigeron canadensis, Eriogonum fasciculatum, E.
gracile, Gutierrezia californica, G. sarothrae, Haplopappus acradenius, H. arborescens,
H. vernonioides, Pluchea camphorata, Salsola kali, Senecio douglasii, Solidago
californica, S. elongata, S. occidentalis.
Melissodes (Eumelissodes) paulula LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 631. 2, 6.
perlusa Cockerell. Alta. and Man., south to Iowa and Ariz. Pollen: Possibly an oligolege of
Helianthus including H. annuus, H. petiolaris, but also visits flowers of Bigelovia,
Brauneria pallida, Grindelia, Medicago sativa, Mentha canadensis, Petalostemon
oligophyllum, Ratibida columnaris.
Melissodes semiagilis perlusa Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 231. 3.

Taxonomy: Cockerell, 1914. Canad. Ent. 46: 413. °.
perpolita LaBerge. Ariz., N. Mex., Colo., (?)Utah. Pollen: Unknown, but visits flowers of Aster,
Chaematazxis, Grindelia, Haplopappus spinulosus.

Melissodes (Eumelissodes) perpolita LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 443. 2, 6.
personatella Cockerell. Central and southern Calif.; Guatemala (Jicaro). Pollen: Unknown, but
visits flowers of Eriogonum gracile, E. subscaposum, Gutierrezia californica, G.

sarothrae, Hagardia squamosa, Haplopappus palmeri, H. vernonioides, Hemizonia
paniculata, Heterotheca grandiflora, Palafoxia linearis.
Melissodes personatella Cockerell, 1901. Canad. Ent. 33: 297. 6.
pexa LaBerge. Ariz.; Mexico (Coahuila, Chihuahua). Pollen: Unknown, but visits flowers of
Lepidium alyssoides.
Melissodes (Eumelissodes) pexa LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 621. 2, d.
pilleata LaBerge. N. C. Pollen: Possibly an oligolege of Compositae, visits flowers of Aster,
Chrysopsis, Gerardia flavia, Haplopappus, Kuhnistera pinnata, Liatris.
Melissodes (Eumelissodes) pilleata LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 459. 2, d.
pullatella LaBerge. Oreg. Pollen: Unknown, but visits flowers of Grindelia.
Melissodes (Eumelissodes) pullatella LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 620. 9.
relucens LaBerge. N. Mex. (Mesilla Park), Tex. (E] Paso and Big Bend Natl. Pk.). Pollen:
Unknown, but visits flowers of Chrysopsis hirsutissima, Haplopappus heterophylla.
Melissodes (Eumelissodes) relucens LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 437. 2, ¢.
robustior Cockerell. Wash., Oreg., Idaho, Nev., Calif. Parasite: Nemognatha hurdi MacSwain,
Sphaeropthalma unicolor (Cress.). Pollen: Oligolege of Compositae, especially
Helianthus including H. annuus, H. bolander, H. gracilentus, H. lenticularis, H.
petiolaris, but also visits flowers of Asclepias, Aster, Brassica adpressa, B. geniculata,
B. incana, Chaenactis artemisifoliae, C. glabriuscula, Chrysothamnus, Cichorium,
Cirsium lanceolatum, Coreopsis grandiflora, C. lanceolata, C. tinctoria, Corethrogyne
bernardina, Cosmos, Encelia californica, E. farinosa, Gaillardia, Godetia bottae,
Grindelia camporum, G. elata, Gutierrezia californica, G. sarothrae, Haplopappus
vernonioides, Hemizonia paniculata, H. pungens, H. wrightii, Heterotheca grandiflora,
Latuca serriola, Lythrum californicum, Marrubium vulgare, Scabiosa atropurpurea,
Senecio douglasii, Solidago californica, S. confinis, Stephanomeria exigua.
Melissodes robustior Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 16: 482. 2, 6.

Biology: MacSwain, 1958. Pan-Pacific Ent. 34: 40. —Chemsak and Thorp, 1962. Pan-Pacific
Ent. 38: 53-55 (nest).

rustica (Say). N. S. to Sask., south to Ga., La., N. Mex. and southern Mexico (Oaxaca), but not
known in Tex. and Okla. Parasite: Triepeolus eldredi Ckll., T. pectoralis (Robt.). Pollen:
Oligolege of Compositae, especially Solidago, Aster, Grindelia, Aplopappus, analyzed
cell provisions of nests in Wyo. showed that females obtained pollen exclusively from
Eriophyllum integrifolium; visits flowers of Abutilon theophrasti, Amphiachyris
dracunuloides, Aplopappus gracilis, Asclepias, Aster anomalus, A. commutatus, A.

Superfamily APOIDEA 2151

crenulis, A. dumosus, A. ericoides, A. e. villosus, A. exiguus, A. grandiflorus, A. laevis,
A. lateriflorus, A. multiflorus, A. novaeangliae, A. paniculatus, A. praeustus, A.
sagittifolius, A. salicifolius, A. turbinellus, Baccharis, Bidens aristosa, B. laevis, B.
asteroides, Centaurea juncea, Chrysopsis, C. mariana, Cirsium arvense, C.
lanceolatum, Cleome lutea, C. serrulata, Coreopsis tripteris, Cosmos, Epilobiwm
perfoliatum, Gaillardia, Grindelia squarrosa, Helenium autumnale, H. tenuifolium,
Helianthus atrorubens, H. divaricatus, H. grosse-serratus, H. maximillianus, H.
petiolaris, H. radula, H. tuberosus, Heliopsis helianthoides, Hieracium scabrum,
Lycopus americanus, Marrubium vulgare, Melilotus alba, M. officinalis, Mentha,
Physostegia parviflora, Polymentha, Ratibida columnaris, R. pinnata, Rudbeckia
laciniata, R. subtomentosa, R. triloba, Silphium perfoliatum, Solidago altissima, S.
canadensis, S. graminifolia, S. juncea, S. nemoralis, S. rigida, S. rugosa, S. serotina,
Spiraea alba, Verbena hastata, V. stricta, Verbesina virginica, Vernonia fasciculata, V.
glauca.

Macrocera rustica Say, 1837. Boston Jour. Nat. Hist. 1: 406. 6, 9.

Melissodes assimilis Smith, 1879. Deser. n. spp. Hym. Brit. Mus., p. 114. 9.

Melissodes ambigua Smith, 1879. Descr. n. spp. Hym. Brit. Mus., p. 116. @.

Melissodes festinata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym. v. 2, p. 300.
Or

Melissodes simillima Robertson, 1897. Acad. Sci. St.-Louis, Trans. 7: 355. 2, 6.

Melissodes asteris Robertson, 1914. Ent. News 25: 373. 2.

Taxonomy: Clement, 1973. Kans. Ent. Soc., Jour. 46: 521, figs. 3-6 (larva).

Biology: Clement, 1973. Kans. Ent. Soc., Jour. 46: 516-525, 6 figs., 1 table (nest architecture,
pollen stores, parasite). :
saponellus Cockerell. Wash., Oreg., Calif., Utah, Colo. Pollen: Unknown, but visits flowers of
Bahia oblongifolia, Chaenactis stevioides, Encelia actoni, Sphaeralcea, Trifolium.
Melissodes saponellus Cockerell, 1908. Canad. Ent. 40: 234. 9.

Biology: LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 241 (interspecific copulation).

semilupina Cockerell. B. C. south to Calif., east to Colo. and N. Mex. Pollen: Oligolege of
Compositae, especially Chrysothamnus and Haplopappus, visits flowers of
Chrysothamnus nauseosus, C. n. consimilis, C. viscidiflorus typicus, Cleome,
Gutierrezia californica, Haplopappus vernonioides, Hemizonia paniculata, Solidago
occidentalis.
Melissodes menuacha semilupina Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 29. ¢.
Melissodes chrysothamni Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 524. °.
snowii Cresson. Alta., Man., Colo., N. Mex., Nebr., N. Dak. Pollen: Probably an oligolege of
Compositae, especially Helianthus and Solidago, visits flowers of Aster, Cleome
serrulata, Gaillardia, Gutierrezia sarothrae, Helianthus petiolaris, H. subrhomboideus,
Lacinaria punctata, Medicago sativa, Solidago missouriensis, S. nemoralis, S. rigida.
Melissodes snowii Cresson, 1872. Acad. Nat. Sci. Phila., Proc. 24: 211. ¢. a
subagilis Cockerell. Minn., Alta. and Wash. south to northern Calif., Ariz. and Tex. Pollen:
Oligolege of Grindelia including G. nana, G. squarrosa, but visits other flowers
including Aplopappus gracilis, A. spinulosus, A. tenuisectus, Aster adscendens,
Baccharis, Baileya multiradiata, Bigelovia wrightii, Chrysopsis hispida,
Chrysothamnus viscidiflorus, Cleome serrulata, Erigeron, Gossypium herbaceum,
Gutierrezia sarothrae, Haplopappus acradenius, Helianthus annuus, H. maximillianus,
H. petiolaris, Heterotheca subaxillaris, Medicago sativa, Pectis papposa, Petalostemon,
Prionopsis ciliata, Ratibida tagetes, Salsola pestifer, Silphiwm, Solidago rigida, S.
serotina, Sphaeralcea, Verbena hastata, Verbesina exauriculata.
Melissodes agilis var. subagilis Cockerell, 1905. Entomologist 38: 145. d.
Melissodes pecosella Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 179. °.
Tetraloniella excurrens melanaspis Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 228.

oF
Xenoglossodes albertensis Cockerell, 1937. Canad. Ent. 69: 87. °.
Biology: Hicks, 1926. Colo. Univ., Studies 15: 225 (nest).

2152 Hymenoptera in America North of Mexico

subillata LaBerge. Que. to Sask., south to N. C. and Ill. Pollen: Oligolege of Compositae, visits
flowers of Aster azureus, Chrysanthemum leucanthemum, Cichorium intybus, Cirsium
arvense, Coreopsis grandiflora, C. lanceolata, Echinacea pallida, Epilobium
angustifolium, Eupatorium maculatum, Gaillardia aristata, Helianthus
maximillianus, H. petiolaris, Hieraceum auriantiacum, Latuca pulchella, Petalostemon
oligophyllum, Platycodon grandiflorium, Psoralea lanceolata, Ratibida columnaris,
Rudbeckia hirta, R. laciniata, Solidago graminifolium, Sonchus arvensis glabrescens,
Teucrium occidentalis, Vernonia fasciculata.
Melissodes (Eumelissodes) subillata LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 568, figs.
99-101. 2, d.
submenuacha Cockerell. Colo., Ariz., N. Mex., western Tex. Pollen: Probably an oligolege of
Compositae, visits flowers of Aplopappus gracilis, Bidens, Cleome serrulata,
Haplopappus heterophylla, Helianthus annuus, H. petiolaris, Hymenothrix wislizeni,
Medicago sativa, Verbesina encelioides, V. oreophila.
Melissodes menuacha var. submenuacha Cockerell, 1897. Entomologist 30: 137. 3.
Melissodes Hewetti Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 527. °.

Taxonomy: LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 241-242 (geogr. and floral
records).
tincta LaBerge. Minn. and Mich. south to Tex. and Fla. Pollen: Oligolege of Compositae, seems
to prefer Chrysopsis and Aster, visits flowers of Aster longicaulus, Chrysopsis
microcephala, Helianthus maximillianus, Verbesina encelioides.
Melissodes (Eumelissodes) tincta LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 493. 2, 6.

trinodis Robertson. Southeast. Canada, Kans. and N. Dak. east to Maine and Ga. Parasite:
Triepeolus nelianthi (Robt.). Pollen: Oligolege of Compositae, especially Helianthus
including H. annuus, H. a. coronarius, H. atrorubens, H. divaricatus, H. grosse-serratus,
H. maximillianus, H. mollis, H. salicifolius, H. tuberosus, but also visits flowers of
Arctium, Asclepias incarnata, Aster anomalus, A. praeatus, Bidens aristosa, B. laevis,
Blephilia hirsuta, Carduus crispus, Cassia chamaecrista, Cirsium lanceolata, Coreopsis
palmata, C. tripteris, Dichophyllum marginatum, Grindelia, Helenium altissimum, H.
autumnale, Heliopsis helianthoides, Lepachys pinnata, Liatris, Monarda fistulosa,
Pepo, Petalostemon purpureum, Ratibida columnaris, Rudbeckia hirta, R. laciniata, R.
submentosa, R. triloba, Solidago canadensis, S. rupestris, S. ulmifolia, Symphoricarpos,
Teucrium canadense, Verbena hastata, V. stricta, Vernonia glauca, V. baldwini interior,
Veronica.

Melissodes trinodis Robertson, 1901. Canad. Ent. 33: 231. 2, d.

Biology: Graenicher, 1905. Wis. Nat. Hist. Soc., Bul. 3: 164 (nest).

tristis Cockerell. Nebr. to Calif., south through Tex. and Ariz. to southern Mexico (Puebla).
Pollen: Most polylectic of all the species of the subgenus Hwmelissodes, seems to prefer
flowers of the Leguminosae, Compositae and Malvaceae, visits flowers of Acacia,
Actinea acaulis, A. richardsonii, Allionia incarnata, Arabis, Argemone, Asclepias
galloides, A. verticillata, Aster canescens, A. spinosa, A. tanacetifolium, Astrgalus,
Baccharis, Bahia, Baileya multiradiata, Berberis trifoliata, B. wilcoxii, Canotia
holocantha, Ceanothus fendleri, C. greggii, Celtis pallida, Cercidium texanum, Cevalia
sinuata, Chilopsis linearis, C. saligua, Chrysopsis hispida, Chrysothamnus, Cirsium
ochrocentrum, C. undulatum, Cleome luteum, C. serrulata, Convolvulus, Croton
luteovirens, Dalea lasianthera, Dasylirion wheeleri, Encelia, Engelmannia pinnatifida,
Erigeron canadensis, Eriogonum trichopes, Euphorbia albomarginata, Eysenhardtia
polystacha, E. spinosa, Gaillardia pinnatifida, G. pulchella, G. suavis, Gaura, Gilia
calcarea, Gossypium herbaceum, Grindelia, Gutierrezia californica, Haplopappus
gracilis, H. laricifolius, Heleniuwm autumnale, H. hoopesii, H. laciniatum, Helianthus
annuus, H. ciliaris, H. petiolaris, Hoffmanseggia densiflora, H. jamesti, Hymenothrix
wislizenii, Iris mios, Kallstroemia grandiflora, Larrea tridentata, Lepidiwm,
Lesquerella ovalifolia, Lippia cuneifolia, L. ligustrina, Lotus, Lupinus, Lygodesmia
juncea, Malvastrum cockerelli, Marrubium vulgare, Medicago sativa, Melilotus alba, M.
officinalis, Mentzelia multiflora, Mertensia franciscana, Mimosa borealis, Monarda
citriodora, M. pectinata, Mortonia scabrella, Nepeta cataria, Nolina microcarpa,

Superfamily APOIDEA 2153

Opuntia basilaris, O. lindheimeri, O. macrorhiza, Parkinsonia, Pectis papposa,
Penstemon superba, Petalostemon candidum, P. flavescens, P. occidentatum, Phacelia
glandulosa, P. popei, Philadelphus microphyllus, Polemonium, Prionopsis, Prosopis
juliflora, Psoralea tenuiflora, Pyracantha, Ratibida columnaris, R. tagetes, Rhus, Rosa,
Salix, Salsola kali, S. pestifer, Salvia lemmoni, Sapindus drummondii, S. sapinarius,
Senecio longilobus, S. werneriaefolius, Solanum elaeagnifolium, Solidago annua, S.
stenolobus, Sphaeralcea angustifolia, S. coccinea, S. emoryi, S. laxa, S. lobata, S.
marginata, Stephanomeria, Tamarix gallica, Thurberia thespesioides, Verbena stricta,
Verbesina encelioides, Vernonia, Vicia, Wislizenia refracta, Zexmenia podocephala,
Zinnia grandiflora.

Melissodes tristis Cockerell, 1894. Ent. News 5: 234. 6.

Melissodes pallidicincta Cockerell, 1896. Entomologist 29: 306. °.

Melissodes tristis var. malvina Cockerell, 1902. Entomologist 35: 177. d.

m elissodes semitristis Cockerell, 1905. Psyche 12: 102. ¢.

Melissodes pallidicincta erythrina Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 231. d.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 161, fig. 8 (sleep). —Hurd and Linsley, 1975.
Smithsn. Contrib. Zool. 193: 41-42, fig. 17 (intrafloral relationships with Larrea).
utahensis LaBerge. Utah and Nev., south to south. Calif. and Ariz., Nebr. Pollen: Unknown,
but visits flowers of Chrysothamnus including C. nauseosus, Chaemataxis, Eriogonum
heermannii, Gutierrezia californica.
Melissodes (Eumelissodes) utahensis LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 602, figs.
109-110. 9, 6.
velutina (Cockerell). South. and cent. Calif. Pollen: Polylege, especially of Polemoniaceae and
Compositae, visits flowers of Acanthopappus sphaerocephalus, Aster abatus,
Chorizanthe staticoides, Cirsium, Coreopsis lanceolata, Cryptantha intermedia, Encelia
actoni, Eremocarpus setigerus, Eriogonum fasciculatum, Gutierrezia californica, G.
sarothrae, G. stricta, Helianthus gracilentus, Heliotropium curassavicum var. oculatum,
Hugelia ambigua, H. virgata, Lotus scoparius, Marrubium vulgare, Mirabilis laevis,
Monarda lanceolata, Prosopis, Salvia columbariae, Trichostema lanatum.
Exomalopsis velutinus Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 58. 2.
verbesinarum Cockerell. Wash. to southern Calif., east to Nev. and Tex.; Mexico (Chihuahua to
Jalisco). Pollen: Oligolege of Compositae, visits flowers of Actinea, Argemone
platyceras, Aster canescens, A. tanacetifolius, Baileya multiradiata, Bigelovia, Cevallia
sinuata, Chaenactis, Chrysothamnus, Croton californicus, Encelia californica,
Erigeron, Eysenhardtia polystachya, Geraea canescens, Grindelia, Gutierrezia lucida,
Haplopappus acradenius, Lygodesmia juncea, Pectis papposa, Psilostrophe cooperi,
Sphaeralcea, Tamarix, Verbesina encelioides.
Melissodes pecosella verbesinarum Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 180. °.
vernalis LaBerge. Wash. and Nev. to southern Calif. and Ariz.; Mexico (Baja California,
Sonora, and Hidalgo). Pollen: Oligolege of Compositae, especially Encelia and Geraea,
visits flowers of Aster abatus, A. agnatus, Baileya multiradiata, Bebbia juncea,
Chilopsis linearis, Encelia farinosa, Geraea canescens, Hyptis emoryi, Melilotus,
Prosopis, Psilostrophe cooperi, Sphaeralcea ambigua, Viguiera deltoides.
Melissodes (Eumelissodes) vernalis LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 608, figs.
113-114. 2, 6.
vernoniae Robertson. Ind., and N. Dak., west to Idaho, south to N. Mex. and Tex. Pollen:
Polylectic, collects pollen of Vernonia, also probably uses pollen from Helianthus and
possibly Ipomoea, visits flowers of Bidens, Gaillardia pulchella, Grindelia, Helianthus
tuberosus, Heliopsis helianthoides, Ipomoea pandurata, Lacinaria, Lactuca pulchella,
Liatris, Monarda fistulosa, Polygonum, Rudbeckia hirta, Silphium laciniatum, S.
speciosum, Verbena stricta, Vernonia altissima, V. fasciculata, V. baldwini interior, V.
longifolia, V. texana.
Melissodes vernoniae Robertson, 1902. Canad. Ent. 34: 323. 2, d.
Melissodes cénfusiformis incondita Cockerell, 1925. Ann. and Mag. Nat. Hist. (9) 16: 230.
OF

2154 Hymenoptera in America North of Mexico

wheeleri Cockerell. Mich. and N. Dak., south to La., Tex. and Ariz. Pollen: Oligolege of
Compositae, especially Gaillardia, Helianthus and Rudbeckia, visits flowers of
Anthemis cotula, Asclepias tuberosa, Cleome serrulata, Coreopsis, Echinacea,
Engelmannia bipinnatifida, Gaillardia pulchella, Grindelia squarrosa, Helianthus
annuus, H. debilis, H. petiolaris, Opuntia, Petalostemon purpureum, Ratibida
columnaris, Rudbeckia bicolor, R. grandiflora, R. hirta, Silphium asperimum,
Sphaeralcea.
Melissodes wheeleri Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 111. 9°.

Taxonomy: Cockerell, 1906. Ann. and Mag. Nat. Hist. (7) 17: 367. 6.

Genus MELISSODES Subgenus PSILOMELISSODES LaBerge

Melissodes subg. Psilomelissodes LaBerge, 1956. Kans. Univ. Sci. Bul. 37: 1173.
Type-species: Melissodes intorta Cresson. Monotypic and orig. desig.
intorta Cresson. Kans., Tex. Pollen: Probably dependent upon the pollen of the Malvaceous
genus Callirhoe including C. digitata, C. involucrata, C. leiocarpa, but also visits
flowers of Asclepias latifolia, Gaillardia, Rorippa sinuata.
Melissodes intorta Cresson, 1872. Amer. Ent. Soc., Trans. 4: 278. 3.
Melissodes wickhami Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 112. °.

Taxonomy: LaBerge, 1963. Nebr. Univ. State Mus., Bul. 4: 234 (geogr. and floral records).

Genus MELISSODES Subgenus CALLIMELISSODES LaBerge

Melissodes subg. Callimelissodes LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 294.
Type-species: Melissodes lupina Cresson. Orig. desig.
ablusa Cockerell. Calif. (Millbrae, Bolinas).
Melissodes metenua ablusa Cockerell, 1926. Pan-Pacifie Ent. 3: 85. 2.
clarkiae LaBerge. Oreg., Calif. Pollen: Probably oligolectic on Clarkia including C. amoena, C.
biloba, C. cylindrica, C. dudleyana, C. gracilis albicaulis, C. speciosa, C. unguiculata, C.
williamsonii.
Melissodes (Callimelissodes) clarkiae LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 315, figs.
50-538. 2, d.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 29-31 (floral
relationships).
coloradensis Cresson. N. C., Ohio, Ind., Ill., Wis., Mo., Nebr., Kans., Wyo., Colo., N. Mex., Ariz.,
Utah, north. Calif. Pollen: Oligolege of Compositae, with Helianthus including H.
annuus, H. atrorubens, H. divaricatus, H. grosse-serratus, H. mollis, H. petiolaris, H.
scaberrimus and H. tuberosus as the primary source of pollen, but also visits flowers of
Cirsium discolor, C. lanceolatum, Coreopsis tripteris, Heliopsis helianthoides, H. laevis,
Liatris pycnostachya, Lythrum alatum, Rudbeckia laciniata, Silphium integrifolium, S.
laciniatum, S. perfoliatum, S. terebinthinaceum, Solanum, Solidago serotina, Verbena
hastata, V. stricta, Vernonia baldwini interior, V. fasciculatum.
Melissodes coloradensis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 200. 2, d.
composita Tucker. Mont. south to Ariz., N. Mex. Ecology: Nests in ground individually, but
many individuals use a communal entrance. Parasite: Triepeolus helianthi Robt.? Pollen:
Unknown, but visits flowers of Grindelia squarrosa, Haplopappus gracilis, Thurberia
thespesioides.
Melissodes lupina composita Tucker, 1909. Kans. Acad. Sci., Trans. 22: 281. 6.

Biology: Hurd and Linsley, 1959. Ent. News 70: 141-146 (nest).
glenwoodensis Cockerell. N. Dak. south to N. Mex., west to Wash. and southern Calif. Pollen:
Oligolege of Compositae, especially Chrysothamnus including C. nauseosus consimilis,
C. n. mojavensis, C. n. speciosus, C. viscidiflorus typicus, but visits other flowers
including Cirsium, Grindelia squarrosa, Haplopappus vernonioides, Helianthus
petiolaris, Solidago trinervata.
Melissodes glenwoodensis Cockerell, 1905. Ann. and Mag. Nat. Hist. (7) 15: 522. 2.

Taxonomy: Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 107.

Superfamily APOIDEA 2155

lupina Cresson. Mont. to Wash., south to Colo., Utah, Nev., Calif.; Mexico (Baja Calif.). Pollen:
Oligolege of Compositae, but visits a wide variety of flowers including Achillea,
Adenostegia, Adenostoma fasciculatum, Arctium, Asclepias, Aster chilensis, A. exilis,
Bigelovia, Brassica, Calochortus nuttalli, Calycadenia, Castanopsis, Centaurea
solstitialis, Chaenactis artemisiaefolia, C. glabriuscula, Chrysopsis villosus,
Chrysothamnus nauseosus speciosus, C. viscidiflorus typicus, Cirsium, Clarkia
amoena, C. bottae, Cleome serrulata, Cordylanthus, Coreopsis lanceolatum, C. tinctoris,
Corethrogyne bernardense, C. filaginifolia, Cosmos, Croton californicus, Cryptantha
intermedia, Cucurbita, Encelia, Epilobium, Eremocarpus setigerus, Erigeron foliosus,
Eriodictyon angustifolium, Eriogonum fasciculatum, E. gracile, E. latifolium,
Eriophyllum confertiflorum, Grindelia camporum, G. elata, G. squarrosa, Gutierrezia
californica, G. sarothrae, Haplopappus arborescens, H. bloomeri angustatus, H. palmeri,
Hemizonia fasciculata, H. fitchi, H. heermannii, H. lobbii, H. luzulaefolia, H.
paniculata, H. pungens, H. wrightii, Heterotheca grandiflora, Holodiscus discolor, Iris
hartwegi, Lessingia, Ligustrum, Limnanthes, Lotus, Lythrum californica, Madia,
Malacothrix, Marrubium vulgare, Medicago sativa, Melilotus alba, M. indica,
Monardella, Nemophila, Perezia microcephala, Phacelia heterophylla, P. humilis,
Phalacroseris bolanderi, Pimpinella, Polygonum auberti, Raphanus, Rhamnus
californica, Salvia carduacea, Senecio, Solidago californica, S. elongata, S. occidentalis,
Stephanomeria exigua, S. virgata, Trichostema laxum, Trifolium repens, Verbena,
Vicia.
Melissodes lupina Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 210. d.
Melissodes intermediella var. catalinensis Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4:
10252.
Melissodes catalinensis vanduzeei Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 86. 2.
lustra LaBerge. Idaho, Oreg., Nev., Calif.; Mexico (Baja Calif.). Pollen: Oligolectic, depending
upon plants of Compositae for pollen, particularly Gutierrezia, Chrysothamnus and
Haplopappus, visits flowers of Aster, Baccharis pilularis, Bigelovia, Chrysothamnus
nauseosus nauseosus, C. n. consimilis, C. n. speciosus, C. viridulus, C. viscidiflorus
typicus, Cleome, Croton californicus, Eriogonum latifolium, Grindelia camporum, G.
sarothrae, Haplopappus bloomeri angustatus, H. palmeri, H. vernonioides, Heterotheca
grandiflora, Melilotus alba, Senecio douglasii.
Melissodes (Callimelissodes) lustra LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 334, fig. 59. 2,
3.
metenua Cockerell. Wash. to northern Calif., Idaho, Wyo., Utah. Pollen: Unknown, but visits
flowers of Cosmos, Eriogonum, Melilotus alba.
Melissodes metenua Cockerell, 1924. Pan-Pacific Ent. 1: 56. °.
minuscula LaBerge. Calif. Pollen: Unknown, but visits flowers of Cressa cretica,
Heliotropium curassavicum, Melilotus.
Melissodes (Callimelissodes) minuscula LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 323. 9,
3.
nigracauda LaBerge. Calif. Pollen: Unknown, but visits flowers of Erigeron, Senecio
douglasii, Stephanomeria exigua.
Melissodes (Callimelissodes) nigracauda LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 327, figs.
54-57. 2, d.
plumosa LaBerge. Wash. to Calif., (?)N. Dak. Pollen: Unknown, but visits flowers of Encelia
californica, Helianthus petiolaris.
Melissodes (Callimelissodes) plumosa LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 308, figs.
42-45. 2, 3.
stearnsi Cockerell. Wash., Oreg., Calif. Pollen: Possibly an oligolege of Compositae, but visits a
wide variety of flowers including Althaea rosea, Asclepias eriocarpa, Aster exilis,
Brassica, Centaurea solstitialis, Cirsium, Cleome, Corethrogyne bernardense, C.
virgata, Cucurbita, Datura meteloides, Eremocarpus setigerus, Eriogonum setiger, Gilia
virgata, Grindelia, Gutierrezia californica, G. sarothrae, Haplopappus palmer, H.
parishii, Helianthus annuus, Heliotropium curassavicum, Hemizonia fasciculata, H.
pungens, Hugelia virgata, Marrubium vulgare, Medicago sativa, Melilotus, Navarretia,
Phacelia, Silybum marianum, Trichostema lanceolatum.
Melissodes stearnsi Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 101. 2.

2156 Hymenoptera in America North of Mexico

tribas LaBerge. Calif. (San Diego).
Melissodes (Callimelissodes) tribas LaBerge, 1961. Kans. Univ. Sci. Bul. 42: 326. 9.

tuckeri Cockerell. Mont., S. Dak., Nebr., Kans., Tex. Pollen: Unknown, but visits flowers of
Aster, Heterotheca subaxillaris.
Melissodes Tuckeri Cockerell, 1909. Canad. Ent. 41: 129. 2.

Genus MELISSODES Subgenus UNASSIGNED

scotti Cockerell. Calif. (San Clemente Isl.).
Melissodes scotti Cockerell, 1939. Calif. Acad. Sci., Proc. (4) 23: 430. °.

UNRECOGNIZED SPECIES OF THE GENUS MELISSODES LATREILLE

The types of the below listed are either lost or destroyed (see LaBerge, 1961. Kans. Univ. Sci.
Bul. 42: 654).

americana (Lepeletier). Carolina, IIl.

Macrocera americana Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 92. 6.
intermedia Cresson. Tex.

Melissodes intermedia Cresson, 1872. Amer. Ent. Soc., Trans. 4: 278. 9°.
pennsylvanica (Lepeletier). Pa.

Macrocera pennsylvanica Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 97. 6.
philadelphica (Lepeletier). Pa.

Macrocera Philadelphica Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 110. 6.

NoMINA NuDA IN THE GENUS MELISSODES LaATREILLE ~

Melissodes maura Bray, 1917. Jour. Ent. Zool. 9: 94.
Melissodes pullata Bray, 1917. Jour. Ent. Zool. 9: 94.

Genus FLORILEGUS Robertson
Revision: Urban, 1970. Bol. Parana Univ. Federal, Zool. 3 (12): 245-280, 4 figs. (included spp.).

Genus FLORILEGUS Subgenus FLORILEGUS Robertson

Florilegus Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 53.
Type-species: Melissodes condigna Cresson. Monotypic.
condignus (Cresson). N. J. to Fla., west to Nebr., Colo., N. Mex., south to Argentina. Pollen:
Polylectic, visits a wide variety of flowers in the United States where it exhibits a
preference for those of the Leguminosae and particularly Medicago sativa.
Melissodes condigna Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 207. 2.
Melissodes palustris Robertson, 1892. Amer. Nat. 26: 273. 6.
Tetralonia quadrata Bertoni and Schrottky, 1910. Zool. Jahrb., Abt. f. Syst. p. 572. d, 2.
Florilegus pavoninus Cockerell, 1914. N. Y. Ent. Soc., Jour. 22: 316. 2.
Florilegus barticanus Cockerell, 1918. Amer. Mus. Nat. Hist. Bul. 38: 690. ¢.
Tetralonia ecuadoria Friese, 1923. Arkiv. f. Zool. 15 (18): 3. 6, 2.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 253-255, fig. 78
(redescription). —LaBerge and Ribble, 1966. Ent. Soc. Amer., Ann. 59: 948-949, figs. 8-11
(cocoon, larva).

Biology: LaBerge and Ribble, 1966. Ent. Soc. Amer., Ann. 59: 944-950, 11 figs., 1 table
(nesting behavior, nest architecture, floral relationships).

Genus GAESISCHIA Michener, LaBerge and Moure
Revision: Urban, 1963. Bol. Parana Univ. Federal, Zool. 3 (4): 79-129, 7 figs., 1 map, 2 tables
(included spp.).

Taxonomy: LaBerge, 1958. Pan-Pacific Ent. 34: 195-201, 3 figs. (diagnostic characteristics,
classification).

Superfamily APOIDEA 2157

Genus GAESISCHIA Subgenus GAESISCHIA Michener, LaBerge and Moure

Gaesischia Michener, LaBerge and Moure, 1955. Dusenia 6: 220.
Type-species: Svastra fulgurans Holmberg. Orig. desig.

Gaesischia Subgenus Prodasynhalonia LaBerge, 1958. Pan-Pacific Ent. 34: 199.
Type-species: Gaesischia mexicana LaBerge. Monotypic and orig. desig.

The typical subgenus occurs south of the United States.
Genus GAESISCHIA Subgenus GAESISCHIANA Michener, LaBerge and Moure

Gaesischia subg. Gaesischiana Michener, LaBerge and Moure, 1955. Dusenia 6: 224.
Type-species: Gaesischia (Gaesischiana) exul Michener, LaBerge and Moure. Orig.
desig.
Gaesischia subg. Agaesischia Moure and Michener, 1955. Dusenia 6: 273.
Type-species: Eucera patellicornis Ducke. Orig. desig.
exul Michener, LaBerge and Moure. Ariz. to Colombia. Pollen: Polylectic, visits flowers of
Andira inermis, Caesalpinia eriostachys, Dalbergia retusa, Myrospermum fructescens,
Parkinsonia aculeata, Piscidia carthagenensis, Pterocarpus rohrii.
Gaesischia (Gaesischiana) exul Michener, LaBerge and Moure, 1955. Dusenia 6: 224. ¢.

Taxonomy: LaBerge, 1958. Pan-Pacific Ent. 34: 198. °.

Biology: Frankie and Baker, 1974. An. Inst. Nac. Univ. Auton. Mexico 45, ser. Botanica 1:
4-5, 1 fig., 1 table (group foraging). —Jones and Buchmann, 1974. Anim. Behaviour 22: 483,
tables 1, 2 (u. v. floral patterns as orientation cues). —Frankie, Opler and Bawa, 1976.
Jour. Ecol. 64: 1049-1057, 1 fig., 4 tables (foraging behavior).

Genus SIMANTHEDON Zavortink

Simanthedon Zavortink, 1975. Calif. Acad. Sci., Proc. 40: 232.
Type-species: Simanthedon linsleyi Zavortink. Monotypic and orig. desig.
linsleyi Zavortink. Southeastern Ariz. (Cochise Co.), N. Mex. (Eddy Co.); Mexico (Durango).

Pollen: Polylectic, especially Menodora scabra, Agave palmeri, Datura meteloides,
Polygala racemosa, but also visits other flowers for nectar and or pollen including
Cevallia sinuata, Conyza, Desmanthus cooleyi, Hoffmanseggia densiflora, Ipomoea
hirsutula, Mentzelia pumila, Salvia reflexa, Solanum elaeagnifolium.

Simanthedon linsleyi Zavortink, 1975. Calif. Acad. Sci., Proc. 40: 236, 2 figs. d, °.

Genus MARTINAPIS Cockerell
Genus MARTINAPIS Subgenus MARTINAPIS Cockerell

Another subgenus, Svastropsis Moure and Michener, is found in Argentina.

Melissodes subg. Martinella Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 450. Preoce.
Type-species: Melissodes luteicornis Cockerell. Monotypic.
Martinapis Cockerell, 1929. Entomologist 62: 19. N. name.

Revision: Zavortink and LaBerge, 1976. Wasmann Jour. Biol. 34: 119-145, 2 figs., 3 tables
(included spp.).

luteicornis (Cockerell). Kans. (Dodge City), west. Tex., N. Mex. and east. Ariz. (Cochise and
Graham Counties); Mexico (Chihuahua and Durango). Pollen: Polylectic, seems to prefer
to visit plants in the families Leguminosae and Zygophyllaceae for pollen and nectar
including Dalea, Hoffmanseggia, Larrea, and Petalostemon, although a female has been
recorded collecting pollen from Solanum elaeagnifolium; visitation records include
flowers of Acacia, Andropus carnosus, Baileya multiradiata, Cevallia sinuata,
Chilopsis linearis, Chrysothamnus, Dalea lanata, D. scoparia, D. terminalis, Datura
inoxia, Fallugia paradoxa, Gaura suffulta, Hoffmanseggia glauca, Larrea tridentata,
Lygodesmia juncea, Melilotus alba, Mentzelia multiflora, Petalostemon, Prosopis
glandulosa var. torreyanum, Solidago occidentalis, Tribulus, Verbesina encelioides,

Xanthisma texanum.
Melissodes luteicornis Cockerell, 1896. Ann. and Mag. Nat. Hist. (6) 18: 293. 6.

2158 Hymenoptera in America North of Mexico

Taxonomy: Zavortink and LaBerge, 1976. Wasmann Jour. Biol. 34: 130-140, fig. 2
(redescription).

Biology: Linsley and Hurd, 1959. Ent. News 70: 67 (pollen source). —Linsley and Cazier,
1970. Kans. Ent. Soe., Jour. 43: table 1 (pollen source). —Hurd and Linsley, 1975. Smithsn.
Contrib. Zool. 193: 41, tables 1, 3, 4, 6, 8 (floral relationship with Larrea, as luteicornis).
—Zavortink and LaBerge, 1976. Wasmann Jour. Biol. 34: 140-143, tables 1-3 (floral
relationships, seasonal periods of activity).

occidentalis Zavortink and LaBerge. South. Calif. (Colorado and south. Mojave Deserts), and
southwest. Ariz. (Lower Colorado Valley); Mexico (Baja California and Sonora). Pollen:
Polylectic, seems to prefer to visit plants in the families Compositae, Leguminosae,
Solanaceae and Zygophyllaceae for pollen and nectar including Cercidiwm, Dalea,
Datura, Larrea, Palafoxia, Solanum; floral visitation records include Cercidium
floridum, Dalea emoryi, Encelia, Helianthus niveus, Larrea tridentata, Palafoxia
linearis, Wislizenia refracta.

Martinapis occidentalis Zavortink and LaBerge, 1976. Wasmann Jour. Biol. 34: 121, figs. 1,

2. 3, 2:

Biology: MacSwain, 1957. Pan-Pacific Ent. 33: 70 (floral relationships, as lwteicornis).

—Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 41, table 12 (floral relationship with
Larrea, as luteicornis in part). —Zavortink and LaBerge, 1976. Wasmann Jour. Biol. 34:
128-129 (floral relationships, daily and seasonal periods of activity).

Genus MELISSOPTILA Holmberg

Revision: Urban, 1968. Rev. Brasil. Ent. 13: 1-94, 70 figs., 3 maps (included spp.).
Taxonomy: LaBerge, 1957. Amer. Mus. Novitates 1837: 37-38 (tax. characters).

Genus MELISSOPTILA Subgenus MELISSOPTILA Holmberg

Melissoptila Holmberg, 1884. Cordoba. Acad. Nac. de Cien., Actas 5: 119.
Type-species: Melissoptila tandilensis Holmberg. Monotypic.
Thyreotremata Holmberg, 1887. Cordoba. Acad. Nac. de Cien., Bol. 10: 225. Nomen nudum.
Thyreothremma Holmberg, 1903. Mus. Nac. Buenos Aires, An. (3) 2: 391.
Type-species: Melissoptila tandilensis Holmberg. Desig. by Sandhouse, 1943.
(=Thyreothremma rhopalocera Holmberg).

The typical subgenus does not occur in North America.
Genus MELISSOPTILA Subgenus PTILOMELISSA Moure

Ptilomelissa Moure, 1943. Rev. de Ent. 14: 482.
Type-species: Ptilomelissa ochromelaena Moure. Monotypic.
otomita (Cresson). Tex. (Brownsville); Mexico, Guatemala, Honduras, Costa Rica, Panama.
Melissodes otomita Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 209. ¢.
Melissodes pinguis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 216. 2, 3.
Melissodes pinguis var. velutinella Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 158. °.
Tetralonia joseana Friese, 1916. Stettin. Ent. Ztg. 77: 330. 6, 2.
Exomalopsis rufitecta Cockerell, 1949. U.S. Natl. Mus., Proc. 98: 454. °.
Exomalopsis rufitecta var. palliditecta Cockerell, 1949. U.S. Natl. Mus., Proc. 98: 454. 9.

Taxonomy: LaBerge, 1957. Amer. Mus. Novitates 1837: 38 (synonymy, selects otomita
instead of pinguis under provision of first reviser). — Urban, 1968. Rev. Brasil. Ent. 13:
67-71, figs. 29, 44, 53-65, map 1 (as pinguis).

Morphology: Graf, 1970. Bol. Federal Univ. Parana, Zool. 3: 281-287, 1 pl., figs. 1-3 (salivary
glands).

@ TRIBE ANTHOPHORINI

This is a large tribe of pollen-collecting bees that is found throughout much of the world.
Although the Anthophorini contains several genera, most of the species belong to the Old World

Superfamily APOIDEA 2159

genus Amegilla and the genus Anthophora which occurs on all the continents except Australia.
In the New World the tribe is represented by the apparently endemic, North American Deltop-
tila and Emphoropsis and by Anthophora which, although present in both North and South
America, is especially well represented by numerous species in North America. While most of
the studied Anthophorini nest in the ground, at least members of the subgenus Clisodon make
their nests in wood.

Genus ANTHOPHORA Latreille

Taxonomy: Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 64-72. —Michener, 1936. Amer. Mus.
Novitates 876: 1-2 (small group only). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul.
152: 323-331, figs. 90-93 (eastern U.S. spp.).

Genus ANTHOPHORA Subgenus ANTHOPHORA Latreille

Podalirius Latreille, 1802. Hist. Hat. Fourmis, p. 430. Name suppressed by Internatl.
Comn. Zool. Nomencl., Op. 151, 1944.
Anthophora Latreille, 1803. Nouv. Dict. Hist. Nat., v. 18, p. 167. Proposed unnecessarily to
replace Podalirius Latreille.
Type-species: Apis acervorum Linnaeus. Desig. by Internat]. Comn. Zool. Nomencl.,
Op. 151, 1944. (=Apis pilipes Fabricius).
Lasius Panzer, [1801-1802]. Faunae Ins. German., h. 86, no. 16. Name suppressed by
Internat]. Comn. Zool. Nomencl., Op. 151, 1944.
Podalirius subg. Paramegilla Friese, 1897. Die Bienen Europas, v. 3, p. 18.
Type-species: Apis ireos Pallas. Desig. by Sandhouse, 1943.
Anthophoroides Cockerell and Cockerell, 1901. Ann. and Mag. Nat. Hist. (7) 7: 48.
Type-species: Podalirius vallorwm Cockerell. Monotypic and orig. desig.
Anthemoessa Robertson, 1905. Amer. Ent. Soc., Trans. 31: 372. Preoce.
Type-species: Anthophora abrupta Say. Monotypic and orig. desig.
Melea Sandhouse, 1943. U. S. Natl. Mus., Proc. 92: 526. N. name.

Taxonomy: Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 51-62 (western U. S. spp.).
—Michener, 1960. Queensland Nat. 16: 63 (status of Amegilla).
abrupta Say. N. Y. to Fla., west to Mich., south to La. and Tex. Parasite: Hornia m.
minutipennis Riley, Monodontomerus mandibularis Gahan, M. montivagus Ashm.,
Physocephala sagittaria (Say), Xeromelecta californica (Cress.)., X. interrupta (Cress.).
Pollen: Unknown, but visits a wide variety of flowers including Aesculus, Asclepias,
Azalea, Blephilia, Cirsium, Convolvulus, Cornus, Delphinium, Dianthera, Diospyros,
Fransera, Gillenia, Hydrophyllum, Iris, Leonurus, Melilotus, Mertensia, Monarda,
Nepeta, Oenothera, Penstemon, Polygonatum, Rosa, Rubus, Scutellaria, Stachys,
Teucrium, Trifolium, Triosteum, Vicia.
Anthophora abrupta Say, 1837. Boston Jour. Nat. Hist. 1: 409. d.
Anthophora sponsa Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 389. °.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1080, figs. 232, 233 (larva).

Biology: Walsh, 1868. Amer. Ent. 1: 9 (as sponsa). —Ashmead, 1894. Psyche 7: 25. —Frison,
1922. Amer. Ent. Soc., Trans. 48: 137-156. —Rau, 1922. Acad. Sci. St. Louis, Trans. 24 (7):
35. —Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 166. —Rau, 1929. Jour. Compar. Psychol. 9:
53. —Rau, 1929. Psyche 36: 156.
abruptella Cockerell. Calif. Parasite: Sphaeropthalma unicolor (Cress.). Pollen: Unknown, but
visits flowers of Amsinckia intermedia, Eriodictyon californicum, Phacelia distans, P.
ramosissima.
Anthophora abruptella Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 72. °.
affabilis Cresson. Calif., Utah, Colo., N. Mex., Tex. Pollen: Collects pollen from flowers of
Camissonia clavaeformis, Oenothera caespitosa, O. deltoides.
Anthophora affabilis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 189. d, °.
aterrima Cockerell. Calif. Pollen: Unknown, but visits flowers of Satureja douglasii,
Thermopsis macrophyllus.
Anthophora aterrima Cockerell, 1924. Pan-Pacifie Ent. 1: 51. °, 6.

2160 Hymenoptera in America North of Mexico

bomboides bomboides Kirby. Ont. to Ga., west to Wis. and Man. Parasite: Monodontomerus
mandibularis Gahan. Pollen: Unknown, but visits flowers of Ceanothus, Penstemon.
Anthophora bomboides Kirby, 1837. In Richardson, Faune Bor.-Amer., v. 4, p. 271. 6.
Anthophora canadensis Cresson, 1869. Amer. Ent. Soc., Trans. 2: 292. 6.

Taxonomy: Cockerell, 1936. Canad. Ent. 68: 276 (key to this and following subspecies of
bomboides).
bomboides neomexicana Cockerell. S. Dak., Wyo., Idaho, Colo., N. Mex., Calif. (Mono Co.),
?Nebr., ?Tex. Parasite: Hornia neomexicana (Ckll.), Monodontomerus montivagus
Ashm., Sphaeropthalma unicolor (Cress.), Xeromelecta californica (Cress.).
Anthophora bomboides var. neomexicana Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5:
408. 6, 2.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 224 (nest). —Hicks, 1931. Canad. Ent. 63: 175

(nest, parasite).

bomboides scutellaris Swenk. Nebr., Colo.
Anthophora scutellaris Swenk, 1909. Ent. News 20: 391. °.

bomboides sodalis Cresson. Wash., Oreg., east. Calif., Nev., ?7Man., ?Alta.
Anthophora sodalis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 212. 3.

bomboides solitaria Ritsema. B. C., ?Calif.
Anthophora insularis Smith, 1879 (not 1857). Deser. New Species Hym. Brit. Mus., p. 124.

2. Preoce.
Anthophora solitaria Ritsema, 1880. Tijdschr. v. Ent. 23: xevii. N. name.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 331.
bomboides stanfordiana Cockerell. B. C., Wash., Oreg., Calif. Parasite: Dasymutilla aureola
(Cress.), Hornia minutipennis occidentalis Linsley, Xeromelecta californica (Cress.).
Pollen: Unknown, but visits flowers of Castilleja latifolia, Delphinium including D.
hesperium, Dudleya, Eriodictyon californicum, Glycyrrhiza lepidota, Penstemon,
Phacelia distans, P. imbricata, Raphanus sativa, Rosa californica, Stanleya pinnata.
Anthophora stanfordiana Cockerell, 1904. Ent. News 15: 32. 6, 2.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1007, 1078, figs. 7-15, 219-222, 225
(larva).

Biology: Kellogg, 1908. Amer. Ins., p. 517. —Nininger, 1920. Psyche 27: 135. —Linsley and
MacSwain, 1942. Calif. Univ. Pubs. Ent. 7: 189 (habits).
bomboides willingi Cockerell. Sask.
Anthophora bomboides Willingi Cockerell, 1911. Canad. Ent. 43: 34. d.
californica albomarginata Timberlake. Ariz., Calif., desert. Pollen: Unknown, but visits
flowers of Aster spinosa, Chilopsis linearis, Dalea, Heliotropium curassavicum, Larrea
tridentata, Phyla nodiflora, Prosopis glandulosa, Salvia columbariae.
Anthophora californica albomarginata Timberlake, 1937. Amer. Mus. Novitates 958: 13. 9,
é.
californica californica Cresson. Colo., N. Mex., Ariz., Calif.; Mexico (Baja Calif.). Pollen:
Unknown, but visits a wide variety of flowers including Amsinckia douglasiana, A.
intermedia, Astragalus pomonensis, Castilleja latifolia, Cercidium floridum, Dalea
californica, Dudleya aloides, Eriodictyon californicum, Isomeris arborea, Lantana
montevidensis, Lotus scoparius, Lupinus, Oenothera, Penstemon spectabilis, Phacelia
distans, P. imbricata, Pholisma racemosa, Raphanus sativus, Rubus ursinus, Salvia
columbariae, S. mellifera, Scrophularia californica, S. laciniata, Senecio douglasii,
Solanum «anti.
Anthophora californica Cresson, 1869. Amer. Ent. Soc., Trans. 2: 290. ¢.
Anthophora 5-fasciata Provancher, 1895. Nat. Canad. 22: 172. 6.
Anthophora tarsata subtarsata Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 88. d, 9.

Taxonomy: Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 154. ¢.

californica erysimi Cockerell. Calif. (San Miguel Is.). Pollen: Unknown, but visits flowers of

Erysimum insulare.
Anthophora californica erysimi Cockerell, 1937. Pan-Pacific Ent. 13: 150. 2.

Superfamily APOIDEA 2161

californica texana Cresson. Colo., N. Mex., Ariz., Tex.; Mexico (Zacatecas). Pollen: Unknown,
but visits flowers of Caesalpinia falcaria, Chilopsis linearis, Larrea tridentata.
Anthophora texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 282. °.

Taxonomy: Cockerell, 1905. Canad. Ent. 37: 335. 6.
capistrata Cresson. Tex.; 7Mexico (Baja Calif.).
Anthophora capistrata Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 187. 3.
centriformis centriformis Cresson. Nev., Calif. (Sierra Nevada Mts.).
Anthophora centriformis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 212. 3.
centriformis vierecki Cockerell. N. Mex., Nev., Calif. Pollen: Unknown, but visits flowers of
Beloperone californica, Hyptis emoryi, Lotus scoparius, Lupinus odoratus, Nama
parryi, Penstemon spectabilis, Scutellaria angustifolia.
Anthophora centriformis vierecki Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 69. 6.
chlorops chlorops Michener. Wash.
Anthophora chlorops Michener, 1936. Brooklyn Ent. Soc., Bul. 31: 92. 9.
chlorops utahensis Michener. Utah.
Anthophora chlorops utahensis Michener, 1936. Amer. Mus. Novitates 876: 2. 9.
citreostrigata Dours. North America. (No recent records; doubtless not Nearctic).
Anthophora citreo-strigata Dours, 1868. Soc. Linn. Nord France, Mem. 71: 95. 9.
coptognatha Timberlake. Ariz., Calif. deserts. Pollen: Unknown, but visits flowers of Dalea
fremontii, Lantana, Larrea tridentata, Lycium, Prosopis, Salvia pilosa.
Anthophora coptognatha Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 54. 3, @.
crotchii Cresson. B. C., Wash., Calif. Pollen: Unknown, but visits a wide variety of flowers
including Amsinckia douglasiana, A. eastwoodae, A. intermedia, Astragalus, Brassica,
Brodiaea capitata, Caulanthus heterophyllus, Delphinium parryi, Eriodictyon, Lantana
montevidensis, Lupinus, Marrubium vulgare, Oenothera, Orthocarpus purpurascens,
Raphanus sativus, Salvia carduacea, S. columbariae, Trichostema parishii.
Anthophora crotchii Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 192. 3.
Anthophora Washingtoni Cockerell, 1905. In Viereck, Canad. Ent. 37: 313. 9.

Taxonomy: Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 161. 3.

dammersi Timberlake. Calif. Pollen: Collects pollen from flowers of Camissonia kernensis, but
also visits flowers of Amsinckia douglasiana, A. intermedia, Beleperone californica,
Chaenactis fremontii, Salazaria mexicana, Salvia carduacea.
Anthophora dammersi Timberlake, 1937. Amer. Mus. Novitates 958: 15. d, 9.

edwardsii edwardsii Cresson. Wash. to Calif., Nev. and Utah. Parasite: Anthrax sp. near fur
O.S., Dasymutilla aureola (Cress.), Melecta separata callura (Ckll.), Meloe fransiscana
Van Dyke, Mythicomyia sp. near pusilla Melander, Nemognatha scutellaris LeC.,
Sphaeropthalma unicolor (Cress.), Tricrania stansburyi Hald., Xeromelecta californica
(Cress.). Pollen: Collects pollen from Amsinckia and Astragalus, but visits these and a
wide variety of other flowers for nectar including Brassica, Brodiaea, Cryptantha,
Erodium, Eriogonum, Lantana, Layia, Lupinus, Mentzelia, Orthocarpus, Phacelia,
Plagiobothrys, Prunus, Raphanus, Ribes, Salix, Salvia, Solanum. Predator: Anthrenus
scrophulariae (L.). Ptinus californicus Pic.

Anthophora edwardsti Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 190. 2, 6.

Taxonomy: Michener, 1958. Kans. Univ. Sci. Bul. 35: 1082, figs. 228, 234, 235 (larva).
—Timberlake, 1951. N. Y. Ent. Soce., Jour. 59: 62.

Biology: Linsley and MacSwain, 1941. South. Calif. Acad. Sci., Bul. 40: 130. —Thorp, 1969.
Amer. Midland Nat. 82: 321-337, 7 figs., 1 map, 2 tables (nest, flight behavior, flower
relationships, enemies).

edwardsii gohrmanae Cockerell. Colo., N. Mex., Calif.
Anthophora gohrmanae Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 454. d.
Anthophora gohrmanae var. coloradensis Michener, 1936. Brooklyn Ent. Soc., Bul. 31: 92.
Os 2s

Taxonomy: Timberlake, 1951. N. Y. Ent. Soce., Jour. 59: 62.

2162 Hymenoptera in America North of Mexico

fedorica Cockerell. Tex.
Anthophora fedorica Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 70. ¢.

Taxonomy: Cockerell, 1908. Ent. Soc. Wash., Proc. 9: 71. 2.
forbesi Cockerell. Ariz., Calif. (Death Valley), Mexico (Baja Calif.). Parasite: Melecta separata

arizonica (Ckll.). Pollen: Unknown, but visits flowers of Larrea tridentata.
Anthophora forbesi Cockerell, 1907. Canad. Ent. 39: 354. °.

Taxonomy: Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 59. ¢.

Biology: Lutz and Cockerell, 1920. Amer. Mus. Nat. Hist., Bul. 42: 574 (parasite).
frontata Say. La.
Anthophora frontata Say, 1837. Boston Jour. Nat. Hist. 1: 409. ¢.
fulvicauda Timberlake. Calif. Pollen: Collects pollen from flowers of Camissonia clavaeformis
but also visits flowers of Chilopsis linearis, Eriodictyon crassifolium, Stephanomeria
exigua.
Anthophora fulvicauda Timberlake, 1937. Amer. Mus. Novitates 958: 14. 9, 6.
fulvicollis Timberlake. Calif., Ariz. Pollen: Collects pollen from flowers of Camissonia
clavaeformis.
Anthophora fulvicollis Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 59. 3.
fumipennis Swenk. Nebr.
Anthophora fumipennis Swenk, 1909. Ent. News 20: 391. 9.
fuscipennis Smith. North America. (No recent records).
Anthophora fuscipennis Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 338. 2.
lesquerellae (Cockerell). N. Mex. Pollen: Unknown, but visits flowers of Lesquerella fendleri,
Lycium, Ungnadia speciosa.
Podalirius lesquerellae Cockerell, 1896. Canad. Ent. 28: 197. 2, 6.

Taxonomy: Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 399. 3.

linsleyi Timberlake. South. Calif., arid regions. Parasite: Anthrax nidicola Cole, Hornia
boharti Linsley, Hylemya cilicrura Rond., Lytta chloris Fall, L. occipitalis Horn, L.
purpurascens Fall, Monodontomerus montivagus Ashm., Myopa rubida (Bigot),
Nemognatha scutellaris LeC., Sphaeropthalma unicolor (Cress.)., Triepeolus mojavensis
Linsley, Xeromelecta californica (Cress.). Pollen: Primarily associated with flowers of
Salvia carduacea, but also visits flowers of Amsinckia intermedia, Dalea saundersii, D.
schottii, Larrea tridentata, Lupinus odoratus, Monardella exilis, Phacelia distans,
Salazaria. Predator: Ptinus californicus Pic, Trogoderma ajax Casey, T. simplex
Jayne.

Anthophora linsleyi Timberlake, 1941. Pan-Pacifie Ent. 17: 34. 6, 9.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1082, figs. 227, 236, 237 (larva).

Biology: Linsley and MacSwain, 1942. Amer. Midland Nat. 27: 402-417 (parasites, predators,
scavengers). —Ferguson, 1962. Calif. Univ. Pubs. Ent. 27: 76 (parasites). —Hurd and
Linsley, 1975. Smithsn. Contrib. Zool. 193: 43 (nest, floral relationships).

marginata Smith. N. Mex., Ariz.; Mexico.
Anthophora marginata Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 339. ¢.

Taxonomy: Cockerell, 1905. Amer. Ent. Soc., Trans. 31: 331. 2, ¢.
montana Cresson. Colo., Tex., N. Mex., Ariz. Parasite: Sphaeropthalma unicolor (Cress.).
Pollen: Unknown, but visits flowers of Cleome serrulata, Delphinium, Lycium vulgare,
Mentzelia pumila, Salvia lanceolata.
Anthophora montana Cresson, 1869. Amer. Ent. Soc., Trans. 2: 290. 2.

Biology: Linsley, 1962. Ent. Soc. Amer., Ann. 55: 161, fig. 9.
nebracensis Swenk. Nebr. Pollen: Unknown, but visits flowers of Monarda fistulosa.
Anthophora nebracensis Swenk, 1909. Ent. News 20: 390. °.

neglecta Timberlake and Cockerell. Calif. Pollen: Collects pollen from flowers of Camissonia
clavaeformis, but also visits flowers of Abronia villosa, Amsinckia douglasiana, A.
intermedia, A. tessellata, Acamptopappus sphaerocephalus, Arctostaphylos, Astragalus

’

Superfamily APOIDEA 2163

pomonensis, Delphinium, Ephedra, Isomeris arborea, Lantana montevidensis, Lotus
rigidus, Lupinus bicolor, Melilotus alba, Phacelia distans, Ranunculus, Rhus ovata,
Salix, Salvia columbariae, S. pilosa, Sisymbrium irio, Trichostema lanatum.

Anthophora neglecta Timberlake and Cockerell, 1936. In Cockerell, Amer. Mus. Novitates
831: 4. 6, 2.

nigrita Dours. “Indiana.” No recent records; probably not North American.
Anthophora nigrita Dours, 1869. Soc. Linn. Nord France, Mem. 2: 137. ¢.
Podilirius nigratus(!) Dalla Torre, 1896. Cat. Hym., v. 10, p. 278.

Taxonomy: Moure, 1960. Studia Ent. 3: 118-119 (status of A. nigrita (Fabr.) and A. nigrita
Dours).
occidentalis Cresson. B. C., Mont., S. Dak., Oreg., Wyo., Kans., Colo., Utah, N. Mex., Ariz.

Parasite: Anthrax fur (O. S.), Chrysura smaragdicolor (Walk.), Dasymutilla dugesii
(Ckll. and Casad), D. fulvohirta (Cress.), Hornia m. minutipennis Riley, H.
neomexicana (Ckll.), Monodontomerus montivagus Ashm., Nemognatha lurida Cress.,
N. piezata Fabr., Sphaeropthalma unicolor (Cress.), Triepeolus sp., Xeromelecta
californica (Cress.). Pollen: Unknown, but visits flowers of Carduus, Cleome serrulata,
Convolvulus sepium.

Anthophora occidentalis Cresson, 1869. Amer. Ent. Soc., Trans. 2: 292. 2, ¢.

Biology: Johnson, 1963. Ent. News 14: 290. —Hungerford and Williams, 1912. Ent. News 23:
259. —Long, 1925. Carnegie Inst. Wash., Yearbook 24: 340. —Mickel, 1928. Ent. News 39:
69. —Porter, 1951. Iowa State Coll. Jour. Sci. 26: 23-30, 19 figs. —Hobbs, Nummi and
Virostek, 1961. Canad. Ent. 93: 142-148 (nests, nesting site, nest associates).
pacifica infernalis (Dalla Torre). Utah, Nev., south. Calif. Parasite: Melecta pacifica fulvida
Cress. Pollen: Unknown, but visits flowers of Amsinckia intermedia, Arctostaphylos
glandulosa, A. glauca, Buddleia, Lantana montevidensis, Lupinus bicolor, Pedicularis
densiflora, Rhus laurina, Ribes indecorum, R. nevadensis, R. speciosum, Solanum
douglasit.
Anthophora carbonaria Cresson, 1879. Amer. Ent. Soc., Trans. 7: 210. 2. Preoce.
Podalirius infernalis Dalla Torre, 1896. Cat. Hym., v. 10, p. 273. N. name.
Anthophora corvicolor Cockerell, 1905. Ent. News 16: 81. 9.
pacifica pacifica Cresson. B. C., Wash., Oreg., north. Calif., Nev. Pollen: Unknown, but visits
flowers of Arctostaphylos glauca, Cercocarpus ledifolius, Cynoglossum grandis,
Eriodictyon californicum, Erodium botrys, Lantana montevidensis, Lasthenia
chrysostoma, Lotus glaber, Lupinus bicolor, L. densiflorus, Pedicularis densiflora,
Ribes indecorum, R. malvaceum, Salix lasiolepis.
Anthophora pacifica Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 190. 3.
Anthophora ignava Cresson, 1879. Amer. Ent. Soc., Trans. 7: 210. 2.
pernicis Timberlake. Ariz., Nev. Pollen: Unknown, but visits flowers of Ribes cereum.
Anthophora pernicis Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 58. 3.
platti Timberlake. South. Calif. Pollen: Unknown, but visits flowers of Lantana montividensis,
Salix, Salvia mellifera.
Anthophora platti Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 60. 3, °.
porterae Cockerell. Wyo., Colo., N. Mex., Calif. (desert). Pollen: Polylectic, visits flowers of
Astragalus including A. crotalariae, A. fremontii, Lupinus, Oenothera deltoides, Ribes.
Anthophora porterae Cockerell, 1900. Ann. and Mag. Nat. Hist. (7) 5: 407. d, 2.
Anthophora porterae var. semiflava Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 183. d.
Anthophora porterae var. watsoni Cockerell 1911. Amer. Ent. Soc., Trans. 37: 238. d.
Anthophora porterae mut. Thalassiana Cockerell, 1920. Nature 105: 518. ¢.
raui Rohwer. Mo., Colo. to N. Y.
Anthophora raui Rohwer, 1923. Ent. Soc. Wash., Proc. 25: 100. 9.

Biology: Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 179. —Rau, 1929. Psyche 36: 555.
smithii Cresson. S. Dak., Kans., Colo., Tex., N. Mex., Ariz. Pollen: Unknown, but visits flowers
of Carduus, Cleome serrulata, Cnicus, Grindelia subalpina, Petalostemon oligophyllus,
Salvia pitcheri, Verbena macdougali.
Anthophora smithii Cresson, 1869. Amer. Ent. Soc., Trans. 2: 289. 6, °.
Podalirius cardui Cockerell, 1897. Amer. Ent. Soc., Trans. 24: 155. ¢.

2164 Hymenoptera in America North of Mexico

Biology: Banks, 1902. N. Y. Ent. Soc., Jour. 10: 212. —Linsley, 1962. Ent. Soc. Amer., Ann.
55: 161-162, fig. 9.
subignava Cockerell. Colo.
Anthophora subignava Cockerell, 1929. N. Y. Ent. Soc., Jour. 37: 443. 9.
urbana catalinae Cockerell. Calif. (Santa Catalina Isl.).
Anthophora catalinae Cockerell, 1901. Canad. Ent. 33: 297. 2.
urbana clementina Cockerell. Calif. (San Clemente Isl.).
Anthophora catalinae clementina Cockerell, 1939. Calif. Acad. Sci., Proce. (4) 23: 430. 2, 3.
urbana nicolai Cockerell. Calif. (San Nicolas Isl.).
Anthophora nicolai Cockerell, 1939. Calif. Acad. Sci., Proc. (4) 23: 429. 2, 3.
urbana urbana Cresson. Idaho, Wash., Colo., Utah, N. Mex., Ariz., Calif.; Mexico (Baja
California and Sonora). Parasite: Meloe niger Kirby, Xeromelecta californica (Cress.),
Zodion obliquefasciatum (Macq.)? Pollen: Highly polylectic, taking pollen and or nectar
from a great variety of annuals and perennials including Allium, Amsinckia intermedia,
Amorpha californica, Arctostaphylos drupacea, A. glandulosa, A. glauca, A. nevadensis,
Artemisia tridentata, Asclepias eriocarpa, A. sublata, Aster canescens, Astragalus
parishii, Baileya, Bebbia juncea, Brassica campestris, B. geniculata, Brodiaea crocea, B.
laxa, B. lutea, Cakile edentula, Calycadenia multiglandulosa, Camissonia, Centaurea
solstitialis, Cercidium floridum, Chaenactis artemisiaefolia, C. glabriuscula,
Chaemabatia foliolosa, Chamaenarion angustifolium, Chilopsis linearis, Chrysopsis
fastigiata, C. villosa, Chrysothamnus nauseosus, Cirsium californicum, Clarkia biloba,
C. cylindrica, C. rhomboidea, C. unguiculata, Clematis lasiantha, Cleomella obtusifolia,
Convolvulus arvense, C. occidentalis, Cordylanthus nevinii, Corethrogyne, Cryptantha
intermedia, Dalea californica, D. emoryi, D. saundersii, Durantia plumieri, Epilobium,
Eriastrum densiflora, E. sapphirinum, E. virgatum, Erigeron divergens, E.
stenophyllus, Eriodictyon californicum, E. parryi, E. trichocalyx, Eriogonum
confertiflorum, E. fasciculatum, E. nodosum, E. nudum, E. subscaposum, Eryngium
aristatulum, Geraea canescens, Gilia capitata, G. tricolor, Gormania obtusata, Grindelia
camporum, Gutierrezia californica, G. lucida, G. sarothrae, Haplopappus acradenius, H.
linearifolius, H. palmeri, H. squarrosus, H. vernonioides, Helianthus annuus, H.
gracilentus, Heliotropium curassavicum, Hemizonia paniculata, H. pungens, H.
wheeleri, Hoffmanseggia, Horkelia bernardina, Hyptis emoryi, Iris, Lantana
montividensis, Larrea tridentata, Lathyrus splendens, Lepidospartum squamatum,
Lessingia germanorum, Lonicera japonica, Lotus argophyllus, L. davidsonii, L.
hamatus, L. leucophaeus, L. purshianus, L. scoparius, Lupinus lobbii, L. superbus,
Lythrum californicum, Malacothamnus densiflorus, Marrubium vulgare, Medicago
sativa, Melilotus alba, Mentzelia multiflora, Mimulus guttatus, Mirabilis laevis,
Monardella lanceolata, M. linoides, M. odoratissima, M. stricta, M. villosa, Navarretia,
Oenothera, Palafoxia linearis, Penstemon antirrhinoides, P. grinnelli, P. heterophyllus,
P. heterodoxus, P. palmeri, P. spectabilis, Pentachaeta aurea, Phacelia cicutaria, P.
ciliata, P. distans, P. frigida, P. imbricata, P. ramosissima, Phyllodoce brewer,
Potentilla glandulosa, Prosopis glandulosa, Ranunculus californicus, Raphanus
sativus, Rhus laurina, Rosa californica, Salazaria mexicana, Salvia apiana, S.
carduacea, S. carnosa, S. clevelandii, S. mellifera, S. pachyphylla, S. pilosa,
Scrophularia californica, Senecio douglasii, S. integerrimus, Sisymbrium, Solanwm
douglasii, Solidago californica, S. multiradiata, S. occidentalis, Stachys bullata, S.
californica, Stephanomeria exigua, S. pauciflora, S. virgata, Streptanthus tortuosus,
Trichostema lanceolatum, T. laxum, T. parishii, Verbena californica, Vicia americana,
Wislizenia refracta, Zauschneria californica. Predator: Trogoderma incisulum LeC., T.
sternale Jayne.
Anthophora urbana Cresson, 1878. Acad. Nat. Sci., Phila., Proc. 30: 188. 2, ¢.
Podalirius alamosanus Cockerell, 1896. Canad. Ent. 28: 195. 2.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1080, figs. 223, 224, 226 (larva).

Biology: Davidson, 1897. South. Calif. Acad. Sci., Proc. 1: 4 (as montana). —Linsley, 1962.
Ent. Soc. Amer., Ann. 55: 161, fig. 9 (sleep). —Mayer and Johansen, 1976. Pan-Pacific Ent.
52: 120-125 (nest site, nest architecture, adult activity, development of life stages,
parasites, predators, associates).

Superfamily APOIDEA = 2165

ursina californiensis Michener. Calif.
Anthophora simillima var. californiensis Michener, 1935. Pan-Pacific Ent. 11: 182. dg, 2.
ursina simillima Cresson. Wash., Wyo., Colo., N. Mex., Ariz., Calif. Pollen: Unknown, but
visits flowers of Astragalus.
Anthophora simillima Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 189. 6.
Anthophora euops Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 451. 6.

Taxonomy: Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 478. 3, °.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 225.
ursina ursina Cresson. Wis. and IIl, east to New England, south tc Ga., 7N. Mex. Parasite:

Melecta pacifica atlantica Linsley? Pollen: Unknown, but visits flowers of Aesculus,
Asclepias, Astragalus, Azalea, Cercis, Collinsia, Delphinium, Dicenira, Dodacatheon,
Geranium, Lithospermum, Lonicera, Mertensia, Monarda, Pedicularis, Penstemon,
Phlox, Physalis, Polygonatum, Polymonium, Ribes, Scutellaria, Trifolium, Vaccinium,
Vicia, Viola.

Anthophora ursina Cresson, 1869. Amer. Ent. Soc., Trans. 2: 291. 3.

Anthophora pyralitarsis Dours, 1869. Soe. Linn. Nord France, Mem. 2: 160. é.

Taxonomy: Robertson, 1905. Amer. Ent. Soc., Trans. 31: 372. 2, 6. —Timberlake, 1951. N. Y.
Ent. Soe., Jour. 59: 57.
vallorum (Cockerell). N. Mex.; Mexico. Parasite: Hornia neomexicana (Ckll.). Pollen:
Unknown, but visits flowers of Chilopsis, Ipomoea, Proboscidea, Solanum
elaeagnifolium.
Podalirius vallorum Cockerell, 1896. Canad. Ent. 28: 195. d, 9.
vannigera Timberlake. Calif., Ariz. Pollen: Unknown, but visits flowers of Jsomeris arborea,
Lycium fremontii, L. torreyi, Phacelia distans, Salvia pilosa.
Anthophora vannigera Timberlake, 1951. N. Y. Ent. Soc., Jour. 59: 56. 3.
walshii Cresson. Mass., Wis., Ill., lowa, Nebr., Mo., Kans., Colo., Tex., ?Ariz. Pollen: Unknown,
but visits flowers of Abutilon, Baptisia, Blephilia, Cassia, Chamaecrista, Cirsium,
Helianthus, Lespedeza, Lobelia, Petalostemon, Ruellia, Teucrium.
Anthophora walshii Cresson, 1869. Amer. Ent. Soc., Trans. 2: 290. d, 2.

Genus ANTHOPHORA Subgenus CLISODON Patton

Clisodon Patton, 1879. U. S. Geol. and Geog. Survey, Bul. 5: 479.
Type-species: Anthophora furcata terminalis Cresson. Monotypic and orig. desig.
(=Anthophora terminalis Cresson).

Anthophora furcata furcata (Panzer) and other subspecies not listed here are Palearctic.

Taxonomy: Cockerell, 1936. Canad. Ent. 68: 276 (key). —Timberlake, 1951. N. Y. Ent. Soc.,
Jour. 59: 51 (status).
furcata neofurcata (Sladen). B. C.
Clisodon neofurcata Sladen, 1919. Canad. Ent. 51: 125. dg, 9.
furcata pernigra Cresson. Nev., Calif.
Anthophora pernigra Cresson, 1879. Amer. Ent. Soc., Trans. 7: 210. @.
furcata sperryi (Cockerell). Ariz.
Clisodon terminalis sperryi Cockerell, 1937. South. Calif. Acad. Sci., Bul. 36: 107. ¢.
furcata syringae (Cockerell). Wash., Calif. Pollen: Unknown, but visits flowers of Castilleia,
Solidago, Syringa.
Podalirius syringae Cockerell, 1898. Acad. Nat. Sci. Phila., Proc. 50: 54. d.
Taxonomy: Cockerell, 1924. Pan-Pacific Ent. 1: 50. —Michener, 1953. Kans. Univ. Sci. Bul. 35:
1082, figs. 229-231 (larva).
furcata terminalis Cresson. Alta. and ?B. C. to Que., south to Colo., Utah, N. Mex., Ariz., IIl.
and N.C. Pollen: Unknown, but visits flowers of Allium, Blephilia, Linum lewisii,
Medicago, Mertensia, Monarda, Penstemon, Polemonium, Pontederia cordata, Prunella,
Rosa, Salvia, Scutellaria, Stachys palustris.
Anthophora terminalis Cresson, 1869. Amer. Ent. Soc., Trans. 2: 292. 9, 6.
Ceratina bidentata Provancher, 1882. Nat. Canad. 13: 234. d.

2166 Hymenoptera in America North of Mexico

Anthophora nudata Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 336. 6.

Anthophora subglobulosa Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., v. 2,
p. 297. 6.

Anthophora nubiterrae Viereck, 1903. Amer. Ent. Soc., Trans. 29: 45. d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 328-330, figs. 91, 92, table 10
(redescription, synonymy).

Biology: Cockerell, 1903. Birds and Nature, v. 14, p. 127. —Medler, 1964. Canad. Ent. 96:
1332-1336, 4 figs.

Genus ANTHOPHORA Subgenus MICRANTHOPHORA Cockerell

Anthophora subg. Micranthophora Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 66.
Type-species: Anthophora curta Provancher. Orig. desig.
abroniae Timberlake. Ariz., Calif.; Mexico. Pollen: Unknown, but visits flowers of Abronia
including A. villosa, Chaenactis fremontii, C. glabriuscula, Geraea canescens,
Heliotropium curassavicum.
Anthophora abroniae Timberlake, 1937. Amer. Mus. Novitates 958: 6. d, 2.

albata Cresson. Kans., Colo.
Anthophora albata Cresson, 1876. Davenport Acad. Sci., Proc. 1: 211. 2.

arthuri Cockerell. Colo. Parasite: Zacosmia maculata (Cress.).
Anthophora arthuri Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 72. 9.

cockerelli Timberlake. Calif., deserts. Pollen: Unknown, but visits flowers of Cleomella
obtusifolia, Dalea, Heliotropium curassavicum, Wislizenia refracta.
Anthophora cockerelli Timberlake, 1937. Amer. Mus. Novitates 958: 4. d, °.

columbariae Timberlake and Cockerell. Calif. Pollen: Unknown, but visits flowers of
Chaenactis fremontii, C. glabriuscula, Cryptantha intermedia, Lotus scoparius,
Lupinus nanus, Phacelia minor, Salvia carduacea, S. columbariae, Salvia mellifera,
Sphaeralcea ambigua, Trifolium variegatum.
Anthophora (Micranthophora) columbariae Timberlake and Cockerell, 1937. In Cockerell,
Amer. Mus. Novitates 948: 9. gd, 2.

curta curta Provancher. Calif.; Mexico (Baja California). Parasite: Zacosmia maculata

(Cress.). Pollen: Polylectic, visits a wide variety of flowers including Abronia villosa,
Acacia greggii, Baileya pleniradiata, Bebbia juncea, Calycadenia tenella, Centaurea
melitensis, C. solstitialis, Chaenactis artemisaefolia, C. fremontii, C. glabriuscula, C.
tenuifolia, Chrysopsis villosa, Chrysothamnus nauseosus, Chrysanthemum coronaria,
Cnicus, Convolvulus arvense, Coreopsis lanceolata, Corethrogyne bernardina, Croton
californicus, Cryptantha intermedia, C. lepida, Curcurbita foetidissima, Encelia actoni,
E. californica, E. farinosa, Eriodictyon trichocalyx, Eriogonum fasciculatum,
Eriophyllum multicaule, Grindelia camporum, G. elata, Gutierrezia californica, G.
lucida, G. sarothrae, Haplopappus palmeri, Helianthus annuus, H. gracilentus,
Heliotropium curassavicum, Hemizonia fasciculata, H. paniculata, H. pungens, H.
wrightii, Heterotheca grandiflora, Jaumea carnosa, Layia platyglossa, Lotus scoparius,
Melilotus alba, Palafoxia linearis, Pentachaeta aurea, Peucephyllum schottii, Phacelia
distans, Phyla nodiflora rosea, Raphanus sativus, Salvia columbariae, Senecio
californica, S. douglasii, Stephanomeria exigua, S. virgata, Trichostema lanceolatum, T.
parishii, Viguiera laciniata, V. multiflora.

Anthophora curta Provancher, 1895. Nat. Canad. 22: 173. °.

Anthophora curta var. ensenadensis Cockerell, 1941. San Diego Soc. Nat. Hist., Trans. 9:
349. 6.

Taxonomy: Cockerell, 1907. Ent. News 18: 396. 6.
curta melanops Cockerell. Colo., Tex., N. Mex., Ariz.; Mexico. Parasite: Zacosmia maculata
(Cress.). Pollen: Unknown, but visits flowers of Cleome, Dithyraea wislizeni, Pectis
papposa, Wedelia incarnata.
Anthophora curta var. melanops Cockerell, 1926. Pan-Pacific Ent. 3: 84. ¢.

Biology: Hicks, 1934. Colo. Univ., Studies 21: 265 (nest, parasite).

Superfamily APOIDEA 2167

erythrothorax Michener. Calif.
Anthophora erythrothorax Michener, 1936. Brooklyn Ent. Soc., Bul. 31: 93. °.
estebana Cockerell. South. Calif. deserts; Mexico (Baja California). Pollen: Unknown, but visits
flowers of Bebbia juncea, Chaenactis fremontii, Dalea, Encelia farinosa, Hyptis emoryi,
Peucephyllum schottii.
Anthophora estebana Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 81. d.
exigua Cresson. N. Mex., Nev., Calif. Pollen: Unknown, but visits flowers of Chaenactis
fremontii, Chrysothamnus nauseosus speciosus, C. viscidiflorus, Haplopappus
linearifolius, H. monactis, Lessingia, Penstemon, Senecio douglasii.
Anthophora exigua Cresson, 1879. Amer. Ent. Soc., Trans. 7: 211. 9, d.
flavocincta Huard. Calif. Pollen: Unknown, but visits Siiwers of Artemisia, Cressa cretica,
Heliotropium curassavicum, Hemizonia paniculata.
Anthophora nigrocincta Provancher, 1895. Nat. Canad. 22: 172. 6. Preoce.
Anthophora flavocincta Huard, 1897. Nat. Canad. 24: 25. N. name.
Anthophora anstrutheri Cockerell, 1906. Amer. Ent. Soc., Trans. 32: 71. @.
flexipes Cresson. Utah, Colo., Nev., Calif. Parasite: Anthrax limatulus Marst., Zacosmia
— maculata (Cress.), Zonitis (Neozonitis) hesperis Selander. Pollen: Polylectic, stores
pollen from a wide variety of flowers.
Anthophora flexipes Cresson, 1879. Amer. Ent. Soc., Trans. 7: 211. 2, 6.

Biology: Torchio and Youssef, 1968. Kans. Ent. Soc., Jour. 41: 289-302, 21 figs. (life history).
hololeuca Cockerell. Ariz., South. Calif.; Mexico (Baja California and Sonora). Pollen: Polylege,
most commonly taken at flowers of Dalea, Acacia and Cercidium, but visits other

flowers including Abronia, Acamptopappus, Bebbia, Croton, Cryptantha, Heliotropium,
Koeberlinia, Larrea, Oenothera, Palafoxia, Stephanomeria.
Anthophora hololeuca Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 82. 2, 6.
maculifrons Cresson. N. Mex., Nev., Calif.; Mexico (Baja California). Pollen: Unknown, but
visits flowers of Chrysothamnus including C. nauseosus, C. viscidiflorus, Helianthus
annuus.
Anthophora maculifrons Cresson, 1879. Amer. Ent. Soc., Trans. 7: 210. 2, d.
mortuaria Timberlake. Calif. Pollen: Unknown, but visits flowers of Heliotropium
curassavicum, Pluchea sericea.
Anthophora mortuaria Timberlake, 1937. Amer. Mus. Novitates 958: 10. d, °.
nigritula Cockerell. Calif.
Anthophora nigritula Cockerell, 1924. Pan-Pacifie Ent. 1: 52. 2.

Taxonomy: Michener, 1936. Brooklyn Ent. Soc., Bul. 31: 93. ¢.
pachyodonta Cockerell. Nev., South. Calif.; Mexico (Baja California), deserts. Pollen:
Unknown, but visits flowers of Bebbia juncea, Encelia farinosa, Heliotropium
curassavicum, Sphaeralcea emoryi, S. orcutti.
Anthophora pachyodonta Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 80. d..2
peritomae Cockerell. Wyo., Utah, Colo., N. Mex. Parasite: Anthrax limatulus artemisia
Marst., Zacosmia maculata (Cress.). Pollen: Stores pollen from flowers of Grindelia
squarrosa, but visits other flowers including Chrysopsis, Chrysothamnus, Helianthus
annuus, Solidago canadensis.
Anthophora curta var. peritomae Cockerell, 1905. Ent. News 16: 272. ¢.
Anthophora peritomae var. interspersa Cockerell, 1907. Ent. News 18: 397. ¢.
Anthophora peritomae var. tinctula Cockerell, 1907. Ent. News 18: 397. °.

Taxonomy: Cockerell, 1907. Ent. News 18: 396. d, 9.

Biology: Torchio, 1971. Los Angeles Co. Mus., Contrib., Sci. 206: 1-14, 13 figs. (life history).
petrophila Cockerell. N. Mex., Calif. Pollen: Unknown, but visits flowers of Acamptopappus
sphaerocephalus, Amsinckia intermedia, A. tessellata, Aster canescens, Astragalus,
Baileya pleniradiata, Bebbia juncea, Chaenactis brachypappa, C. carphoclinia, C.
glabriuscula, Chrysothamnus nauseosus, Dalea saundersii, Erigeron foliosus,
Eriophyllum confertiflorum, Geraea canescens, Gutierrezia lucida, Haplopappus
acradenius, H. cooperi, H. linearifolius, H. monactis, Lessingia tenuis, Lotus davidsonii,

2168 Hymenoptera in America North of Mexico

L. scoparius, Monardella exilis, Salvia pilosa, Stephanomeria exigua, S. pauciflora S.

virgata.
Anthophora curta petrophila Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 15. °.

Taxonomy: Cockerell, 1907. Ent. News 18: 395. ¢.
phenax (Cockerell). Tex. to South. Calif., Mexico. Pollen: Unknown, but visits flowers of
Lycium including L. torreyi.
Podalirius phenax Cockerell, 1898. Canad. Ent. 30: 146. 3.
rhodothorax Michener. Calif. Pollen: Unknown, but visits flowers of Chrysopsis villosa,
Erigeron, Lessingia glandulifera, Monardella stricta.
Anthophora rhodothorax Michener, 1936. Brooklyn Ent. Soc., Bul. 31: 94. 2, 6.
Anthophora emarginata Timberlake, 1937. Amer. Mus. Novitates 958: 11. 6, °.
salazariae Timberlake. Calif., deserts. Pollen: Unknown, but visits flowers of Baileya, Bebbia
juncea, Chaenactis carphoclinia, C. fremontii, Dalea saundersii, Erodium cicutarium,
Haplopappus cooper, Larrea tridentata, Lotus scoparius, Oenothera clavaeformis, O.
desertorum, Phacelia distans, Palafoxia linearis, Salazaria mexicana, Salvia
columbariae, Senecio longilobus.
Anthophora salazariae Timberlake, 1937. Amer. Mus. Novitates 958: 7. 2, d.

Genus EMPHOROPSIS Ashmead

Emphoropsis Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 60.
Type-species: Habropoda floridana Smith. Desig. by Cockerell and Cockerell, 1901.
Meliturgopsis Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 62.
Type-species: Emphoropsis murina Cockerell. First included species, Cockerell,
1909. (=Emphoropsis murihirta murina Cockerell).
Psithyrus subg. Laboriopsithyrus Frison, 1927. Amer. Ent. Soc., Trans. 53: 69.
Type-species: Bombus laboriosus Fabricius. Orig. desig.

Taxonomy: Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 99-100.

birkmanni Cockerell. Tex.
Emphoropsis Birkmanni Cockerell, 1905. Canad. Ent. 37: 265. 9.

Taxonomy: Cockerell, 1934. Amer. Mus. Novitates 732: 2. 2, 6.

Biology: Cockerell, 1934. Amer. Mus. Novitates 732: 2 (nest).
cineraria (Smith). B. C., Wash., Calif. Parasite: Melecta thoracica (Sm.), M. pacifica Cress.
Pollen: Unknown, but visits flowers of Arctostaphylos mariposa, A. patula, Ribes.
Anthophora cineraria Smith, 1879. Deser. New Species Hym. Brit. Mus., p. 124. 2, 6.

Taxonomy: Cockerell, 1904. Canad. Ent. 36: 302.

Biology: Torchio and Youssef, 1968. Kans. Ent. Soc., Jour. 41: 300 (parasite).
citula Cockerell. Colo.
Emphoropsis citulus Cockerell, 1929. N. Y. Ent. Soc., Jour. 37: 442. ¢.
cressoni (Dalla Torre). Wyo., Colo. 7B. C. Pollen: Unknown, but visits flowers of Delphinium,
Ribes longiflorum.
Anthophora mucida Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 191. 2. Preoce.
Podalirius cressoni Dalla Torre, 1896. Cat. Hym., v. 10, p. 264. N. name.
Emphoropsis mucida var. johnsoni Cockerell, 1905. Entomologist 38: 59. 2, 6.
Emphoropsis mucida var. ventralis Michener, 1936. Amer. Mus. Novitates 876: 3. °.

Taxonomy: Cockerell, 1907. Colo. Univ., Studies 4: 256. 6. —Michener, 1936. Amer. Mus.
Novitates 876: 3. 2, 6.
dammersi Timberlake. Calif. Pollen: Unknown, but visits flowers of Arctostaphylos including
A. glauca, A. pungens.
Emphoropsis dammersi Timberlake, 1937. Amer. Mus. Novitates 958: 2. 2, ¢.
depressa (Fowler). Calif. Pollen: Unknown, but visits flowers of Amsinckia intermedia,
Arbutus menziesii, Arctostaphylos glauca, Berberis californica, Brassica, Ceanothus

— Superfamily APOIDEA 2169
soredatus, Eriodictyon californicum, Lupinus albifrons, Phacelia, Ribes indecorum,
Salvia columbariae, S. mellifera, S. sonomensis, Vicia.

Habropoda depressa Fowler, 1899. Canad. Ent. 31: 288. 2, ¢.

Biology: Fowler, 1899. Canad. Ent. 31: 283 (nest, adult habits).
excellens Timberlake. Calif. (Kern Co.). Pollen: Unknown, but visits flowers of
Chrysothamnus.
Emphoropsis excellens Timberlake, 1962. Ent. News 73: 36. 2, 6.
interspersa Cockerell. Calif. Pollen: Unknown, but visits flowers of Amsinckia douglasiana, A.

intermedia, Astragalus, Salvia mellifera.
Emphoropsis interspersa Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 99. 6.

Taxonomy: Cockerell, 1924. Pan-Pacifie Ent. 1: 50.
laboriosa fedorensis Cockerell. Tex.
Emphoropsis floridana var. fedorensis Cockerell, 1905. Canad. Ent. 37: 265. 6, 9.
laboriosa laboriosa (Fabricius). Ill. to New England, south to Miss. and Fla. Pollen: Unknown,
but visits flowers of Cercis, Cirsium, Dicentra, Gelsemium sempervirens, Lupinus,
Malus, Melilotus, Prunus, Vaccinium, Vicia.
Bombus laboriosus Fabricius, 1804. Systema Piezatorum, p. 352.
Anthophora floridana Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 389. 2, d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 332-338, fig. 95
(redescription, synonymy).

Biology: Ornduff, 1970. Arnold Arboretum, Jour. 51: 1-17 (floral visitation).
miserabilis (Cresson). Wash., Oreg. Calif.; Mexico (Baja Calif.). Pollen: Stores pollen of
Abronia, Convolvulus, Lupinus, but visits flowers of Arctostaphylos, Cirsium,

Erysimum, Lantana, Lotus, Lupinus, Phacelia, Raphanus for nectar.
Anthophora miserabilis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 191. d.

Taxonomy: Torchio and Stephen, 1961. Ent. Soc. Amer., Ann. 54: 683-687, 7 figs. (larva and
pupa).
Biology: Stephen and Torchio, 1961. Ent. Soc. Amer., Ann. 54: 687-692, 2 figs. (nest, life
history).
morrisoni (Cresson). Colo.
Anthophora morrisoni Cresson, 1878. Acad. Nat. Sci. Phila., Proc. 30: 192. ¢.
murihirta Cockerell. B. C., Calif.
Emphoropsis murthirta Cockerell, 1905. Ent. News 16: 81. d.

murina Cockerell. Calif.
Emphoropsis murthirta murina Cockerell, 1909. U. S. Natl. Mus., Proc. 36: 414. ¢.

Taxonomy: Cockerell, 1924. Pan-Pacific Ent. 1: 49. d, 9.
pallida Timberlake. South. Calif., Ariz., Utah, deserts. Parasite: Lytta magister Horn, Zonitis
(Neozonitis) n. sp. Pollen: Apparently an oligolege of Larrea tridentata, but visits
flowers of other plants including Compositae, Lupinus, Camissonia, Phacelia Crenulata

for nectar and sometimes for pollen.
Emphoropisis pallida Timberlake, 1937. Amer. Mus. Novitates 958: 1. 2, d.

Biology: Bohart, Torchio, Maeta and Rust, 1972. Kans. Ent. Soc., Jour. 45: 381-392, 14 figs.
(life history). —Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 44 (pollen
relationships).

pascoensis (Cockerell). Wash.
Habropoda floridana var. pascoensis Cockerell, 1898. Acad. Nat. Sci. Phila., Proc. 50: 54.
2
rugosissima Cockerell. Calif., Nev.; Mexico (Baja California). Pollen: Collects pollen from
flowers of Camissonia clavaeformis, C. kernensis, but also visits flowers of Abronia
maritima, Amsinckia intermedia, Lupinus bicolor, L. nanus, Lotus scoparius, Phacelia
distans, Salvia mellifera, Stanleya pinnata.
Emphoropsis rugosissima Cockerell, 1905. Biol. Soc. Wash., Proc. 18: 182. 9.

2170 Hymenoptera in America North of Mexico

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 46 (floral
relationships).
salviae Michener. Calif. Pollen: Unknown, but visits flowers of Salvia.
Emphoropsis salviae Michener, 1936. Brooklyn Ent. Soc., Bul. 31: 95. d.
salviarum (Cockerell). N. Mex., Ariz. Pollen: Unknown, but visits flowers of Salvia.
Habropoda salviarum Cockerell, 1898. N. Mex. Agr. Expt. Sta., Bul. 24: 42. 6, °.
semifulva Cockerell. Calif.
Emphoropsis semifulva Cockerell, 1905. South. Calif. Acad. Sci., Bul. 4: 99. 6.
tristissima Cockerell. Calif. Pollen: Unknown, but visits flowers of Amsinckia intermedia,
Arctostaphylos patula, Collinsia bicolor, Lupinus bicolor, L. nanus, Phacelia distans,
Salvia sonomensis.
Emphoropsis infernalis tristissima Cockerell, 1904. South. Calif. Acad. Sci, Bul. 3: 161. °.

vierecki Cockerell. Colo., N. Mex.
Emphoropsis vierecki Cockerell, 1909. U. S. Natl. Mus., Proc. 36: 414. 2.

TRIBE MELECTINI

All the members of this tribe are cleptoparasites in the nests of other anthophorid bees. The
Melectini occur on all the continents except Australia.

Revision: Linsley, 1939. Ent. Soc. Amer., Ann. 32: 429-468, 9 figs. (Nearctic spp.). —Hurd and
Linsley, 1951. Calif. Ins. Survey, Bul. 1: 119-140, 1 pl., 5 maps (U. S. spp.). —Lieftinck,
1972. Tijdschr. Ent. 115: 253-324, 57 figs., 2 pls., 2 maps, 1 table (generic revision of
smaller genera of Old World melectines, distribution and host relationships, comments on
N. Amer. genera).

Taxonomy: Hurd, 1953. Kans. Ent. Soc., Jour. 26: 35-37, 1 map (generic distribution in
Americas). —Rozen, 1969. Amer. Mus. Novitates 2382: 1-24, 56 figs. (larva).

Genus MELECTA Latreille
Genus MELECTA Subgenus MELECTA Latreille

Melecta Latreille, 1802. Hist. Nat. Fourmis, p. 427.
Type-species: Apis albifrons Forster. Desig. by Latreille, 1810 (=Apis punctata
Fabricius), see Day and Fitton, 1977. Biol. Jour. Linn. Soe. 9: 40.
Symmorpha Klug, 1807. Mag. f. Insektenk. 6: 198, 227.
Type-species: Apis albifrons Forster. Monotypic (=Apis punctata Fabricius).
Bombomelecta Patton, 1879. U. S. Geol. and Geog. Survey, Bul., p. 370.
Type-species: Melecta thoracica Cresson. Monotypic.
bohartorum Linsley. Calif., Nev.
Melecta (Melecta) bohartorum Linsley, 1939. Ent. Soc. Amer., Ann. 32: 442. 9°.
pacifica atlantica Linsley. Kans. and Nebr., to N. J., south to Ga. and Ala. Host: Anthophora
ursina ursina Cress.?
Melecta (Melecta) atlantica Linsley, 1943. N. Y. Ent. Soc., Jour. 51: 225. @.

Taxonomy: Michener, 1948. Ent. Soc. Wash., Proc. 50: 17. 2, ¢. —Michener, 1954. Kans. Ent.
Soc., Jour. 27: 66. —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 492-493, fig. 117
(redescription).

pacifica fulvida Cresson. Wash., Oreg. northeast. Calif., Idaho, Wyo., Colo., Utah, Nev., Ariz.,
N. Mex., Tex. Host: Anthophora pacifica infernalis (Dalla Torre), Emphoropsis
cineraria Smith.
Melecta fulvida Cresson, 1878. Amer. Ent. Soc., Trans. 7: 204. °.
Bombomelecta semifulva Cockerell, 1921. Ann. and Mag. Nat. Hist. (9) 7: 212. ¢.

Taxonomy: Michener, 1948. Ent. Soc. Wash., Proc. 50: 17. ¢. —Hurd and Linsley, 1951. Calif.
Ins. Survey, Bul. 1: 128, map 4.

Biology: Torchio and Youssef, 1968. Kans. Ent. Soc., Jour. 41: 30 (induced parasitization; as
M. pacifica).

Superfamily APOIDEA 2171

pacifica pacifica Cresson. Calif.
Melecta pacifica Cresson, 1878. Amer. Ent. Soc., Trans. 7: 91. 3.
Taxonomy: Hurd and Linsley, 1951. Calif. Ins. Survey, Bul. 1: 127, map 4.

separata alfredi (Cockerell). N. Mex.
Bombomelecta alfredi Cockerell, 1895. Psyche 7 (sup.): 11. d.

separata arizonica (Cockerell). Ariz. Host: Anthophora forbesii Ckll.
Bombomelecta arizonica Cockerell, 1902. Canad. Ent. 34: 267. 2.
Biology: Lutz and Cockerell, 1920. Amer. Mus. Nat. Hist., Bul. 42: 574 (host).
separata callura (Cockerell). B. C., to Calif., Idaho, Utah, Colo., Ariz., N. Mex. Host:
Anthophora edwardsii Cress. Parasite: Sphaeropthalma unicolor (Cress.).
Bombomelecta callura Cockerell, 1926. Pan-Pacifie Ent. 3: 58. 3.
Taxonomy: Rozen, 1969. Amer. Mus. Novitates 2382: 8-10, figs. 2-8 (larva).

Biology: Thorp, 1969. Amer. Midland Nat. 82: 330-331 (host). —Thorp, 1969. Amer. Midland
Nat. 82: 338-345, 7 figs. (behavior, ecology).
separata johnsoni (Cockerell). Colo.
Bombomelecta johnsoni Cockerell, 1905. Ent. News 16: 270. 6.
separata mojavensis Linsley. Calif. (Mojave Desert).
Melecta (Melecta) separata mojavensis Linsley, 1939. Ent. Soc. Amer., Ann. 32: 446. 2, d.
separata separata Cresson. Calif., Oreg., Wash., Idaho.
Melecta separata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 204. 2.
thoracica Cresson. Wash., Oreg., Calif. (Sierra Nevada Mts.), Nev., Utah, Colo., Wyo., Nebr., S.
Dak., Kans. Host: Emphoropsis cineraria (Sm.).
Melecta thoracica Cresson, 1875. Rpt. Geog. Geol. Expl. and Survey west of 100th
Meridian, v. 5, p. 726.
Melecta (Melecta) sierrae Linsley, 1939. Ent. Soe. Amer., Ann. 32: 437. 2, ¢.
Taxonomy: Linsley, 1945. Ent. News 56: 151, fig. 1. 2. —Hurd and Linsley, 1951. Calif. Ins.
Survey, Bul. 1: 126, map. 2.

Biology: Linsley, 1943. Pan-Pacific Ent. 19: 160 (habits, as M. sierrae).

Genus MELECTA Subgenus MELECTOMIMUS Linsley

Melecta subg. Melectomimus Linsley, 1939. Ent. Soc. Amer., Ann. 32: 448.
Type-species: Melecta edwardsii Cresson. Monotypic and orig. desig.

edwardsii Cresson. Calif., Nev.; Mexico (Baja California). Host: ?7E mphoropsis sp.
Melecta edwardsii Cresson, 1878. Amer. Ent. Soc., Trans. 7: 92. ¢.
Bombomelecta zygos Viereck, 1903. Amer. Ent. Soc., Trans. 29: 179. °.

Genus XEROMELECTA Linsley
Another subgenus, Nesomelecta, is found in the West Indies.
Genus XEROMELECTA Subgenus XEROMELECTA Linsley

Melecta subg. Xeromelecta Linsley, 1939. Ent. Soc. Amer., Ann. 32: 450.
Type-species: Bombomelecta larreae Cockerell. Monotypic and orig. desig.
larreae (Cockerell). N. Mex. to Nev., south. Calif.
Bombomelecta larreae Cockerell, 1900. Canad. Ent. 32: 361. °.
Bombomelecta azygos Viereck, 1903. Amer. Ent. Soc., Trans. 29: 181. ¢.

Genus XEROMELECTA Subgenus MELECTOMORPHA Linsley

Melecta subg. Melectomorpha Linsley, 1939. Ent. Soc. Amer., Ann. 32: 451.
Type-species: Melecta californica Cresson. Orig. desig.
californica (Cresson). B. C. to Calif. east to IIl., Mo., Okla., Tex.; Mexico (Baja California,
Puebla and Zacatecas). Host: Anthophora abrupta Say, A. bomboides neomexicana Ckll.,

2172 Hymenoptera in America North of Mexico

A. bomboides stanfordiana Ckll., A. edwardsii Cress., A. linsleyi Timb., A. occidentalis
Cress., A. urbana Cress. Parasite: Sphaeropthalma anthophora (Ashm.), S. wnicolor
(Cress.).

Melecta californica Cresson, 1878. Amer. Ent. Soc., Trans. 7: 91. d, 2.

Melecta miranda Fox, 1893. Ent. News 4: 148. °.
Pseudomelecta pasadenensis Cockerell, 1910. Ann. and Mag. Nat. Hist. (8) 5: 27. 2.

Melecta sladeni Viereck, 1924. Canad. Ent. 56: 15. °.

Taxonomy: Hurd and Linsley, 1951. Calif. Ins. Survey, Bul. 1: 128, map 5. —Hurd, 1953.
Kans. Ent. Soc., Jour. 26: 35-36 (Mexican distrib.). —Michener, 1953. Kans. Univ. Sci. Bul.
35: 1083, figs. 238-242 (larva). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
494-496, fig. 119 (redescript.). —Rozen, 1969. Amer. Mus. Novitates 2382: 10.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 224 (as californica miranda). —Mickel, 1928.
Ent. News 39: 73 (as californica miranda). —Linsley and MacSwain, 1942. Amer. Midland
Nat. 27: 408, f. 10. —Hobbs, Nummi and Virostek, 1961. Canad. Ent. 93: 146. —Linsley,
1962. Ent. Soc. Amer., Ann. 55: 162-168, fig. 8.

interrupta (Cresson). Minn., Wis., and IIl., south to Tex., west to Wyo., Colo. and Ariz.; Mexico
(Chihuahua, Durango and Zacatecas). Host: Anthophora abrupta Say.
Melecta interrupta Cresson, 1872. Amer. Ent. Soc., Trans. 4: 275. 2, 6.
Melecta interrupta var. fallugiae Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 23. 9.
Melecta interrupta var. rociadensis Cockerell, 1904. Ann. and Mag. Nat. Hist. (7) 14: 23. 6.

Taxonomy: Hurd and Linsley, 1951. Calif. Ins. Survey, Bul. 1: 133, map 5. —Hurd, 1953.
Kans. Ent. Soc., Jour. 26: 36 (Mexican distrib.). —Mitchell, 1962. N. C. Agr. Expt. Sta.

Tech. Bul. 152: 496-497, fig. 119 (redescription).
Genus BRACHYMELECTA Linsley

Brachymelecta Linsley, 1939. Ent. Soc. Amer., Ann. 32: 458.
Type-species: Melecta? mucida Cresson. Monotypic and orig. desig.

mucida (Cresson). Nev.
Melecta? mucida Cresson, 1878. Amer. Ent. Soce., Trans. 7: 205. 6.

Genus ZACOSMIA Ashmead

Zacosmia Ashmead, 1898. Psyche 8: 282.
Type-species: Melecta maculata Cresson. Monotypic and orig. desig.

Micromelecta Baker, 1906. Invertebrata Pacifica, v. 1, p. 143.
Type-species: Melecta maculata Cresson. Monotypic and orig. desig.
maculata desertorum Cockerell. North. Mexico to south. Calif., Ariz., Tex.
Zacosmia maculata desertorum Cockerell, 1916. Canad. Ent. 48: 391. ¢.
maculata maculata (Cresson). Alta. and Wash. to Calif., Idaho, Utah, Colo. and Wyo. Host:
Anthophora (Micranthophora) arthuri Ckll., A. (M.) curta melanops Ckll., A. (M.)
flexipes Cress., A. (M.) peritomae Ckll. Parasite: Anthrax limatulus larrea Marston.
Melecta maculata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 204. 2, 6.
Melecta (Pseudomelecta?) suffusa Viereck, 1924. Canad. Ent. 56: 15. 9. N. syn.

Taxonomy: Hurd and Linsley, 1951. Calif. Ins. Survey, Bul. 1: 122. —Hurd, 1953. Kans. Ent.
Soc., Jour. 26: 37, fig. (Mexican distrib.). —Rozen, 1969. Amer. Mus. Novitates 2382: 14-15,
figs. 24-30 (larva).

Biology: Hicks, 1934. Colo. Univ., Studies 21 (4): 265-267. —Torchio and Youssef, 1968. Kans.
Ent. Soc., Jour. 41: 289-302, 21 figs. (ecology, immature stages). —Torchio, 1971. Los
Angeles Co. Mus., Contrib. Sci. 206: 1-10, 13 figs., 1 table (ecology).

TRIBE CENTRIDINI

This is a New World group of pollen-collecting bees which are especially abundant in the trop-
ics although some species are present in the warm temparate regions of both continents. The
tribe contains two genera, Centris and Epicharis, but only the genus Centris occurs in America

north of Mexico.

Superfamily APOIDEA 2173
Taxonomy: Rozen, 1965. Amer. Mus. Novitates 2233: 13-26, figs. 35-72 (larvae).

Genus CENTRIS Fabricius

Taxonomy: Timberlake, 1940. Pan-Pacific Ent. 16: 138-141 (Calif. spp.). —Michener, 1951.
Kans. Ent. Soc., Jour. 24: 1-11 (subgenera). —Snelling, 1956. Pan-Pacifie Ent. 32: 1-8 (Calif.
spp.). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 333-388, figs. 96, 97 (eastern
U. S. spp.). —Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 112: 1-33 (nomencl. and
tax. of some N. Amer. spp.). —Snelling, 1974. Los Angeles Co. Mus., Contrib. Sci. 259: 1-41,
44 figs. (distr. and tax. of some N. Amer. spp.).

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 44-46 (intrafloral ecol. of
Larrea visiting spp.). —Alcock, Jones and Buchmann, 1976. Kans. Ent. Soc., Jour. 49:
469-474, 5 figs., 1 table (nesting behavior).

Genus CENTRIS Subgenus XEROCENTRIS Snelling

Centris subg. Xerocentris Snelling, 1974. Los Angeles Co. Mus., Contrib. Sci. 259: 3.
Type-species: Centris californica Timberlake. Orig. desig.

Taxonomy: Snelling, 1974. Los Angeles Co. Mus., Contrib. Sci. 259: 3-5 (key to N. Amer.
spp.).
californica Timberlake. Calif., Ariz.; Mexico (Baja California). Pollen: Apparently oligolectic on
Wislizenia refracta, but also visits flowers of Cleomella obtusifolia.
Centris californica Timberlake, 1940. Pan-Pacifiec Ent. 16: 139. °.
hoffmanseggiae Cockerell. N. Mex., Ariz., south Calif. and Santa Lucia Mts. in coastal central
Calif. Pollen: Polylectic, visits flowers of Astragalus, Cercidium, Dicentra chrysantha,
Larrea tridentata, Lotus scoparius, Lupinus densiflorus, Medicago sativa, Melilotus,
Penstemon antirrhinoides, Prosopis glandulosa.
Centris hoffmanseggiae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 395. 3 (9
misdet.).
Centris hoffmanseggiae davidsoni Cockerell, 1904. South. Calif. Acad. Sci., Bul. 3: 160. d.

Taxonomy: Cockerell, 1900. Canad. Ent. 32: 363. ¢, 2.
pallida Fox. Ariz., south Calif. deserts and Santa Catalina Island; Mexico (Baja California and

Sonora). Pollen: Polylectic, chiefly Acacia greggii, Cercidium torreyanum, Dalea emoryi,
D. spinosa, Larrea tridentata, Olneya tesota, Prosopis pubescens, but also visits other
flowers including Asclepias erosa, Aster spinosa, Cirsium, Parkinsonia aculeata.

Centris pallida Fox, 1899. Acad. Nat. Sci. Phila., Proc. 51: 66. 2.

Centris pallida callognatha Cockerell, 1923. Calif. Acad. Sei., Proc. (4) 12: 78. °.

Centris trichosoma Cockerell, 1928. Calif. Acad. Sci., Proc. (4) 12: 78. 6.

Biology: Alcock, Jones and Buchmann, 1976. Jour. Zool., London 179: 189-199, 3 pls., 2 figs., 2
tables (nest site, adult behavior including emergence, mating). —Alcock, Jones and
Buchmann, 1976. Kans. Ent. Soc., Jour. 49: 472-474, figs. 4, 5, table 1 (nest architecture,
nesting behavior). —Alcock, 1976. Psyche 83: 121-131, 2 figs., 1 table (social organization of
male populations).

rhodomelas Timberlake. Calif. Pollen: Unknown, but visits flowers of Psoralea macrostachya.
Centris rhodomelas Timberlake, 1940. Pan-Pacific Ent. 16: 139. 2, d.
tiburonensis Cockerell. Ariz., south. Calif.; Mexico (Baja California). Pollen: Polylectic, chiefly
Dalea including D. emoryi, D. schottii, D. spinosa, but visits other flowers for nectar
and or pollen including Acacia greggii, Asclepias, Cercidium torreyanum, Koeberlinia
spinosa, Olneya tesota.
Centris tiburonensis Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 78. 2.

Genus CENTRIS Subgenus PARACENTRIS Cameron

Paracentris Cameron, 1903. Amer. Ent. Soc., Trans. 29: 235.
Type-species: Paracentris fulvohirta Cameron. Monotypic.

Centris subg. Penthemisia Moure, 1950. Dusenia 1: 390.
Type-species: Hemisia Chilensis Spinola. Orig. desig.

Centris subg. Trichocentris Snelling, 1956. Pan-Pacifie Ent. 32: 4.

2174 Hymenoptera in America North of Mexico

Type-species: Centris rhodoleuca Cockerell. Orig. desig.

Taxonomy: Snelling, 1974. Los Angeles Co. Mus., Contrib. Sci. 259: 5-20, figs. (key to N.
Amer. spp.).
angustifrons Snelling. Ariz. (Huachuca Mts.).
Centris (Paracentris) angustifrons Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 112:
13292

aterrima Smith. South. Ariz.; Mexico.
Centris aterrima Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 378. ¢.
atripes Mocsary. Tex., N. Mex., Ariz., Calif. (Brawley); Mexico to Costa Rica. Pollen: Polylectic,
visits flowers of Dalea, Hyptis, Kallstroemia grandiflora, Koeberlinia spinosa, Larrea
tridentata, Solanum elaeagnifolium.
Centris atripes Mocsary, 1899. Termes. Fuzetek 22: 254. 6.
Centris atriventris Fox, 1899. Acad. Nat. Sci. Phila., Proc. 51: 68. 2, ¢. Preoce.
Centris foxi Friese, 1900. K. K. Naturhist. Hofmus., Ann. 15: 350. N. name.

caesalpiniae Cockerell. Tex., N. Mex., Ariz.; Mexico (Chihuahua). Pollen: Polylectic, visits
flowers of Acacia, Asclepias brachystephana, Baccharis, Cevallia sinuata, Eriogonum,
Hoffmanseggia densiflora, H. falcaria, Robinia, Solanum elaeagnifolium, S. rostratum, ,
Verbesina encelioides.
Centris caesalpiniae Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 394. °, d.
Centris morsei Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 355. ¢.
Centris marginata Fox, 1899. Acad. Nat. Sci. Phila., Proc. 51: 67. ¢.

cockerelli cockerelli Fox. Tex., N. Mex.; Mexico (Chihuahua, Nuevo Leon, Tamaulipas);
intergrades with following subspecies in western N. Mex., Chihuahua, eastern Ariz. and
Sonora. Pollen: Presumably polylectic, visits flowers of Dalea formosa, Hoffmanseggia,
Prosopis glandulosa var. torreyanum.
Centris Cockerelli Fox, 1899. Acad. Nat. Sci. Phila., Proc. 51: 68. °.

cockerelli resoluta Cockerell. Ariz., Nev., south. Calif.; Mexico (Baja California and Sonora).
Pollen: Polylectic, chiefly Cercidium torreyanum, Dalea, Krameria, Larrea tridentata,
Prosopis, but visits other flowers for nectar and possibly pollen.
Centris cockerelli resoluta Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 76. 2, 6.

Biology: Alcock, Jones and Buchmann, 1976. Kans. Ent. Soc., Jour. 49: 471-472, fig. 3, table 1
(nest architecture, nesting behavior).

lanosa Cresson. Kans., Tex., and western Fla.
Centris lanosa Cresson, 1872. Amer. Ent. Soc., Trans. 4: 284. ¢.
Centris subhyalina Fox, 1899. Acad. Nat. Sci. Phila., Proc. 51: 69. 2.
Centris birkmannii Friese, 1900. Termes. Fuzetek 238: 44. 6, 9.

mexicana Smith. Ariz., N. Mex., Tex.; Mexico (Chihuahua, D. F., Durango, Michoacan, Nuevo
Leon, and Sonora). Pollen: Unknown, but visits flowers of Cevallia sinuata,
Hoffmanseggia densiflora, Melilotus alba, Menodora scabra.
Centris mexicana Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 378. “2” = 6.

rhodopus Cockerell. Tex., N. Mex., Ariz., south. Calif.; Mexico (Baja California, Chihuahua, and

Sonora). Pollen: Polylectic, visits flowers of a wide variety of plants including Acacia
greggii, Asclepias, Bebbia juncea, Centaurea, Cercidium torreyanum, Cevallia sinuata,
Chilopsis linearis, Croton californicus, Dalea schotti, D. spinosa, Eriogonum inflatum,
Eriodictyon, Eriophyllum, Hoffmanseggia densiflora, H. falcaria, Larrea tridentata,
Lupinus, Palafoxia linearis, Prosopis glandulosa, Solanum elaeagnifolium, S.
rostratum, Solidago confinis, Tamarix, Wislizenia refracta.

Centris caesalpiniae var. rhodopus Cockerell, 1897. Ann. and Mag. Nat. Hist. (6) 19: 394.
Oem)

Centris rhodopus var. pulchrior Cockerell, 1900. Canad. Ent. 32: 363. ¢.

Centris rhodoleuca Cockerell, 1923. Calif. Acad. Sci., Proc. (4) 12: 75. ¢.

Biology: Alcock, Jones and Buchmann, 1976. Kans. Ent. Soc., Jour. 49: 469-471, figs. 1, 2,
table 1 (nest architecture, nesting behavior).

Superfamily APOIDEA 2175

zacateca Snelling. Ariz., N. Mex.; Mexico (Chihuahua, Durango, Jalisco and Zacatecas). Pollen:
Unknown, but visits flowers of Baccharis, Hoffmanseggia densiflora, Melilotus alba,
Penstemon tenutfolius.
Centris (Paracentris) zacateca Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 112: 11.

ear
Genus CENTRIS Subgenus ACRITOCENTRIS Snelling

Centris subg. Acritocentris Snelling, 1974. Los Angeles Co. Mus., Contrib. Sci. 259: 36.
Type-species: Centris (Melanocentris) ruthannae Snelling. Orig. desig.
ruthannae Snelling. Ariz. Pollen: Unknown, but visits flowers of Kallstroemia grandiflora.
Centris (Melanocentris) ruthannae Snelling, 1966. Los Angeles Co. Mus., Contrib. Sci. 112:
28, fig..1d andi. 6, °.

Genus CENTRIS Subgenus CENTRIS Fabricius

Centris Fabricius, 1804. Systema Piezatorum, p. 354. Name placed on Official List of
_Generic Names in Zool. by Internatl. Comn. Zool. Nomencl., Op. 567, 1959.
Type-species: Apis haemorrhoidalis Fabricius. Desig. by Internatl. Comn. Zool.
Nomencl., Op. 567, 1959.
Hemisia Klug, 1807. Mag. Insektenk. 6: 227.
Type-species: Apis haemorrhoidalis Fabricius. Desig. by Cockerell, 1906.
decolorata Lepeletier. Tex. (Cameron Co.); Mexico.
Centris decolorata Lepeletier, 1841. Hist. Nat. Ins., Hym. v. 2, p. 160. 6.
eiseni Fox. Ariz.; Mexico (Baja California, Morelos, Puebla, Sinaloa, and Sonora).
Centris eiseni Fox, 1894. Calif. Acad. Sci., Proc. 4: 22. °.
flavofasciata Friese. Ariz. (Nogales); Mexico (Guerrero, Morelos and Sonora), south to
Panama.
Centris flavifrons var. flavofasciata Friese, 1899. Termesz. Fuzetek 23: 46. d.
versicolor (Fabricius). South. Fla.; widespread in American tropics. Pollen: Polylectic, visits
flowers of Borrichia, Byrsonima, Carymbola, Erigenia, Ocimum, Securidaca and
Thrysallis in the United States.
Apis versicolor Fabricius, 1775. Systema Ent., p. 386.
Centris versicolor vincentana Cockerell, 1938. Entomologist 71: 282. °.

Taxonomy: Moure, 1960. Studia Ent. 3: 123-125 (synonymy).
Genus CENTRIS Subgenus HEMISIELLA Moure

Hemisiella Moure, 1945. Rev. de Ent. 16: 407.
Type-species: A pis lanipes Fabricius. Orig. desig.
confinis Perez. Ariz. (Patagonia and Tumacacori); Mexico (Chiapas, Guanajuato, Morelos,
Puebla). Pollen: Presumably polylectic, is known to collect pollen from flowers-of
Parkinsonia aculeata, but also visits these and other flowers for nectar including
Eysenharatia polystacha, Jacaranda.
Centris confinis Perez, 1905. Mus. Hist. Nat. Paris, Bul. 11: 40. °.
transversa Perez. Ariz. (Superior); Mexico (Puebla and Tehuacan). Pollen: Unknown, but
Arizona specimen was taken at flowers of tamarisk.
Centris transversa Perez, 1905. Mus. Hist. Nat. Paris, Bul. 11: 39. °, ¢.

Genus CENTRIS Subgenus UNASSIGNED

limbata Friese. Tex.
Centris limbata Friese, 1900. Termes. Fuzetek 23: 44. 2.

TRIBE CTENIOSCHELINI

This is a chiefly Neotropical group of parasitic bees and only the genera Fricrocis and
Mesoplia extend northward into the United States.

Taxonomy: Rozen, 1969. Amer. Mus. Novitates 2382: 1-24, 56 figs. (larva).

2176 Hymenoptera in America North of Mexico

Genus ERICROCIS Cresson

Ericrocis Cresson, 1887. Amer. Ent. Soc., Trans. Sup. Vol., pp. 131, 134.
Type-species: Crocisa? lata Cresson. Monotypic.

Revision: Linsley, 1939. Ent. Soc. Amer., Ann. 32: 463-468.
arizonensis Baker. South. Calif., Ariz.; Mexico. Host: Anthophora spp., Centris spp.
Ericrocis arizonensis Baker, 1906. Invertebrata Pacifica, v. 1, p. 143. ¢.
Ericrocis melectoides Baker, 1906. Invertebrata Pacifica, v. 1, p. 144. 2 (¢ misdet.).
lata (Cresson). Fla. to south. Calif.
Crocisa? lata Cresson, 1878. Amer. Ent. Soc., Trans. 7: 91. 6.

Genus MESOPLIA Lepeletier

Mesoplia Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 457.
Type-species: Mesocheira azurea Lepeletier and Serville. Monotypic.
Melissa Smith, 1854. Cat. Hym. Brit. Mus. 2: 279.
Type-species: Mesocheira azurea Lepeletier and Serville. Desig. by Sandhouse,
1943.
dugesi (Cockerell). South. Ariz.; Mexico.
Mesonychium dugesi Cockerell, 1917. Ann. and Mag. Nat. Hist. (8) 19: 477. ¢.

SUBFAMILY XYLOCOPINAE

Although these bees are found throughout much of the world they are especially well
represented by numerous species in the tropics. Most of the species are pollen-collecting bees
and, unlike most other non-parasitic anthophorids, they usually make their nests in wood of vari-
ous sorts. The subfamily contains two tribes, the Ceratinini (small carpenter bees) and the
Xylocopini (large carpenter bees) both of which are present in America north of Mexico.

TRIBE CERATININI

This is a large group of mostly small, almost hairless bees which, except for parasitic species,
make their nests in hollow, pithy or rotten stems and sometimes in vines. While there are about
a dozen genera of these bees present in the Old World including the cleptoparasitic Eucon-
dylops, Inquilina and Nasutapis, only the nearly worldwide genus Ceratina and the endemic
Chilean genus Manuelia are present in the New World.

Pithitis smaragdula (Fabr.) was intentionally introduced into the United States from Ludhi-
ana, India on April 10, 1969 at Davis, California as a potentially important pollinator of economi-
cally useful plants, especially legumes and cucurbits (Daly, Bohart and Thorp, 1971. Ent. Soc.
Amer., Ann. 64: 1145-1150). The introduction apparently was unsuccessful since no recoveries of
this species have been reported.

Genus CERATINA Latreille

Revision: Daly, 1973. Calif. Univ. Pubs. Ent. 74: 1-113, 31 figs. 2 pls., 3 tables (U. S. spp.).

Taxonomy: Smith, 1907. Amer. Ent. Soc., Trans. 33: 115-124 (N. Amer. spp.). —Sandhouse,
1935. Ent. Soc. Wash., Proc. 37: 93-95 (east. U. S. species). —Michener, 1936. Amer. Mus.
Novitates 844: 1-13 (west. U. S. spp.). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul.
152: 500-507, figs. 121-125. —Hirashima, 1971. Faculty Agr., Kyushu Univ., Jour. 16:
349-375, 20 figs. (subgeneric classification).

Biology: Malyshev, 1912. Trav. Soc. Imper. Nat. St. Petersbourg, Compt. Rend. Seances 43:

252-5, 276-7 (natural enemies). — Daly, 1967. Ent. Soc. Amer., Ann. 60: 1273-1282, 7 figs., 1
table (natural enemies).

Genus CERATINA Subgenus CERATINA Latreille

Clavicera Latreille, 1802. Hist. Nat. Fourmis, p. 433. Name suppressed by Internatl. Comn.
Zool. Nomencl., Op. 1001, 1973.

Ceratina Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 380. Name placed on Official List of
Generic Names in Zool. by Internat]. Comn. Zool. Nomencl., Op. 1001, 1973.

Superfamily APOIDEA 2177

Type-species: Apis cucurbitina Rossi. Monotypic. (=Hylaeus albilabris Fabricius).
Neoceratina Perkins, 1912. Ann. and Mag. Nat. Hist. (8) 9: 117.

Type-species: Neoceratina australensis Perkins. Monotypic.
Ceratinula Moure, 1941. Arq. Mus. Paranaense 1: 78.

Type-species: Ceratina lucidula Smith. Orig. desig.

Taxonomy: Michener, 1965. Amer. Mus. Nat. Hist., Bul. 130: 220-221 (synonymy).
arizonensis Cockerell. Tex. to Calif.; Mexico; inadvertently and successfully introduced into
the Hawaiian Islands (Oahu). Pollen: Polylectic, visits a wide variety of flowers
including Antirrhinum nutallianum, Asclepias, Baccharis sarothroides, Baeria
chrysostoma, Brassica, Calochortus venustus, Ceanothus crassifolius, Chaenactis,
Chorizanthe parryi, Cirsium, Clarkia, Cleomella obtusifolia, Condalia, Cressa, Croton
californicus, Cryptantha flaccida, C. intermedia, C. muricata, Encelia farinosa,
Ericameria parishii, Eriogonum fasciculatum, E. f. var. foliolosum, E. gracile,
Eriophyllum confertiflorum, E. lanosum, Euphorbia albomarginata, E. palmer, E.
pediculifera, Larrea tridentata, Lotus glaber, Lycium fremontii, Malacothrix
californica, Mesembryanthemum, Nolina microcarpa, Phacelia distans, P. douglasii,
Prosopis juliflora, Rhus, Salix lasiolepis, Salvia mellifera, Senecio californicus,
Solanum elaeagnifolium, Sphaeralcea laxa, Tamarix gallica.
Ceratina arizonensis Cockerell, 1898. Canad. Ent. 30: 238. do.
Ceratina arizonensis vanduzeei Cockerell, 1924. Calif. Acad. Sci. (4) 12: 543. 2.
cockerelli Smith. S. C., Ga. and Fla., west to Tex. Pollen: Apparently polylectic, visits a wide
variety of flowers including Agastache breviflora, Bidens, Callirhoe involucrata,
Citrullus vulgaris, Erigeron quercifolius, Euphorbia, Gaillardia, Heterotheca, Lippia,
Lupinus, Marilaunidium origanifolium, Melilotus alba, Monarda citriodora, Phacelia
patuliflora, Polygala, Rubus, Sabatia campestris, Sida heterophylla, Stachys floridana.
Ceratina lunata Smith, 1907. Amer. Ent. Soc., Trans. 33: 119. 6, 2. Preoce.
Ceratina cockerelli Smith, 1907. Canad. Ent. 39: 260. N. name.

Genus CERATINA Subgenus ZADONTOMERUS Ashmead

Zadontomerus Ashmead, 1899. Amer. Ent. Soc., Trans. 26: 69.
Type-species: Ceratina tejonensis Cresson. Monotypic and orig. desig.
Zaodontomerus Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 406. Emend.

SPECIES GROUP DUPLA

calcarata Robertson. Que. to Ga., west to Man., south to Tex. Ecology: Nests in borings in
sumac pith. Parasite: Aprostocetus americanus Ashm., Axima zabriskiei Howard,
Coelopencyrtus hylaei Burks, Grotea anguina Cress., Omalus iridescens (Nort.)?,
Parasierola sp., Rhydinofoenus tarsatorius (Say). Pollen: Polylectic, visits a wide
variety of flowers including Aesculus glabra sargentii, Alliaria officinalis, Amelanchier
canadensis arborea, Amorpha fruticosa, Amphiachyris dracunculoides, Antennaria,
Apocynum, Arabis, Aruncus, Aster azureus, A. laevis, A. macrophyllus, A. paniculatus,
A. saggitifolius, Barbarea vulgaris, Bicuculla cucullaria, Blephilia, Brassica, Cacalia,
Callirhoe digitata, Caltha palustris, Camassia, Capsella bursa-pastoris, Cardamine,
Ceanothus ovatus, Celastrus scandens, Cercis, Chionanthus virginicus,
Chrysanthemum leucanthemum, Chrysopsis, Cirsium, Claytonia caroliniana virginica,
Collinsia, Convolvulus sepium, Crataegus, Cypripedium, Dentaria diphylla, Deutzia
gracilis, Dirca, Echium vulgare, Ellisia, Erigenia, Erigeron canadensis, E.
philadelphicus, E. pulchellus, E. ramosus, Erythronium, Eulophus, Eupatorium
ageratoides, Euphorbia marginata, Forsythia, Fragaria virginiana, Geranium
maculatum, Geum, Gillenia Grossularia, Haplopappus, Helenium, Helianthus,
Hepatica, Heterotheca, Hydrangea paniculata, Ilex, Inula helenium, Isopyrum,
Kolkwitzia amabilis, Lepargyrea canadensis, Lepidium, Lonicera siberica, L. tatarica,
Maculara pomifera, Malus, Malva rotundifolia, Melilotus alba, M. officinalis,
Osmorrhiza, Oxalis europaea, Polemonium, Potentilla recta, Prunus americana, P.
gracilis, P. persica, P. serotina, P. tomentosa, Pulsatilla hirsutissima, Pyracantha,
Pyrus ioensis, Ranunculus, Rhus aromatica, R. canadensis, R. glabra, Ribes

2178 |§ Hymenoptera in America North of Mexico

missouriense, Robinia pseudacacia, Rubus, Rudbeckia, Salix discolor, S. nigra, Salvia,
Sambucus, Sanguinaria, Scilla campanulata, Senecio aureus, Smillacina, Solidago
canadensis, S. rigida, S. ruprestis, Spiraea vanhouttei, Taenidia, Taraxacum officinale,
T. taraxacum, Tussilago farfara, Vaccinium, Verbena, Viburnum molle, Viola
papilionacea, Waldsteinia, Zanthoxylum, Zizia aurea.

Ceratina calcarata Robertson, 1900. Acad. Sci. St. Louis, Trans. 10: 54. 6.

Biology: Rau, 1926. Acad. Sci. St. Louis, Trans. 25: 184. —Rau, 1928. Ent. Soc. Amer., Ann.
21: 380 (nest, parasite). —Krombein, 1960. Ent. News 71: 68 (nest, parasite).
dupla Say. Que. to Fla., west to Man., south to Tex., La. and Miss. Parasite: Aprostocetus
americanus Ashm., Axima zabriskiei Howard, Diomorus zabriskiei Cress., Grotea
anguina Cress., Habritys latrus Wallace, Hoplocryptus gracilis (Prov.), Merisus sp.
Pollen: Polylectic, visits a wide variety of flowers including Aesculus glabra sargentii,
Amelanchier canadensis, Ammannia, Amorpha, Anemonella, Antennaria
plantaginifolia, Anthemis cotula, Arctium, Arabis, Aruncus, Asclepias, Aster
paniculatus Barbarea vulgaris, Benzoin aestivale, Bidens mitis, B. pilosa, Blephilia,
Brassica, Brauneria, Cacalia, Camassia, Capsella bursa-pastoris, Cardamine,
Ceanothus, Cephalanthus occidentalis, Cerastrium, Chrysanthemum leucanthemum,
Chrysopsis, Circaea, Cirsium, Claytonia caroliniana, Collinsia, Convolvulus, Coreopsis,
Cornus, Crataegus punctata, Cryptotaenia, Cypripedium, Daucus carota, Delphinium,
Dentaria diphylla, D. laciniata, Dianthera, Diospyros, Dirca, Ellisia, Emilia coccinae,
Erigenia, Erigeron annuus, E. canadensis, E. philadelphicus, E. quercifolius,
Eryngium, Erythronium, Eupatorium, Flaveria linearis, Fragaria virginiana, Galactia,
Geranium maculatum, Gerardia, Gillenia, Hedeoma, Helenium, Helianthus, Heliopsis, .
Heracleum, Houstonia, Hydrangea paniculata, Hydrophyllum, Hypoxis, Isopyrum,
Krigia, Lactuca, Leonurus, Lepidium, Liatris, Lithospermum, Lobelia, Lycopus,
Lythrum, Malus, Marrubium, Medicago lupulina, Melilotus alba, M. officinalis,
Monarda, Nelumbo, Nepeta, Oenothera, Opuntia, Oxalis, Pastinaca, Penstemon,
Petalostemon, Phacelia covillei, Phyrma, Poinciana, Polemonium, Polygonatum,
Polygonium, Potentilla recta, Prunus americana, P. serotina, Prunella, Psoralea,
Ptelea, Pulsatilla, Pycnanthemum, Ranunculus, Rhannus, Rhus, Rosa, Rubus,
Rudbeckia, Sabatia, Sagittaria, Salix discolor, S. sericea, Salvia lyrata, S. pratensis,
Sambucus, Scrophularia, Scutellaria, Senecio plattensis, Sida, Silphium, Sinapis,
Smilax, Solidago, Specularia, Spiraea, Stachys floridana, Stellaria, Stokesia laevis,
Taraxacum dens-leonis, T. officinale, Teucrium, Tradescantia, Trifolium, Triosteum,
Tussilago farfara, Vaccinium corymbosum, Verbena, Verbesina, Vernonia, Veronica,
Viburnum, Viola rafinesquii, Waldsteinia, Zizia aurea. Predator: Atomosia puella
(Wied.).
Ceratina dupla Say, 1837. Boston Jour. Nat. Hist. 1: 397. 2 (¢ misdet.).
Halictus Ontariensis Provancher, 1882. Nat. Canad. 13: 203. ¢.
Ceratina dupla var. halophila Cockerell, 1911. Canad. Ent. 43: 390. °.
Ceratina dupla floridana Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 505. ¢, 9.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1063, figs. 173-179 (larva).

Biology: Angus, 1869. Amer. Nat. 2: 49. —Ashmead, 1894. Psyche 7: 25. —Comstock and
Comstock, 1895. Manual for Study of Insects, p. 669. —Packard, 1897. N. Y. Ent. Soc.,
Jour. 5: 112. —Graenicher, 1905. Ent. News 16: 43. —Graenicher, 1905. Wis. Nat. Hist.
Soc., Bul. 3: 158. —Hicks, 1926. Colo. Univ., Studies 15: 250.

strenua Smith. N. Y. to Ga., west to Wis., Mo., Kans., Okla. and Tex. Parasite: Axima
zabriskiei Howard, Grotea anguina Cress. Pollen: Polylectic, visits a wide variety of
flowers including Amorpha fruticosa, Arctium minus, Asclepias tuberosa, Aster,
Brauneria purpurea, Callirhoe digitata, Ceanothus ovatus, Cercis canadensis,
Chrysanthemum, Chrysopsis, Claytonia virginica, Coreopsis grandiflora, Crataegus,
Daucus carota, Erigeron philadelphicus, Euphorbia, Fragaria virginiana, Geranium
maculatum, Helenium, Helianthus, Heterotheca, Hydrangea arborescens, Kolkwitzia
amabilis, Leucanthemum, Marrubium vulgare, Melilotus albus, M. officinalis,
Oenothera, Oxalis, Phacelia, Potentilla pumila, P. canadensis, Prunus, Psoralea
floribunda, Ptelea trifoliata, Pyracantha, Rhus aromatica, R. canadensis, R. glabra,

Superfamily APOIDEA ~—_ 2179

Robinia pseudoacacia, Rubus nigrobaccus, Salix nigra, Senecio, Silphium laciniatum,
Solidago, Taraxacum dens-leonis, Vaccinium, Verbena urticifolia, Vernonia baldwinii,
Viola sororia, Vitex agnus-castus, Waldsteinia, Zizia aurea.

Ceratina strenua Smith, 1879. Descr. New Species Hym. Brit. Mus., p. 97. 3.

Ceratina metallica Smith, 1907. Amer. Ent. Soc., Trans. 33: 121. ¢.

Biology: Krombein, 1960. Ent. News 71: 68 (nest). —Gordh and Barrows, 1976. Kans. Ent.
Soc., Jour. 49: 344-345, 1 fig. (phoresitized female, as metallica).
texana Daly. Tex. Pollen: Presumably polylectic, visits flowers of Agastache breviflora,
Callirhoe involucrata, Lippia, Monarda citriodora, Opuntia, Phacelia patuliflora,
Rubus.
Ceratina texana Daly, 1973. Calif. Univ. Pubs. Ent. 74: 100, figs. 9c, 131, 26, 281. 2, 3.

SPECIES GROUP NANULA

apacheorum Daly. N. Mex., Ariz., Calif. and Nev. Pollen: Apparently polylectic, visits a wide
variety of flowers including Acacia greggii, Aster, Baccharis glutinosa, B. sarothroides,
Baileya multiradiata, Cassia leptocarpa, Cercidium floridum, Chaenactis stevioides,
Chilopsis linearis, Condalia lycioides, Croton, Cryptantha intermedia, Encelia farinosa,
Eriodictyon crassifolium, Gutierrezia lucida, Haplopappus cooperi, Heliotropium
curassavicum, Heterotheca, Larrea tridentata, Medicago sativa, Melilotus, Mimosa
biunciflora, Opuntia, Penstemon centranthifolius, P. parryi, Prosopis juliflora,
Rhamnus crocea, Salix, Senecio monoensis, Sphaeralcea emoryi, S. laxa, S. pedata,
Tamarix gallica,Werbesina, Wislizenia refracta.

Ceratina apacheorum Daly, 1973. Calif. Univ. Pubs. Ent. 74: 36, figs. 13i, 25, 28d, 30b. 9,

3.

Biology: Hurd and Linsley, 1975. Smithsn. Contrib. Zool. 193: 46 (floral ecology).

hurdi Daly. Calif. (Jamesburg and Pinnacles Natl. Mon.).
Ceratina hurdi Daly, 1973. Calif. Univ. Pubs. Ent. 74: 58, figs. 13d, 28c, 30a, 3le, 3li. 2, 3.

nanula Cockerell. B. C. to Calif., Idaho, Mont., south to Ariz. and N. Mex.; Mexico (northern).
Parasite: A prostocetus americanus Ashm., Coelopencyrtus hyleoleter Burks?, Eurytoma
apiculae Bugbee, Grotea californica Cress. Pollen: Polylectic, visits a wide variety of
flowers including Achillea borealis arenicola, Allium acuminatum, A phyllon
multiflorum, Aragallus lambertti, Arctostaphylos, Argemone platyceras, Asclepias
fascicularis, A. tuberosa, Aster adscendens, Astragalus goniatus, Baccharis glutinosa,
B. pilularis, B. sarothroides, Berberis, Besseya plantagenea, Brassica, Calochortus
lutea, C. paludicola, Camissonia clavaeformis, Ceanothus, Centarium davyi, Centaurea
melitensis, Chrysopsis, Chrysothamnus nauseosus speciosus, C. viscidiflorus, C. v.
typicus, Chorizanthe cuspidata, C. staticoides, Cirsium arvense, C. vulgare, Clarkia
amoena huntiana, C. biloba, C. rubicunda, Cleome lutea, C. serrulata, Collinsia
heterophylla, Convolvulus occidentalis, Corethrogyne virgata, Cowania stansburyana,
Cryptantha intermedia, Dalea greggii, Descuraina sophia, Diplaucus auriantiacus,
Encelia californica, E. farinosa, Epilobium, Erigeron argentatus, E. divergens, E.
foliosus, E. inornatus, E. miser, E. stenophyllus, E. strigosus, Eriodictyon californicus,
Eriogonum fasciculatum, E. heermannii, E. nudum, E. vimineum, Eriophyllum
confertiflorum, Eschscholzia californica, Euphorbia albomarginata, Gilia altissima, G.
exilis, Gnaphalium californicum, Grindelia camporum, G. latifolia, Helenium
puberulum, Helianthella californica, Helianthus petiolaris, Heliotropium curassavicum,
Hesperochiron californicus, Heterotheca grandiflora, Hieracium albiflorum, Hulsea
callicarpha, Iris, Kerria japonica, Lasthenia chrysostoma, Lepechinia calycina,
Linanthus aureus, Lomatium dissectum, Lotus nevadensis, L. scoparius, L. strigosus
var. hirtellus, Lupinus arboreus, Malacothrix saxatilis, Melilotus alba, M. officinalis,
Mentzelia laevicaulis, Mimulus guttatus, Monardella villosa, Montia perfoliata,
Navarretia heterodoxa, Nemophila integrifolia, Oenothera ovata, Opuntia basalaris,
Penstemon centranthifolius, P. floridus, P. grinnellii, Phacelia congesta, P. distans, P.
fremontii, P. minor, P. ramosissima, Physaria, Plantago lanceolata, Plectritis ciliosa,
Prunus subcordata, Ranunculus californicus, Rhamnus crocea, Salix, Salvia mellifera,
Sapindus saponaria, Scrophularia californica, Senecio longilobus, Solidago,

2180 Hymenoptera in America North of Mexico

Sphaeralcea ambigua, Stephanomeria cichoriacea, S. virgata, Thelypodium
linearifolium, Trichostema, Trifolium microcephalum, Verbena lasiostachys, V.
prostrata, Viola nuttallti, Wedelia incarnata, Wyethia.

Ceratina nanula Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 352. ¢.

Ceratina nanula rigdenae Michener, 1936. Amer. Mus. Novitates 844: 3. 2, 6.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 251 (nest).
shinnersi Daly. Tex. Pollen: Apparently polylectic, visits flowers of Amblyolepis setigera,

Amphiachyris, Argemone intermedia, A. mexicana, Aster exilis, Astragalus, Callirhoe
mvolucrata, Cercis canadensis, Chamaesaracha conioides, Chrysanthemum maximum,
Chrysopsis pilosa, Citrullis vulgaris, Coreopsis, Croptilon divaricatum var.
divaricatum, Cucumis melo, Erigeron strigosus, Eupatorium serotinum, Helianthus
annuus, Heterotheca subaxillaris, Leucophyllum frutescens, Marilaunidium
organifolium, Marrubium vulgare, Melilotus alba, Mentha spicata, Monarda,
Neomphila phacelioides, Opuntia Macrorhiza, Phacelia, Phyla incisa, Prosopis
glandulosa, Pyrrhopappus geiseri, Rubus trivialis, Salix, Scutellaria, Sophora affinis,
Sphaeralcea angustifolia, Tamarix gallica, Tetragonotheca ludoviciana, Tetraneurus
linearifolia, T. scaposa, Verbesina encelioides, Vicia ludoviciana, Vitex agnus-castus.

Ceratina shinnersi Daly, 1973. Calif. Univ. Pubs. Ent. 74: 90, figs. 13f, 21, 22m-N, 28b. °,
3.

SPECIES GROUP PACIFICA

pacifica Smith. B. C. to Calif., Idaho and Wyo., south to Ariz. Parasite: Aprostocetus

americanus Ashm.?, Rhydinofoenus kaweahensis Bradley. Pollen: Apparently polylectic,
visits flowers of Agave nevadensis, Arctostaphylos, Asclepias, Aster andersonii,
Balsamorhiza sagittaria, Calochortus luteus, C. macrocarpus, C. nuttallii, Ceanothus
fendleri, Centaurea melitensis, Chaenactis, Chrysothamnus, Cirsium arvense, C.
vulgare, Convolvulus occidentalis, Clematis linguisticifolia, Cleome lutea,
Corethrogyne, Cryptantha intermedia, C. lepida, C. racemosa, Encelia farinosa,
Erigeron Eriodictyon trichocalyx var. lanatum, Eriogonum heermannii, Grindelia
camporum, G. squarrosa, Haplopappus linearifolius, Helianthus annuus, H. petiolaris,
Lomatium dissectum, Malacothrix tenuifolia, Navarretia heterodoxa, Medicago sativa,
Mentzelia, Opuntia basilaris, Penstemon leonardi, Phacelia linearis, P. tanacetifolia,
Rubus leucodermis, Senecio, Sisymbrium altissimum, Solidago petradoria, Sphaeralcea
ambiguua, S. angustifolia, Stanleya pinnata, Stephanomeria virgata, Trifolium,
Verbena lasiostachys, Viguiera nevadensis, Wyethia glabra.

Ceratina pacifica Smith, 1907. Amer. Ent. Soc., Trans. 33: 128. 2.

Ceratina subpunctigena Michener, 1936. Amer. Mus. Novitates 844: 6. 2, d.

Ceratina utahensis Michener, 1936. Amer. Mus. Novitates 844: 7. °.

punctigena Cockerell. Calif. Parasite: Aprostocetus americanus Ashm.?, Coelopencyrtus
hyleoleter Burks, Eurytoma apiculae Bugbee, Grotea californica Cress. Pollen:
Apparently polylectic, visits flowers of Agoseris glauca, Aplopappus parishii,
Arctostaphylos patula, Asclepias erosa, Calendula, Calochortus, Ceanothus, Chaenactis,
Cirsium californicum, Clarkia dudleyana, Convolvulus occidentalis, Coreopsis
lanceolata, Corethrogyne virgata, Croton californicus, Cryptantha intermedia,
Cucurbita, Encelia farinosa, Eriodictyon trichocalyx var. lanatum, Eriophyllum
tridactylum, Gilia, Hazardia squarrosa, Helianthus gracilentus, H. occidentalis, Hulsea
callicarpha, Malacothrix saxatilis, M. s. tenuifolia, Penstemon incertus, P. spectabilis,
Phacelia, Rhus trilobata, Salvia mellifera, Scrophularia californica, Senecio douglasii,
Sphaeralcea, Stenotopsis linearifolius, Stephanomeria virgata. Predator: Oxyopes
scalaris Hentz.

Ceratina neomexicana punctigena Cockerell, 1916. Pomona Jour. Ent. Zool. 8: 62. 9.

Taxonomy: Michener, 1936. Amer. Mus. Novitates 844: 5. 2, 6.

Biology: Linsley, MacSwain and Raven, 1963. Calif. Univ. Pubs. Ent. 33: 47 (floral
relationships). —Bugbee, 1966. Pan-Pacific Ent. 42: 211 (natural enemies).

Superfamily APOIDEA 2181

SPECIES GROUP TEJONENSIS

micheneri Daly. Calif., Oreg. Pollen: Apparently polylectic, visits flowers of Brassica
campestris, Clarkia amoena, Convolvulus occidentalis, Corethrogyne, Cryptantha
intermedia, Encelia californica, Grindelia robusta, Helianthus, Hypochoeris radicata,
Potentilla, Rosmarinus officinalis, Salvia stachyoidea, Vaccinium ovatum.
Ceratina micheneri Daly, 1973. Calif. Univ. Pubs. Ent. 74: 59, figs. 6a, 8a, 9g, lla, 11i, 12h,
20an2otee Gr

tejonensis Cresson. Calif., Oreg. Parasite: Grotea californica Cress. Pollen: Apparently

polylectic, visits flowers of Amsinckia, Arctostaphylos drupacea, Barbarea orthoceras,
Brassica, Brodiaea coronaria, B. grandiflora, B. laxa, Ceanothus crassifolius, Cirsium,
Clarkia rhomboidea, Convolvulus occidentalis, Cucurbita, Diplacus aurantiacus,
Epigonia, Eriodictyon, Helianthus gracilentus, Lepechinia calycina, Oenothera hooker,
Paeonia brownii, Penstemon grinnellii, Phacelia distans, Prunus amygdalus, Rubus
ursinus, Salix, Salvia mellifera, Scrophularia californica.

Ceratina tejonensis Cresson, 1864. Ent. Soc. Phila., Proc. 2: 390. ¢.

Ceratina gigantea Smith, 1907. Amer. Ent. Soc., Trans. 33: 123. 9.

SPECIES GROUP UNASSIGNED

acantha Provancher. B. C. to Calif., Idaho, Nev. and Utah; Mexico (Baja California). Parasite:
Aprostocetus americanus Ashm., Coelopencyrtus hyleoleter Burks, Grotea californica
Cress. Pollen: Polylectic, visits a great variety of flowers including Achillea millefolium,
Ageratum, Allium dichamydium, Anaphalis, Anthemis cotula, Anthriscus scandicina,
Aralia californica, Arctostaphylos, Asclepias mexicana, Aster yosemitanus, Astragalus
bolanderi, Baccharis douglasii, B. pilularis, Barbaraea orthoceras, Bidens, Brassica
adpressa, B. alba, B. campestris, B. geniculata, B. nigra, Brodiaea, Calochortus albus,
C. luteus, Canna, Carduus tenuifolius, Ceanothus crassifolius, C. parvifolius, C.
soredatus, Centaurea melitensis, C. solstitialis, Centaurium davyi, Chaenactis
glabriuscula, Chaemabatia foliolosa, Chrysanthemum maximum, Chrysothamnus,
Cicuta douglasii, Cirsium arvense, C. lanceolatum, C. vulgare, Clarkia amoena
huntiana, C. rhomboidea, C. rubicunda, Claytonia, Clematis, Collinsia heterophylla,
Convolvulus arvensis, C. occidentalis, Coreopsis maritima, Corethrogyne virgata,
Cosmos, Cotoneaster, Cotula, Crepis vesicaria taraxacifolia, C. virens, Croton
californicus, Cryptantha flaccida, C. intermedia, Daucus carota, Deinandra fasciculata,
Diplacus auriantiacus, D. longiflorus, Dudleya, Echium giganteum, Encelia
californica, Epilobium, Erigeron stenophyllus, Eriogonum fasciculatum, E. latifolium
nudum, E. vimineum, Eriodictyon californicum, Eriophyllum confertiflorum, E.
lanatum, E. multicaule, E. staechadifolium, Eryngium aristatulum, Eschscholzia
californicum, Foeniculum vulgare, Fragaria californica, Gaillardia, Galium, Geranium
dissectum, G. molle, Gilia achilleaefolia, G. capitata, G. multicaulis, Glycyrrhiza
lepidota, Gnaphalium californicum, Grindelia camporum, G. robusta latifolia,
Haplopappus palmeri, Helenium bigelovii, H. puberulum, Helianthus gracilentus,
Heliotropium curassavicum, Hemizonia fasciculata, H eracleum lanatum, Heteromeles
arbutifolia, Heterotheca grandiflora, Holodiscus discolor, Hypericum, Hypochoeris
radiata, Lactuca, Lasthenia chrysostoma, L. gracilis, Layia platyglossa, Lepechinia
calycina, Lepidium, Linanthus parviflorus, Lomatium dissectum, Lonicera hispidula,
Lotus corniculatus, L. glaber, L. scoparius Madia, Malacothamnus arcuatus,
Malacothriz saxatilis, Malvastrum fasciculatum, M. thurberi, Marrubium vulgare,
Melilotus alba, Mentha spicata, Mesembryanthemum crystallinum, Micrampelis,
Monardella lanceolata, M. villosa, Montia perfoliata, Myosotis laxa, Nasturtium,
Navarretia heterodoxa Nemophila exilis, N. heterophylla, Oenothera cheiranthifolia,
Onkerria japonica, Opuntia, Paeonia brownii, Penstemon spectabilis, Perideridia
gairdneri, Phacelia distans, Physocarpus capitatus, Plagiobothrys nothofulvus,
Polygonum auberti, Potentilla, Prunella vulgaris, Prunus ilicifolia, P. subcordata,
Radicula nasturtium-aquaticum, Ranunculus californicus, Rhamnus californica,
Ribes, Rosa, Rubus parviflorus, R. ursinus, Salix argophylla, S. lasiolepis, Salvia
leucophylla, S. mellifera, Sambucus mexicana, Satureja douglasii, Scrophularia
californica, Sedum, Senecio platylobus, Sidalcea malvaeflora, Silybum marianum,

2182 Hymenoptera in America North of Mexico

Sisymbrium altissimum, Sisyrinchium, Solidago, Sonchus arvensis, Sphacele calycina,
Spiraea sorbifolia, Stachys bullata, Stellaria media, Stephanomeria virgata, Syringa,
Taraxacum vulgare, Trifolium involucratum, ft microcephalum, Venegasia
carpesioides, Verbena lasiostachys, V. prostrata, Vicia villosa, Vitis, Wyethia
amplexicaulis glabra. Predator: Cymatodera ovipennis Say.

Ceratina acantha Provancher, 1895. Nat. Canad. 22: 190. 2.

Ceratina submaritima Cockerell, 1897. Acad. Nat. Sci. Phila., Proc. 49: 352. d, 9.

Ceratina submaritima ehrhorni Cockerell, 1903. Ann. and Mag. Nat. Hist. (7) 12: 453. 6.

Biology: MacSwain, 1945. Pan-Pacific Ent. 21: 97 (natural enemies). —Daly, 1966. Ent. Soc.
Amer., Ann. 59: 1138 (nest).
diodonta Smith. Tex. Pollen: Unknown, but visits flowers of Agastache breviflora, Lippia,
Monarda citriodora.
Ceratina diodonta Smith, 1907. Amer. Ent. Soc., Trans. 33: 121. 6, °.
melanoptera Cockerell. Ariz., N. Mex., Tex. (El Paso); Mexico (Sonora). Pollen: Unknown, but
visits flowers of Argemone, Cassia leptocarpa, Dalea greggii, Encelia farinosa,
Eriogonum, Haplopappus gracilis, Proboscidea parviflora, Salix taxifolia, Wedelia
incarnata.
Ceratina melanoptera Cockerell, 1924. Calif. Acad. Sci., Proc. (4) 12: 543. 3, 2.
neomexicana Cockerell. Idaho south to Calif., Nev., Ariz., N. Mex. and Tex. Pollen: Polylectic,
visits a wide variety of flowers including Agave, Amorpha fruticosa, Anemone
intermedia, Aquilegia, Artemisia, Aster, Astragalus goniatus, Berberis trifoliata,
Carduus Ceanothus, Chaenactis glabriuscula, Chrysopsis, Chrysothamnus nauseosus,
Erigeron canus, E. inornatus, Gilia calcarea, Grindelia perennis, Haplopappus
bloomeri angustatus, Helianthus petiolaris, Iris, Ligustrum vulgare, Malva cockerelli,
Melilotus officinalis, Monarda, Nothocalais cuspidata, Oenothera, Opuntia erinacea,
Penstemon gracilis, Phacelia, Physaria, Platycodon grandiflorum, Potentilla, Prunus,
Pulsatilla hirsutissima, Rubus, Salix bebbiana, Senecio longilobus, Silphium, Solidago,
Taraxacum officinale, Tetraneuris, Viola nuttallit.
Ceratina neomexicana Cockerell, 1901. Ent. News 12: 43. °.

Taxonomy: Cockerell, 1906. Canad. Ent. 38: 165. ¢.

Biology: Hicks, 1926. Colo. Univ., Studies 15: 251 (nest).

sequoiae Michener. Wash., Idaho, Oreg. and Calif. Parasite: Aprostocetus americanus Ashm.
Pollen: Oligolectic, obtains pollen from flowers of the genus Clarkia including C. biloba,
C. b. australis, C. cylindrica, C. dudleyana, C. gracilis albicaulis, C. purpurea, C.
rhomboidea, C. unguiculata, C. williamsonu, C. xantiana, C. viminea, but also visits
other flowers for nectar such as Amsinckia, Asclepias mexicana, Cryptantha,
Epilobium, Eriogonum, Gilia tricolor, Hesperochiron, Layia platyglossa, Lepechina
calycina, Lessingia leptoclada, Lotus glaber, Madia, Mimulus, Nemophila, Oenothera,
Penstemon, Phacelia, Ranunculus, Salvia, Sambucus, Scrophularia californica,
Senecio, Stachys ajugoides, Trichostema laxum, Verbena lasiostachys. Predator:
Apiomerus crassipes (Fabr.).

Ceratina sequoiae Michener, 1936. Amer. Mus. Novitates 844: 1. °.

Biology: MacSwain, Raven and Thorp, 1973. Calif. Univ. Pubs. Ent. 70: 34-37 (floral
relationships).
timberlakei Daly. Oreg., Calif. Pollen: Apparently polylectic, visits flowers of Arctostaphylos,
Asclepias mexicana, Calochortus luteus, Clarkia, Cucurbita, Eriodictyon californicum,
Gilia capitata, Haplopappus arborescens, Lotus glaber, Prunus subcordata, Salvia,

Senecio douglasti, Sidalcea calycosa.
Ceratina timberlakei Daly, 1973. Calif. Univ. Pubs. Ent. 74: 102, figs. 7c, 10i, 12a, 27a, 28e,
Staple lene no

Genus CERATINA Subgenus EUCERATINA Hirashima, Moure and Daly

Ceratina subg. Euceratina Hirashima, Moure and Daly, Jn Hirashima, 1971. Faculty Agr.
Kyushu Univ., Jour. 16: 369.
Type-species: Apis callosa Fabricius. Orig. desig.

Superfamily APOIDEA = 2183

The majority of the species included in this subgenus occur in Asia and Europe, but a few spe-
cies are present in northern Africa. The species listed below is adventive, having been inadver-
tently introduced into California where it is successfully established.

Taxonomy: Hirashima, Moure and Daly, Jn Hirashima, 1971. Faculty Agr. Kyushu Univ.,
Jour. 16: 369-371, figs. 18-20 (deser., synopsis).
dallatorreana Friese. South. Europe and N. Africa; Calif. Successfully adventive in Calif.
(Central Valley). Parasite: Aprostocetus americanus Ashm., Ewrytoma apiculae Bugbee,
Grotea californica Cress. Pollen: Presumably polylectic, visits flowers of Centaurea
solstitialis, Convolvulus arvensis, Cucurbita, Melilotus officinalis in Calif.
Ceratina dallatorreana Friese, 1896. Termesz. Fusetek, 19: 38, 50. d, 2.

Biology: Daly, 1966. Ent. Soc. Amer., Ann. 59: 1138-1154, 17 figs., 5 tables (nest, floral
records, immature stages, life history).

TRIBE XYLOCOPINI

This tribe consists of generally large to very large, robust, pollen-collecting bees which occur
on all the continents and many of the islands. Included are three genera, Lestis with two species
in eastern Australia, Proxylocopa with 22 members centered in the more arid parts of the
southwestern Palaearctic Region and Xylocopa with more than 600 nominal forms, the majority
of which are found in the Ethiopian and Neotropical Regions.

Even though most members of this tribe (Lestis and Xylocopa) construct their nests in a wide
variety of usually dead plant materials, one group of species (Proxylocopa), insofar as known,
make their nests in the ground.

Revision: Hurd and Moure, 1963. Calif. Univ. Pubs. Ent. 29: 1-365, 244 figs., 1 frontispiece
(classification, includes biological information).

Genus XYLOCOPA Latreille

Taxonomy: Ackerman, 1916. N. Y. Ent. Soc., Jour. 24: 196-232, 1 pl. (U.S. spp.). —Hurd,
1955. Calif. Ins. Surv. Bul. 4: 35-72, 4 pls., 4 maps (U. S. spp.). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 507-511, figs. 126-127 (eastern U. S. spp.).

Biology: Malyshev, 1931. Ztschr. f. Morph. u. Okol. der Tiere (Abt. A. der Ztschr. f. wiss.
Biol.). 23 (3-4): 754-809, 15 figs. (nest, life history). —Hurd, 1958. Ent. Soc. Amer., Ann. 51:
365-375, 5 figs. (nesting habits). —Hurd, 1959. Kansas Ent. Soc., Jour. 32: 53-58, 1 fig.
(beefly parasitism).

Morphology: Wille, 1958. Ent. Soc. Amer., Ann.'51: 539-545 (dorsal vessel).

Genus XYLOCOPA Subgenus XYLOCOPA Latreille

Xilocopa Latreille, 1802. Hist. Nat. Fourmis, p. 432. Name suppressed by Internatl. Comn.
Zool. Nomencl., Op. 743, 1965.
Xylocopa Latreille, 1802. Hist. Nat. Crust. Ins., v. 3, p. 379. Name placed on Official List
of Generic Names in Zool. by Internat]. Comn. Zool. Nomencl., Op. 743, 1965.
Type-species: Apis violacea Linnaeus. Desig. by Westwood, 1840.

This subgenus occurs in the western Palaearctic.

Genus XYLOCOPA Subgenus SCHOENHERRIA Lepeletier

Xylocopa subg. Schoenherria Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 207.
Type-species: Xylocopa micans Lepeletier. Desig. by Sandhouse, 1943.
loripes Smith. Southeast. Ariz.; Mexico south to Honduras. Pollen: Unknown, but visits flowers
of Bouganvillea, Crotalaria.
Xylocopa loripes Smith, 1874. Ent. Soc. London, Trans., p. 298. d.
Xylocopa formosa Smith, 1874. Ent. Soc. London, Trans., p. 299. °.
micans Lepeletier. Southeast. Va. to Fla., west along Gulf Coast states to Texas; Mexico
(Tamaulipas). Ecology: Nests in sound, dead privet. Pollen: Polylectic, visits flowers of
Aesculus, Azalea, Bidens leucantha, Calopogon pulchellus, Carduus spinosissimus,

2184 Hymenoptera in America North of Mexico

Cephalanthus, Cercidium, Chaemaecrista brachiata, Cicuta, Crotalaria pumila,
Dalbergia ecastophyllum, Galactia, Glycine, Gossypium, Hibiscus, Ilex, Ligustrum,
Lythrum alatum, L. lineare, Medicago, Melilotus, Parkinsonia, Passiflora incarnata,
Phaseolus, Philibertella clausa, Prosopis, Prunus, Rhus, Richardia, Sabal palmetto,
Senecio, Serenoa serrulata, Solanum bahamense, S. wendlandii, Solidago, Stachys,
Suriana, Verbena brasiliensis, Vernonia.

?Apis nasuta Christ, 1791. Naturg. Klassif., Nomencl. Ins. Bienen-,Wespen-,
Ameisengeschl., v. 8, p. 130.

Xylocopa micans Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 208. ¢.

Xylocopa vidua Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 210. 2.

Xylocopa purpurea Cresson, 1872. Amer. Ent. Soe., Trans. 4: 284. ?.

Biology: Hurd, 1958. Ent. Soc. Amer., Ann. 51: 373, fig. 4 (nest).
Morphology: Maidl, 1912. Zool.-bot. Gesell., Verh. 62: 19-26, figs. 1-5 (gynandromorph).

Genus XYLOCOPA Subgenus NOTOXYLOCOPA Hurd

Xylocopa subg. Notoxylocopa Hurd, 1956. Amer. Mus. Novitates 1776: 2.
Type-species: Xylocopa tabaniformis Smith. Orig. desig.

Revision: O’Brien and Hurd, 1965. Ent. Soc. Amer., Ann. 58: 175-196, 38 figs. (taxonomy,
distr., nesting substrates).
tabaniformis androleuca Michener. West. N. Mex. to Utah, Nev., Ariz. and south. Calif.;

Mexico (Baja California and Sonora). Ecology: Nests in structural Douglas fir timbers
and possibly native juniper. Pollen: Apparently polylectic, visits flowers of Asclepzas,
Cucurbita pepo, Larrea tridentata, Oenothera parryi, Parkinsonia aculeata, Penstemon
barbatus, Stanleya pinnata.

Xylocopa orpifex androleuca Michener, 1940. South. Calif. Acad. Sci., Bul. 39: 127. d, @.

Biology: Hurd, 1958. Ent. Soc. Amer., Ann. 51: 373 (nesting habits). —Hurd and Linsley,
1975. Smithsn. Contrib. Zool. 193: 47 (floral relationships with Larrea tridentata).
tabaniformis orpifex Smith. Oreg., Calif.; Mexico (Baja California). Ecology: Nests in sound,

dead, native softwoods including Alnus rhombifolia, Juniperus, Libocedrus decurrens,
Populus, Pseudotsuga menziesii, Sequoia sempervirens, Sequoiadendron gigantea.
Parasite: Anthrax delila (Loew.), A. tigrinus (DeG.), Monodontomerus montivagus
Ashm., Nemognatha scutellaris Say. Pollen: Polylectic, visits a wide variety of native
and introduced flowers including Althaea rosea, Antirrhinum, Arctostaphylos patula,
Asclepias, Brassica, Ceanothus crassifolius, C. integerrimus, Cestrum, Chrysothamnus,
Cirsium, Citrus lemon, Clarkia breweri, C. concinna, C. elegans, C. gracilis, C. g.
albicaulis, C. mildrediae, C. unguiculata, Cordylanthus nevinii, Cucurbita, Delphinium,
Dicentra chrysantha, Diplacus auriantiacus, D. puniceus, Eriastrum virgatum,
Eriodictyon californicum, Fremontia, Fuchsia, Haplopappus parishii, Isomeris arborea,
Larrea tridentata, Lathyrus odoratus, Lepidospartum squamatum, Lonicera japonica,
Lotus scoparius, Lupinus densiflorus, L. grayi, Medicago sativa, Mentzelia laevicaulis,
Mimulus cardinalis, Mirabilis, Monardella, Oenothera deltoides, O. hookeri, Penstemon
breviflorus, P. bridgesii, P. grinnellii, P. labrosus, Phacelia davidsonii, P. distans, P.
heterophylla, P. ternatus, Rhododendron occidentale, Ribes, Rubus, Salvia mellifera,
Solanum umbelliferum, Solidago, Stachys albens, S. ajugoides, Thermopsis
macrophyllus, Trichostema lanceolatum, Verbascum blattaria, Wistaria, Wyethia.

Xylocopa orpifex Smith, 1874. Ent. Soc. London, Trans., p. 298. ?, ¢.

Xylocopa orpifera(!) Dalla Torre, 1896. Cat. Hym., v. 10, p. 216. Lapsus calami.

Xylocopa orpitex(!) Malyshev, 1931. Ztsch. Morph. Okol. Tiere (A) 238: (3-4): 788. Lapsus
calami.

Biology: Davidson, 1893. Ent. News 4: 151-153 (nest, life history, parasite). —Nininger, 1916.
Pomona Jour. Ent. Zool. 8: 158-165 (life history, parasite). —Hurd, 1959. Kans. Ent. Soc.,
Jour. 32: 54-56 (beefly parasitism). —Straw, 1956. Amer. Midland Nat. 90: 47-53 (floral
relationships with Penstemon). —Cruden, 1966. Pan-Pacifie Ent. 42: 111-119, 3 figs.
(territorial behavior). —Marston, 1970. Smithsn. Contrib. Zool. 43: 116 (beefly parasitism).

Superfamily APOIDEA = 2185

tabaniformis parkinsoniae Cockerell. South. Tex.; Mexico. Ecology: Nests in dead cottonwood.
Pollen: Unknown, but visits flowers of Cercidium, Citrus, Parkinsonia aculeata,
Verbesina encelioides. Other subspecies, including typical tabaniformis, occur from
Mexico to Ecuador.
Xylocopa tabaniformis parkinsoniae Cockerell, 1917. In W. P. Cockerell. N. Y. Ent. Soc.,
Jour. 25: 192. °.

Genus XYLOCOPA Subgenus XYLOCOPOIDES Michener

Xylocopa subg. Xylocopoides Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 155.
Type-species: Apis virginica Linnaeus. Orig. desig.

Taxonomy: Hurd, 1961. Amer. Ent. Soc., Trans. 87: 247-257, 2 pls. (synopsis). —Hurd and
Moure, 1963. Calif. Univ. Pubs. Ent. 29: 140-142 (revised keys to spp. and subspp.).

californica arizonensis Cresson. Tex. to south. Calif., Nev., and Utah; Mexico. Ecology: Nests
in sound, dead, native softwoods including Agave, Dasylirion, Nolina Yucca, but will
also nest in structural redwood when used for construction within its geographic range.
Parasite: Anthrax xylocopae Marston. Pollen: Polylectic, visits a wide variety of flowers
including those of Acacia angustissima, A. greggti, Agave nevadensis, Aloysia wrightii,
Argemone platyceras, Asclepias subverticillata, Baccharis, Bigelovia, Cercocarpus,
Chilopsis linearis, C. saligua, Chrysothamnus nauseosus, Cirsium, Citrus, Cleome,
Cryptantha oblata, Cucurbita maxima, C. mixta, C. moschata, C. pepo, Dalea albiflora,
Datura meteloides, Delphinium andesicola, Fendlera, Fouquieria splendens, Gaillardia
pulchella, Gaura coccinea, Gossypium, Heterotheca subaxillaris, Hyptis emoryi,
Kallstroemia grandiflora, Koeberlinia spinosa, Larrea tridentata, Lepidospartum
squamatum, Lippia lycioides, Lupinus magnificus, Medicago sativa, Melilotus alba,
Nama parryi, Mentzelia pumila, Parkinsonia aculeata, Penstemon, Prosopis
glandulosa var. torreyanum, Ratibida columnaris, Rhus laurina, R. trilobata, Robinia
neomexicana luxurians, Rubus, Salvia lemmoni, Senecio longilobus, S. salignus,
Solanum elaeagnifolium, Solidago, Sphaeralcea, Stanleya pinnata, Thurberia, Viguiera
longifolia, Wislizenia refracta, W. r. var. mamillata, Yucca brevifolia, Y. elata, Y.
whipplei.

Xylocopa arizonensis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 212. 2, 6.

Biology: Hurd, 1958. Ent. Soc. Amer., Ann. 51: 369-371 (nest). —Hurd, 1959. Kans. Ent. Soc.,
Jour. 32: 57-58 (nest, beefly parasitism). —O’Brien and O’Brien, 1966. Pan-Pacifie Ent. 42:
27-29 (nest, territoriality). —Marston, 1970. Smithsn. Contrib. Zool. 43: 118 (beefly
parasitism). —Hurd and Linsley, 1975. Smithsn. Contr. Zool. 193: 47, fig. 18 (floral
relationships).

californica californica Cresson. Oreg. (Cascade Mts.), Calif. (North, Coast Ranges and Sierra

Nevada Mts.). Ecology: Nests in sound, dead, native softwoods including Libocedrus
decurrens, Sequoia sempervirens, Sequoiadendron gigantea. Pollen: Polylectic, visits
flowers of Aesculus californica, Agastache urticifolia, Arctostaphylos, Aesclepias
speciosa, Ceanothus integerrimus, Cercis occidentalis, Chrysothamnus, Clarkia
dudleyana, C. elegans, C. unguiculata, Epigonia, Epilobium angustifolium, Eriodictyon
californicum, Lupinus albifrons, Medicago sativa, Mentha pulegium, Monardella
lanceolata, Penstemon breviflorus, P. grinnellii, Phacelia heterophylla, Ranunculus,
Solidago occidentalis, Yucca whipplet.

Xylocopa californica Cresson, 1864. Ent. Soc. Phila., Proc. 3: 40. °.

Xylocopa amblardi Perez, 1901. Act. Soc. Linn. Bordeaux 56: 115. d.

Xylocopa libocedri Cockerell, 1914. Insecutor Inscitiae Menstruus 2: 101. d, °.

Biology: Cruden, 1966. Pan-Pacific Ent. 42: 111-119, 3 figs. (nest, territorial and copulatory
behavior).
californica diamesa Hurd. Calif. (Centr. and south. coast ranges); Mexico (Baja California).
Ecology: Nests in dead floral scapes of Yucca whipplei. Pollen: Polylectic, visits
flowers of Arctostaphylos patula, Asclepias mexicana, Apocynum androsaemifolium,
Argemone platyceras, Ceanothus, Chrysothamnus bernardinus, C. nauseosus,
Cucurbita foetidissima, Dicentra chrysantha, Eriogonum fasciculatum, Fremontia

2186 Hymenoptera in America North of Mexico

californica, Haplopappus linearifolius, H. parishii, Isomeris arborea, Larrea tridentata,
Lepidospartum, Ligustrum, Lupinus austromontanus, L. grayi, L. parishii, Penstemon
palmeri, Phacelia heterophylla, Salvia, Stachys, Stanleya pinnata, Yucca whipplei.
Xylocopa californica diamesa Hurd, 1954. Pan-Pacifie Ent. 30: 202. d, ?.
virginica krombeini Hurd. Southern Fla. Predator: Horstia virginica Baker.
Xylocopa (Xylocopoides) virginica krombeini Hurd, 1961. Amer. Ent. Soc., Trans. 87: 251.
Onnce

Biology: Krombein, 1967. Trap-nesting wasps and bees, pp. 347-348, pl. 20, fig. 101 (nest
architecture, life history, predator).
virginica texana Cresson. South. Kans. to Tex. Pollen: Apparently polylectic, visits flowers of
Asclepias, Calyophus serrulata, Cercis canadensis, Eryngium, Passiflora incarnata,
Rubus, Salvia pitcheri, Vitex.
Xylocopa texana Cresson, 1872. Amer. Ent. Soc., Trans. 4: 283. 6, 2.
virginica virginica (Linnaeus). New England and adjacent Canada (Great Lakes region), south
to centr. Fla., west to Nebr., Kans., Okla. and east. Tex. Ecology: Nests in dead, sound
wood of many kinds of trees, including structural timbers. Parasite: Anthrax tigrinus
(DeG.), Villa sinuosus (Wied.). Pollen: Polylectic, visits flowers of a wide variety of both
native and introduced plants including Abelia, Amsonia tabernaemontana, Amygdalus
persica, Aquilegia vulgaris, Asclepias incarnata, A. pulchra, A. tomentosa, Azalea,
Barbarea barbarea, Bidens, Brauneria, Ceanothus ovatus, Cornus, Cyrilla racemiflora,
Diervilla japonica, Diospyros kaki, Eupatorium perfoliatum, Ilex, Lespedeza, Lonicera,
flava, L. japonica, Lythrum alatum, Malus, Medicago sativa, Melilotus alba, Mertensia,
Monarda, Opuntia, Passiflora incarnata, Pedicularis canadensis, Petalostemon,
Petunia, Phlomis tuberosa, Phlox divaricata, Physostegia, Plumbago capensis, Prunus,
Rhododendron nudiflorum, Rhus glabra, Robinia pseudoacacia, Rubus, Salvia azurea
grandiflora, Sassafras, Solidago canadensis, Stachys, Symphytum officinale, Syringa
vulgaris, Trifolium incarnatum, T. pratense, Vaccinium corymbosum, V. vacillans,
Verbena, Viburnum molle, Vicia villosa, Wistaria sinensis. Predator: Woodpeckers and
orioles.
Apis Virginica Linnaeus, 1771. Mant. Plant., v. 2, p. 450.
Apis Virginica Drury, 1773. Illust. Nat. Hist. Exotic Ins., v. 1 (index), p. 2.
Apis analis Fabricius, 1775. Systema Ent., p. 384.
Centris carolina Fabricius, 1804. Systema Piezatorum, p. 357. 6.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1062, figs. 167-172 (larva). —Hurd, 1955.
Calif. Ins. Surv., Bul. 4: 50, map 1 (distribution, floral records, nesting substrates).
—Moure, 1960. Studia Ent. 3: 160 (synonymy).

Biology: Angus, 1868. Amer. Nat. 1: 157 (parasite). —Walsh, 1868. Amer. Ent. 1: 9, f. 4
(habits, nest). —Packard, 1868. Amer. Nat. 1: 369-371, pl. 10, figs. 3-7 (nest, life history,
parasite). —Angus, 1869. Amer. Nat. 2: 48 (parasite). —Angus, 1869. Amer. Nat. 2: 49
(hibernation). —Packard, 1869. Guide to the study of insects, pp. 132-134, pl. 4 (habits).
—Campbell, 1873. Nature 8: 253 (predation by Baltimore orioles). —Packard, 1873. Our
common insects, p. 21-24 (habits). —Gentry, 1875. Amer. Nat. 9: 264-266 (pollination).
—Turner, 1878. Amer. Nat. 12: 627-628 (nest, predation by woodpeckers). —Mann, 1882.
Psyche 3: 298 (flower piercing habits). —Murtfeldt, 1882. Psyche 3: 343 (flower piercing
habits). —Pammel, 1888. Acad. Sci. St. Louis, Trans. 5: 241-277 (pollination, flower piercing
habits). —Howard, 1892. Ent. Soc. Wash., Proc. 2: 331 (hibernation). —Howard, 1902. The
insect book. N. Y.: Doubleday, Page and Co., p. 9 (nest). —Ashmead, 1894. Psyche 7: 23
(nesting habits). —Comstock and Comstock, 1895. Manual for Study of Insects, p. 670
(nesting habits). —Adams, 1915. Ill. State Lab. Nat. Hist., Bul. 11: 45, 47, 104, 198-199 (life
history). —Burrill, 1925. Science (n. s.) 62 (1597): 134 (flower piercing habits). —Rau, 1926.
Acad. Sci. St. Louis, Trans. 25: 162 (life history, parasite). —Rau, 1929. Jour. Compar.
Psychol. 9: 37 (homing experimentation). — Rau, 1931. Jour. Compar. Psychol. 12: 257
(homing experimentation). —Rau, 1933. The jungle bees and wasps of Barro Colorado
Island. Publ. by Phil Rau, Kirkwood, St. Louis Co., Mo., pp. 225-226, figs. (habits and
instincts, life history). —Burrill, 1934. Missouri State Museum, Bul. 11: 3-4 (flower piercing
habits). —Weiss and Smith, 1940. U.S. Dept. Agr. Circular 556: 16-19, fig. (flower-spot

Superfamily APOIDEA 2187

disease transmission). —Franklin, 1951. Kans. Agr. Expt. Sta. Tech. Bul. 70: 1-64
(pollination). —Hurd, 1955. Calif. Ins. Surv., Bul. 4: 50 (nesting substrates). —Chandler,
1958. Pest Control 26: 36, 38, 40, 47, 3 figs. (economic damage). —Hurd, 1958. Ent. Soc.
Amer., Ann. 51: 368, 373-374, fig. 5 (nesting substrates). —Hurd, 1959. Kans. Ent. Soc.,
Jour. 32: 56-57 (beefly parasitism). —Balduf, 1961. Brooklyn Ent. Soc., Bul. 56: 81-84 (nest
associates). —Balduf, 1962. Ent. Soc. Amer., Ann. 55: 263-271, 3 figs. (life history).
—Sabrosky, 1962. Ent. Soc. Wash., Proc. 64: 184 (mating behavior). —Dorr and Martin,
1966. Michigan Quart. Bul. 48: 445 (flower piercing habits). —Krombein, 1967. Trap-nesting
wasps and bees, pp. 345-346 (nest architecture, life history). —Marston, 1970. Smithsn.
Contrib. Zool. 43: 119 (beefly parasitism). —Gerling and Hermann, 1976. Entomophaga 21:
227-223, 3 figs., 3 tables (beefly parasitism).

Morphology: Milliron, 1958. Brooklyn Ent. Soc., Bul. 53: 66-68 (gynandromorph).
Genus XYLOCOPA Subgenus NEOXYLOCOPA Michener

Xylocopa subg. Neoxylocopa Michener, 1954. Amer. Mus. Nat. Hist., Bul. 104: 157.
Type-species: Apis brasilianorum Linnaeus. Orig. desig.
mexicanorum Cockerell. Tex. to east. Ariz.; Mexico.
Xylocopa mexicanorum Cockerell, 1912. Ann. and Mag. Nat. Hist. (8) 9: 555. 2.
varipuncta Patton. Ariz, Nev., Calif.; Mexico. Ecology: Nests in rotting or decaying wood.
Parasite: ?7Anthrax simson habrosus Marston. Pollen: Polylectic, visits a wide variety of
flowers including Acacia, Argemone, Asclepias, Astragalus parishii, Brassica
campestris, Buddleia, Calliopsis, Ceanothus hookeri, Cercidium Torreyanum, Cucurbita
foetidissima, C. maxima, C. mixta, C. moschata, C. pepo, Datura meteloides,
Eschscholzia californica, Gossypium, Lantana camara, Larrea tridentata, Lathyrus
odoratus, Lonicera japonica, Lupinus paynei, Lycopersicum esculentum, Medicago
sativa, Oenothera hookeri, Parkinsonia aculeata, Passiflora, Penstemon antirrhinoides,
Prosopis glandulosa var. torreyanum, Salvia, Sesbania macrocarpa, Solanum
douglasti, S. elaeagnifolium, Sphaeralcea emoryi, Trichostema lanceolatum, Vitex
pyramidata, Wislizenia refracta, W. r. var. mamillata, Wistaria. X. sonorina Smith,
described from Hawaii, has been regarded by some investigators as a senior synonym.
Xylocopa varipuncta Patton, 1879. Canad. Ent. 11: 60. 9.

Biology: Nininger, 1916. Pomona Jour. Ent. Zool. 8: 164-165 (nest, life table data). —Hurd,
1958. Ent. Soc. Amer., Ann. 51: 369, fig. 1. —Janzen, 1964. Pan-Pacific Ent. 40: 65-66 (nest).
—Marston, 1970. Smithsn. Contrib. Zool. 43: 5 (possible parasite). —Hurd and Linsley,
1975. Smithsn. Contrib. Zool. 193: 47-48 (intrafloral ecology).

Morphology: Gordh and Gulmahamad, 1975. Ent. Soc. Wash., Proc. 77: 269-2738, 8 figs.
(gynandromorph).

Genus XYLOCOPA Subgenus STENOXYLOCOPA Hurd and Moure

Xylocopa subg. Stenoxylocopa Hurd and Moure, 1960. Ent. Soe. Amer., Ann. 53: 809.
Type-species: Xylocopa artifex Smith. Orig. desig.

Taxonomy: Hurd and Moure, 1960. Ent. Soc. Amer., Ann. 53: 809-821, 14 figs., 2 tables
(synopsis and summary of biological information).
artifex Smith. Mts. of south. Ariz. to Argentina. Ecology: Nests in hollow culms, usually dead,

of bamboo and related plants including Arundo donax, Chusquea bambusoidea, Guadua,
Merostachys clausseni. Pollen: Presumably polylectic, visits in South America flowers
of Baccharis dracunculifolia, Cassia bicapsularis, C. splendida, Crotalaria paulina,
Leonurus sibircus, Passiflora, Solanum atropurpureum, S. balbisii, S. paniculatum,
Tecoma ipe.

Xylocopa artifex Smith, 1874. Ent. Soc. London, Trans., p. 289. 2.

Xylocopa erratica Smith, 1874. Ent. Soc. London, Trans., p. 293. d.

Taxonomy: Hurd, 1955. Calif. Ins. Surv., Bul. 4: 62-63 (synonymy, distribution).

2188 Hymenoptera in America North of Mexico

Biology: Schrottky, 1902. Rev. Mus. Paulista 5: 468 (floral records). —Schrottky, 1904.
Allgem. Ztschr. Ent. 9: 344-349 (nest). —Strand, 1912. Zool. Jahrb. Abt. Syst. 33: 267
(nest). —Bertoni, 1918. An. Cient. Paraguayos (2) 3: 219 (nest). —Moure, 1942. Papeis
Avulsos Dept. Zool., Secretaria Agr. Sao Paulo 2: 301 (nest). —Hurd and Moure, 1960. Ent.
Soe. Amer., Ann. 53: 819-820 (nest substrates).

Family APIDAE

Species of this family are found throughout much of the world from the high Arctic latitudes
to or near the southern limits of the major land masses of the Southern Hemisphere. The family
consists of two subfamilies, the Bombinae which includes the orchid bees (Euglossini) and bum-
blebees (Bombini) and the Apinae which contains the stingless honeybees (Meliponini) and the
familiar stinging honeybees (Apini). Some of these bees are of exceptional value to man not only
because of their production of honey—and other_products, but also because they pollinate many
agricultural and other plants. Although the family contains some social parasites (e.g., Aglae,
Exaerete and Psithyrus) and nest robbers (e.g., Lestrimelitta), the pollen-collecting females, un-
like those of any other family of bees, transport pollen by means of specialized pollen baskets
(corbiculae) located on the hind tibiae. Virtually all stages of social development are exhibited by
the family. These include all of the highly eusocial bees (Apinae) which live in perennial colonies
as well as the primitively eusocial bumblebees and the solitary and parasocial Euglossini.

In America north of Mexico the family is represented most conspicuously by the introduced
European honeybee (Apis mellifera Linnaeus) and the many native species of bumblebees. The
only other member of this family present in the United States is a species of the Neotropical
genus Eulaema which was found years ago in the vicinity of Brownsville, Texas. In spite of re-
peated attempts to introduce various species of meliponine bees into the United States none of
these introductions has been successful.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 513-546, figs. 128-134, table
18 (east. U. S. spp.). —Winston and Michener, 1977. Natl. Acad. Sci., U.S. A., Proc. 74:
1135-1137 (phylogeny).

Biology: Michener, 1974. The Social Behavior of Bees, 404 pp., Cambridge, Massachusetts,
The Belknap Press of Harvard University Press.

Morphology: Lello, 1976. Kans. Ent. Soe., Jour. 49: 85-99, 22 figs., 3 tables (adnexal glands of
the sting apparatus).

SUBFAMILY BOMBINAE

This subfamily contains two tribes, the Euglossini which is found only in the Neotropical Re-
gion and the Bombini which, although chiefly Holarctic in occurrence, is represented by a
number of species in both the Oriental and Neotropical Regions. Several species of this latter
tribe have been successfully introduced into the Australian Region (New Zealand).

TRIBE EUGLOSSINI

These bees, which are found only in the Americas (Chihuahua, Sonora, and Texas to Argen-
tina) are centered in the tropics and are most abundant and diverse mh the forested regions. The
tribe consists of six genera, four of which contain pollen-collecting species (Eufriesea, Euglossa,
Eulaema, and Euplusia) and two others (Aglae and Exaerete) whose species are social parasites
in the nests of the pollen-collecting Euglossini. Many of the species are large and brillantly
metallic while others are clothed with contrasting patterns of black and white or yellow and
white pubescence reminiscent of bumblebees. The males of many, if not all, pollen-collecting spe-
cies, unlike most females, are selectively attracted to the flowers of one or more species of
orchids, many of which produce little or no nectar and evidently no food. The attraction of the
males to these flowers is not fully understood. It is believed that lek behavior is involved and
that the males seek certain substances from these flowers, as well as from other sources, and
store these in the organs of their greatly enlarged hind tibiae. It has been suggested that these
substances are converted into sex pheromones, possibly species-specific, which are used to at-
tract females responsive to mating. The males of different species visit the flowers of different
species of orchids and often dislodge pollinia which adhere to the head, thorax or legs. Thus

Superfamily APOIDEA 2189

transported to other flowers of the same species, the pollinia are readily available to effect pol-
lination. This bee-plant (bee-flower) relationship obviously has important implications relative to

the evolution of orchids and these bees.

Taxonomy: Moure, 1950. Dusenia 1: 181-200 (included genera). —Moure, 1963. Rev. Biol.
Tropical 11: 211-216 (tax. characters, key to included genera). —Moure, 1967. Atas
Simposio Biota Amazonica 5 (Zool.): 395-415 (checklist of included genera and spp.).

Biology: Dodson and Frymire, 1961. Mo. Bot. Garden, Bul. 49: 133-152 (floral relationships).
—Dodson and Frymire, 1961. Mo. Bot. Garden, Ann. 48: 137-172 (floral relationships).
—Dodson, 1962. Mo. Bot. Garden, Ann. 49: 35-36 (orchid pollination). — Vogel, 1963. Osterr.
Bot. Ztschr. 110: 308-337 (orchid pollination). —Bennett, 1965. Ins. Sociaux 12: 81-91 (social
behavior). —Pijl and Dodson, 1966. Orchid flowers; their pollination and evolution, 214 pp.
(intrafloral relationships). — Vogel, 1966. Osterr. Bot. Ztschr. 113: 302-361 (orchid
pollination). —Dodson, 1966. Kans. Ent. Soc., Jour. 39: 607-629 (ethology). —Dressler, 1967.
Atas Simposio Biota Amazonica 5 (Zool.): 171-180 (pollination syndrome). —Zucchi and
Camargo, 1969. Fac. Sci. Hokkaido Univ., ser. 6, Zool. 17: 271-380, 2 pls., 21 figs., 9 tables
(review of biology). —Dodson, 1970. Biochemical Coevolution, Oreg. State Univ. Press, pp.
83-107 (role of chemical attractants in orchid pollination). —Janzen, 1971. Science 171:
203-205 (long distance pollination). —Evoy and Jones, 1971. Anim. Behavior 19: 579-584
(motor patterns evoked in males by floral fragrances). —Bennett, 1972. N. Y. Ent. Soc.,
Jour. 80: 137-145 (attraction and collection by use of baited McPhail fruitfly traps).
—Williams and Dodson, 1972. Evolution 26: 84-95, 4 figs., 5 tables (selective attraction of
males to orchid floral fragrances). —Michener, 1974. The Social Behavior of Bees, Chapter
21: 257-260 (natural history of orchid bees). —Dodson, 1975. Jn Gilbert and Raven,
Coevolution of plants and animals, pp. 91-99, 2 figs. (coevolution with orchids).

Morphology: Lanham, 1951. Pan-Pacifie Ent. 27: 181-182 Gugal brush). —Cruz-Landim, Stort,
Costa Cruz and Kitajima, 1965. Rev. Brasil. Biol. 25: 323-342 (tibial organs of males).
—Hills, Williams and Dodson, 1968. Amer. Orchid Soc., Bul. 37: 967-971 (identification of
odor compounds in orchid fragrances). —Dodson, Dressler, Hills, Adams and Williams,
1969. Science 164: 1243-1249 (chemical substances in orchid fragrances attractive to males).

Genus EULAEMA Lepeletier

Taxonomy: Moure, 1950. Dusenia 1: 181-200 (key to related genera, key to included spp.,
synonymy and tax., bibliography of included spp.). —Moure, 1960. Studia Ent. 3: 145-147
(Fabrician Types). —Moure, 1967. Atas Simposio Biota Amazonica 5 (Zool): 410-418, fig. 1
(checklist of included spp.).

Genus EULAEMA Subgenus EULAEMA Lepeletier

Eulaema Lepeletier, 1841. Hist. Nat. Ins., Hym., v. 2, p. 11. Name placed on Official List
of Generic Names in Zool. by Internat]. Comn. Zool. Nomencl., Op. 567, 1959.
Type-species: Apis dimidiata Fabricius. Desig. by Taschenberg, 1883, as validated
by Internatl. Comn. Zool. Nomencl., Op. 567, 1959.
Eulaenis(!) Spinola, 1851. In Gay, Hist. Fiscia Polit. Chile, Zool., v. 6, p. 167.
Eulema Dalla Torre, 1896. Cat. Hym., v. 10, p. 309. Emend.

This subgenus does not occur in America north of Mexico.
Genus EULAEMA Subgenus APEULAEMA Moure

Eulaema subg. Apeulaema Moure, 1950. Dusenia 1: 184.
Type-species: Centris cingulata Fabricius. Orig. desig. (=~Eulaema fasciata

Lepeletier).

polychroma (Mocsary). Tex. (Brownsville); Mexico (Sonora) south to Ecuador and Peru.
Euglossa polychroma Mocsary, 1899. Termes. Fuzetek 22: 170. d.

Taxonomy: Moure, 1967. Atas Simposio Biota Amazonica 5 (Zool.): 400, 412 (tax. status,
geogr. range).

2190 Hymenoptera in America North of Mexico
TRIBE BOMBINI

The Bombini includes common, large or medium-sized, social bees which are conspicuously
marked with yellow and black, and sometimes, in addition, with red, or white hair. While most of
the species are pollen-collecting bees, the species of the genus Psithyrus live as social parasites
in the nests of the genus Bombus. Species of this tribe are found in North and South America,
Eurasia including Japan and Taiwan, the Philippine Islands, Indonesia east to Java and a few
species are successfully adventive in New Zealand.

Revision: Franklin, 1912. Amer. Ent. Soc., Trans. 38: 177-486 (New World spp., first part).
—Franklin, 1913. Amer. Ent. Soc., Trans. 39: 73-200, 22 pls. (New World spp., concluding
part). —Milliron, 1970. Ent. Soc. Canada, Mem. 65: i-lii (bibliography of New World spp.).
—Milliron, 1971. Ent. Soc. Canada, Mem. 82: 1-80, pls. I-XI, 14 figs., 6 maps (Part I:
contains information on development of color, distribution, external morphology,
hybridization, life histories, phylogeny and clagsification; treats west. hemisphere spp. of
genus Bombus and genus Megabombus subg. Bombias). —Milliron, 1972. Ent. Soe.
Canada, Mem. 89: 81-237, pls. XII-XV, maps 7-35 (Part II: treats west. hemisphere spp. of
genus Megabombus subg. Megabombus). —Milliron, 1973. Ent. Soc. Canada, Mem. 91:
239-333, pls. XVI-XX, maps 36-56 (Part III: tyeats west. hemisphere spp. of genus
Pyrobombus subg. Cullumanobombus).

6

Taxonomy: Frison, 1919. Ill. Acad. Sci., Trans. 12: 157-165 (Illinois spp.). —Jackson, 1920.
Ent. Soe. Wash., Proce. 22: 162-168 (District of Columbia and vicinity). —Lutz and
Cockerell, 1920. Amer. Mus. Nat. Hist., Bul. 42: 502-544 (catalog of N. Amer. spp).
—Fattig, 1923. Fla. Ent. 7: 25 (Florida spp.). —Neave, 1926. Ent. News 87: 252-254
(Alberta spp.). —Frison, 1927. Amer. Ent. Soc., Trans. 53: 51-78, 2 pls. (systematic
relationships of spp. in Amer. north of Mexico). —Scullen, 1927. Pan-Pacifie Ent. 4: 69-76,
121-128 (Oregon spp.). —Neave, 1933. Canad. Jour. Res. 8: 62-72 (Manitoba spp.).
—Milliron, 1939. Mich. Acad. Sci. Arts and Letters, Papers 24 (2): 168-182 (Michigan spp.).
—Chandler, 1950. Indiana Acad. Sci., Proc. 60: 167-177 (Indiana spp.). —Franklin, 1954.
Amer. Ent. Soc., Trans. 80: 43-51 (evolution, distribution). —Stephen, 1957. Oreg. Agr.
Expt. Sta. Tech. Bul. 40: 1-163, 31 figs., 18 maps (revision of western N. Amer. spp.).
—Milliron, 1961. Kans. Ent. Soc., Jour. 34: 49-61. (generic reclassification). —LaBerge and
Webb, 1962. Nebr. Agr. Expt. Sta. Res. Bul. 205: 1-38 (Nebraska spp.). —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 513-544, figs. 128-134, table 18 (east. U. S. spp.).
—Medler and Carney, 1963. Wis. Agr. Expt. Sta. Res. Bul. 240: 1-47 (Wisconsin spp.).
—Pisani, Sakagami, Crestana, Aily and Lorenzetti, 1968. Acad. Sci. Brasil. Cienc. Letras,
An. 40: 373-389 (morphometrics). —Faester and Hammer, 1970. Ent. Meddel. 38: 257-302
(cent. and north. European spp.). —Stephen and Koontz, 1973. Melanderia 13: 1-12, 52 figs.
(larvae). —Plowright and Stephen, 1973. Canad. Ent. 105: 733-743 (numerical taxonomic
analysis of evolutionary relationships). —Stephen and Cheldelin, 1973. Biochem.
Systematics 1: 69-76 (phenetic groups based on a-glycerophosphate dehydrogenase).

Biology: Putnam, 1864. Essex Inst. Salem, Proc. 4: 98-105 (habits). —Packard, 1864. Essex
Inst. Salem 4: 107-140, pl. 3 (life histories of New England spp., parasites). —Sladen, 1912.
The humble-bee, its life history and how to domesticate it, xiii and 283 pp., London,
Macmillan. —Frison, 1926. Ent. Soc. Amer., Ann. 19: 203-234, pl. XVII (natural enemies of
Ill. spp.). —Plath, 1927. Ent. Soc. Amer., Ann. 20: 181-192 (hibernation). —Free and Butler,
1959. Bumblebees, xiv and 208 pp., London, Collins. —Plath, 1934. Bumblebees and their
ways, 201 pp., New York, Macmillan. —Macfarlane, 1973. Ent. Soc. Ont., Ann. 104: 27-30
(predators). —Michener, 1974. The social behavior of the bees, chapter 28: 314-328, figs.,
Cambridge, Massachusetts, The Belknap Press of Harvard University Press (natural
history). —Pouvreau, 1974. Apidologie 5: 39-81 (enemies of adults). — Alford, 1975.
Bumblebees, 352 pp., London, Davis-Poynter Ltd. (life histories of British spp.).

Morphology: Barendrecht, 1931. Acta Zool. 12: 153-204, 23 figs. (corpora pedunculata).
—Palm, 1949. Opuse. Ent. 14: 27-47, 8 figs. (pharyngeal gland).

Superfamily APOIDEA 2191

Genus BOMBUS Latreille

The genus Bombus comprises medium to large-sized, nest-building, social species, each of
which normally has a worker caste and a primary reproductive caste of males and females. The
females, or queens, and the workers have pollen-collecting baskets on the hindlegs. These bees
usually live in colonies in or on the ground, often in the deserted nests of rodents. Most of the
species in America north of Mexico depend upon a comparatively wide variety of flowers for
nectar and pollen. The classification and arrangement of the subgenera in this catalog is that of
Richards (1968. Brit. Mus. Nat. Hist. Ent., Bul. 22: 209-276, 39 text-figs.).

Revision: Franklin, 1912. Amer. Ent. Soc., Trans. 38: 177-486 (New World spp., first part ).
—Franklin, 1913. Amer. Ent. Soc., Trans. 39: 73-200, 22 pls. (New World spp., concluding
part). —Milliron, 1970. Ent. Soc. Canada, Mem. 65: i-lii (bibliography of New World spp.).
—Milliron, 1971. Ent. Soc. Canada, Mem. 82: 1-80, pls. I-XI, 14 figs., 6 maps (Part I:
contains information on development of color, distribution, external morphology,
hybridization, life histories, phylogeny and classification; treats west. hemisphere spp. of
genus Bombus and genus Megabombus subg. Bombias). —Milliron, 1972. Ent. Soc.
Canada, Mem. 89: 81-237, pls. XII-XV, maps 7-35 (Part II: treats west. hemisphere spp. of
genus Megabombus subg. Megabombus). —Milliron, 1973. Ent. Soc. Canada, Mem. 91:
239-333, pls. XVI-XX, maps 36-56 (Part III: treats west. hemisphere spp. of genus
Pyrobombus subg. Cullumanobombus). —Loken, 1973. Norsk Ent. Tidsskr. 20: 1-218, 99
figs., 26 Tables (Scandinavian spp., including certain circumpolar spp.).

Taxonomy: Lutz, 1916. Amer. Mus. Nat. Hist., Bul. 35: 501-521 (geogr. distribution). —Frison,
1919. Ill. Acad. Sci., Trans. 12: 157-165 (Illinois spp.). —Jackson, 1920. Ent. Soc. Wash.,
Proc. 22: 162-168 (District of Columbia and vicinity). —Lutz and Cockerell, 1920. Amer.
Mus. Nat. Hist., Bul. 42: 502-539 (catalog of N. Amer. spp.). —Fattig. 1923. Fla. Ent. 7: 25
(Florida spp.). —Neave, 1926. Ent. News 37: 252-254 (Alberta spp.). —Frison, 1927. Amer.
Ent. Soc., Trans. 53: 51-78, 2 pls. (systematic relationships of spp. in Amer. north of
Mexico). —Scullen, 1927. Pan-Pacifie Ent. 4: 69-76, 121-128 (Oregon spp.). —Plath, 1927.
Biol. Bul. 52: 394-410, 6 figs., 2 tables (natural groupings based on biological characters).
—Richards, 1931. Tromso Mus. Arshefter 50 (1927): 1-32, 2 pls. (spp. allied to Bombus
alpinus). —Ritcher, 1933. Ent. Soc. Amer., Ann. 26: 53-63, 33 figs. (larvae). —Neave, 1933.
Canad. Jour. Res. 8: 62-72 (Manitoba spp.). —Skorikov, 1937. Ent. Meddel. 20: 37-64
(Greenland spp. and their circumpolar relationships). —Milliron, 1939. Mich. Acad. Sci. Arts
and Letters, Papers 24 (2): 168-182 (Michigan spp.). —Chandler, 1950. Indiana Acad. Sci.,
Proc. 60: 167-177 (Indiana spp.). —Kruseman, 1952. IXth Internat]. Congr. Ent., Trans. 1:
101-102 (subgeneric classification). —Franklin, 1954. Amer. Ent. Soc., Trans. 80: 43-51
(evolution and distribution). —Chandler, 1954. Indiana Acad. Sci., Proc. 63: 165-167 (color
variation). —Franklin, 1955. Ent. News 66: 65-68 (morphometric studies of males).
—Chandler, 1956. Indiana Acad. Sci., Proc. 65: 116-117 (parallel color variation). —Stephen,
1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 1-163, 31 figs., 18 maps (west. N. Amer. spp.).
—Milliron, 1960. Brooklyn Ent. Soc., Bul. 55: 87-99 (types of west. hemisphere spp.).
—Milliron, 1961. Kans. Ent. Soc., Jour. 34: 49-61 (classification). —Thorp, 1962. Pan-Pacific
Ent. 38: 21-28 (distribution of some west. N. Amer. spp.). —LaBerge and Webb, 1962.
Nebr. Agr. Expt. Sta. Res. Bul. 205: 1-38 (Nebraska spp.). —Medler, 1962. Ent. Soc. Amer.,
Ann. 55: 212-218, 1 fig., 4 tables (morphometric studies of radial cell, glossa, prementum
and first segment of labial palpus). —Medler, 1962. XIth Internat]. Kongr. Ent. Wien,
Verh. 2 (1960): 517-521 (morphometric analyses of mouthparts). —Milliron, 1962. Canad.
Ent. 94: 728-735 (synonymies of New World spp.). —Mitchell, 1962. N. C. Agr. Expt. Sta.
Tech. Bul. 152: 513-538, figs. 128-133, table 18 (eastern U.S. spp.). —Medler and Carney,
1963. Wis. Agr. Expt. Sta. Res. Bul. 240: 1-47 (Wisconsin spp.). —Hazeltine and Chandler,
1964. Kans. Ent. Soc., Jour. 37: 77-87, 6 pls. (queens). —Hobbs, 1964. Canad. Ent. 96:
115-116 (phylogeny based on brood-rearing behavior). —Richards, 1968. Brit. Mus. Nat.
Hist., Bul. (Ent.) 22: 211-276, 39 figs. (subgeneric classification). —Thorp, 1969. Pan-Pacific
Ent. 45: 87-96 (key to spp. of flavifrons group). —Thorp, 1970. Pan-Pacifie Ent. 46: 177-180
(notes on Ariz. records of Bombus balteatus, B. franklini, B. sylvicola). —Loken, 1973.
Norsk Ent., Tidsskr. 20: 1-218, 99 figs. (Scandinavian spp., including certain cireumpolar
spp.). —Stephen and Koontz, 1973. Melanderia 13: 1-12, 52 figs. (larvae).

2192 Hymenoptera in America North of Mexico

Biology: Putnam, 1864. Essex Inst. Salem, Proc. 4: 98-105 (habits). —Packard, 1864. Essex
Inst. Salem, Proe. 4: 107-140, pl. 3 (life histories of New England spp., parasites).
—Putnam, 1865. Essex Inst. Salem, Proe. 4: 98-105 (life histories of New England spp.).
—Coville, 1890. Ent. Soe. Wash., Proce. 1: 197-208 (life histories). —Sladen, 1912. The
humble bee, its life history and how to domesticate it, xiii and 283 pp., London Macmillan.
—Fantham and Porter, 1914. Tropical Med. Parasit., Ann. 8: 623-638 (parasitism by
Nosema bombi). —Wheeler, 1919. Psyche 26: 145-152 (phoresy of Antherophagus).
—Betts, 1920. Bee World 1: 171 (Nosema). —Frison, 1921. Amer. Nat. 55: 188-192
(Antherophagus in nests). —Plath, 1922. Psyche 29: 189-202 (nesting habits). —Plath, 1923.
Psyche 30: 145-154 (colony trumpeter). —Plath, 1923. Biol. Bul. 45: 325-341 (queen
breeding experiments). —Plath, 1923. Psyche 39: 193-202 (egg-eating habits). —Plath, 1923.
Amer. Nat. 57: 571-574 (skunk predation). —Plath, 1924. Biol. Bul. 47: 65-78, 2 figs. (life
histories). —Plath, 1925. Amer. Nat. 59: 441-451 (role of pollination in certain cultivated
crops). —Frison, 1926. Ent. Soc. Amer., Ann. 19: 203-234, pl. XVII (natural enemies of III.
spp.). —Frison, 1926. Econ. Ent., Jour. 19: 149-155 (artificial domiciles). —Plath, 1927. Ent.
Soe. Amer., Ann. 20: 181-192 (hibernation). —Plath, 1927. Psyche 34: 122-128, 2 figs.
(nesting habits of some New England spp.). —Frison, 1927. Econ. Ent., Jour. 20: 522-526
(fertilization and hibernation of queens under controlled conditions). —Frison, 1930. Canad.
Ent. 62: 49-54 (flight orientation). —Balduf, 1939. Canad. Ent. 71: 66-74 (predation by
Phymata). —Plath, 1934. Bumblebees and their ways, 201 pp., New York, MacMillan.
—Balduf, 1941. Ent. Soc. Amer., Ann. 34: 204-214 (predation by Phymata). —Linsley,
1944. Brooklyn Ent. Soe., Bul. 39: 48-49 (vernal flight of males). —Cumber, 1949. Roy. Ent.
Soe. London, Proc. 100: 1-45, 10 figs. (life history, production of worker caste). —Cumber,
1949. New Zealand Sci. Rev. 7: 96-97 (overwintering nest). —Pedersen and Bohart, 1950.
Agron. Jour. 42: 608 (use as pollinators of small seed plots). —Loken, 1954. Bergen Univ.
Pub., Biol. Sta. 13: 1-6 (behavior during solar eclipse). —Brian, 1954. Bee World 35: 61-67,
81-91 (foraging behavior). —Cumber, 1954. New Zealand Jour. Sci. Tech. (B) 36: 95-107
(life cycle). —Fye and Medler, 1954. Econ. Ent., Jour. 47: 672-676 (field domiciles). —Fye
and Medler, 1954. Wis. Acad. Sci. Arts and Letters 43: 75-82 (spring emergence). —Fye
and Medler, 1954. Econ. Ent., Jour. 47: 847-852 (temperature studies of domiciles). —Free,
1955. Roy. ent. Soe. London, Proce. (A) 30: 19-25 (queen production). —Free 1955. Ins.
Sociaux 2: 195-212 (division of labor within colonies). —Free, 1955. Brit. Jour. Anim.
Behav. 3: 147-153 (behavior of egg-laying workers). —Free, 1955. Ins. Sociaux 2: 303-311
(collection of food). —Manning, 1956. Behaviour 9: 164-201 (foraging behavior). —Free,
1957. Roy. Ent. Soe. London, Proce. 32: 182-184 (social facilitation and ovary development).
—Medler, 1957. Ins. Sociaux 4: 252-254 (pollination of alfalfa and red clover). —Free, 1958.
Behaviour 12: 233-242 (defense of colony). —Medler, 1958. Xth Internat]. Congr. Ent.,
Proe. 4: 973-981 (use in pollination of agricultural crops). —Free and Butler, 1959.
Bumblebees, xiv and 208 pp., London, Collins. —Hobbs, Virostek and Nummi, 1960. Canad.
Ent. 92: 868-872 (artificial domiciling). —Holm, 1960. Arsskr. Kgl. Vet.-Landbhojsk, pp.
1-19 (domestication). —Hasselrot, 1960. Opuse. Ent. Suppl. 17: 1-192 (domestication).
—Pringle, 1961. Nat. Hist. 70: 20-29 (flight behavior). —Montgomery, 1961. Outdoor
Indiana 4: 16-22 (bombiculture). —Hobbs, 1962. Canad. Ent. 94: 538-541 (food-gathering
behavior). —Medler, 1962. Canad. Ent. 94: 825-833, 1 fig., 3 tables (egg development and
absorption). —Hobbs, Nummi and Virostek, 1962. Canad. Ent. 94: 1121-1132 (management
for pollination). —Cumber, 1963. New Zealand Jour. Sci. 6: 66-74 (domiciling of an
unusually large nest). —Hobbs, 1964. Canad. Ent. 96: 115-116 (brood-rearing behavior).
—Hobbs, 1964. Canad. Ent. 96: 1465-1470 (ecology of Alpinobombus). —Hobbs, 1965.
Canad. Ent. 97: 120-128 (ecology of Bombias). —Armitage, 1965. Kans. Ent. Soc., Jour. 38:
89-100, 4 figs., 4 tables (predation by Philanthus bicinctus). —Hobbs, 1965. Canad. Ent.
97: 1297-1302 (ecology of Cullwmanobombus). —Knee and Medler, 1965. Canad. Ent. 97:
1149-1155 (seasonal size increase in workers). —Plowright and Jay, 1966. Jour. Apicult.
Research 5: 155-165 (domestication). —Hobbs, 1966. Canad. Ent. 98: 33-39 (ecology of
Fervidobombus). —Holm, 1966. Ann. Rev. Ent. 11: 155-182 (management of red clover
and alfalfa pollinating spp. for seed production). —Milliron and Oliver, 1966. Canad. Ent.
98: 207-213, 6 figs. (nests, usurpation and life histories of Ellesmere Island spp.). —Hobbs,
1966. Canad. Ent. 98: 288-294 (ecology of Subterraneobombus). —Johansen, 1967. Wash.
State Univ. Tech. Bul. 57: 1-12 (ecology of southwest. Wash. spp.). —Hobbs, 1967. Canad.

Superfamily APOIDEA 2193

Ent. 99: 943-951 (management of red clover pollinating spp.). —Hobbs, 1967. Canad. Ent.
99: 1271-1292 (ecology of Pyrobombus). —Hobbs, 1968. Canad. Ent. 100: 156-164 (ecology
of Bombus, s. str.). —Husband, 1968. Mich. Acad. Sci. Arts Letters, Papers, 53: 109-112
(Acarina associated with Mich. spp.). —Plowright and Jay, 1968. Ins. Sociaux 15: 171-192
(caste differentiation: determination of female size). —Alford, 1969. Jour. Anim. Ecol. 38:
149-170 (hibernation). —Eaton and Stewart, 1969. Canad. Ent. 101: 149-150 (damage of
blueberry blossoms by bumblebees). —Alford, 1970. Ins. Sociaux 17: 1-10 (incipient stages
of colonial development). —Free, 1970. Jour. Anim. Ecol. 39: 395-402 (flower constancy).
—Free, 1971. Behaviour 40: 55-61 (stimuli eliciting mating behavior). —Kevan, 1972. Jour.
Ecol. 60: 831-847, 1 fig., 4 tables (pollination of high arctic flowers). —Richards, 1973.
Quaest. Ent. 9: 115-157, 29 figs. (life histories of arctic spp.). —Richards, 1974. Kans. Ent.
Soc., Jour. 47: 141-142 (nest site selection, Alta. spp.). —Macior, 1974. Melanderia 15: 1-59,
10 tables (pollination ecology of spp. in front range of Colorado Rocky Mountains).
—Michener, 1974. The social behavior of the bees, chapter 28: 314-328, figs., Cambridge,
Massachusetts, The Belknap Press of Harvard University Press (natural history).
—Heinrich, 1975. In Gilbert and Raven, Coevolution of animals and plants, pp. 141-158, 4
figs., 2 tables (energetics). —Macior, 1975. Amer. Jour. Bot. 62: 1009-1016, 19 figs., 7 tables
(role in pollination of Delphinium tricorne). —Macior, 1975. Amer. Jour. Bot. 62:
1065-1072, 21 figs., 7 tables (role in pollination of Pedicularis). —Richards, 1976. Kans.
Univ. Sci. Bul. 50: 731-773, 31 figs. (parasitid mites associated with bumblebees).
—Richards and Richards, 1976. Kans. Univ. Sci. Bul. 51: 1-18, 6 figs., 6 tables (relationships
of parasitid mites associated with bumblebees). —Heinrich, 1976. Amer. Sci. 64: 384-395, 14
figs. (foraging behavior and economics of sociality). —Heinrich, 1976. Ecol. Monogr. 46:
105-128 (foraging). —Oster and Heinrich, 1976. Ecol. Monogr. 46: 129-133 (foraging).

Morphology: Cockerell and M’Nary, 1902. Canad. Ent. 34: 71-72 (mouthparts). —Barendrecht,
1931. Acta Zool. 12: 153-204, 28 figs. (corpora pedunculata). —Palm, 1948. VIIIth Internatl.
Congr. Ent., Proc., pp. 289-292, 2 figs. (effect of parasitic action on queens).—

Palm, 1948. Opuse. Ent., Sup. 7: 3-101 (histology of ovaries). —Palm, 1949. Opusce.

Ent. 14: 27-47, 8 figs. (pharyngeal gland). —Medler, 1962. Canad. Ent. 94: 825-833
(development and absorption of eggs). —Kullenberg, Bergstrom and Stallberg-Stenhagen,
1970. Acta Chem. Scandinav. 24: 1481-1482 (volatile components of cephalic marking
secretion of males). —Heinrich, 1972. Jour. Comp. Physiol. 77: 65-79 (patterns of
endothermy in queens, drones and workers). —Heinrich, 1974. Kans. Ent. Soc., Jour. 47:
396-404, 3 figs. (pheromone induced brooding behavior). —Heinrich, 1976. Jour. Expt. Biol.
64: 561-585, 24 figs. (heat exchange).

Genus BOMBUS Subgenus BOMBUS Latreille

Bremus Jurine, 1801. Intell. Blatt. Litt.-Ztg. Erlangen 1: 164. Name suppressed by
Internat]. Comn. Zool. Nomencl., Op. 135, 1939.
Type-species: Apis terrestris Linnaeus. Desig. by Morice and Durrant, 1915.
Bombus Latreille, 1802. Hist. Nat. Fourmis, p. 437.
Type-species: Apis terrestris Linnaeus. Monotypic.
Bremus Panzer, 1804(?). Faunae Ins. German., h. 85.
Type-species: Apis agrorum Fabricius. Desig. by Sandhouse, 1943.
Bombus subg. Leucobombus Dalla Torre, 1880. Naturhistoriker, v. 2, p. 40.
Type-species: Apis terrestris Linnaeus. Desig. by Sandhouse, 1943.
Bombus subg. Terrestribombus Vogt, 1911. Gesell. Naturf. Freunde, Sitzber., p. 55.
Type-species: Apis terrestris Linnaeus. Desig. by Frison, 1927.

This subgenus occurs in the Holarctic Region as well as parts of southeastern and
southwestern Asiaz.

Biology: Hobbs, 1968. Canad. Ent. 100: 156-164 (ecology of south. Alta. spp.).

affinis Cresson. Que. and Ont. south to Ga., west to S. Dak and N. Dak. Ecology: Frequently
usurps colonies of Bombus terricola Kirby. Parasite: Psithyrus ashtoni (Cress.).
Bombus affinis Cresson, 1863. Ent. Soc. Phila., Proe. 2: 103. 6, &.
Bombus affinis var. novae-angliae Bequaert, 1920. Psyche 27: 6. d, §.

2194 Hymenoptera in America North of Mexico

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 518-519, table 18 (synonymy,
redescription). —Milliron, 1971. Ent. Soe. Canada, Mem. 82: 67-72, pl. X, map 4
(redescription, geogr. and floral records).

Biology: Plath, 1922. Psyche 29: 190-192 (nest, parasite). —Plath, 1927. Ent. Soc. Amer., Ann.
20: 188, 191 (hibernation, parasite). —Plath, 1934. Bumblebees and their ways, p. 135 (life
history).

lucorum lucorum (Linnaeus). Holarctic; Alaska southward to parts of south. B. C. and Alta.,

east through Yukon and N. W. T. In addition to the following subspecies, other
subspecies occur in Arctic Eurasia.

Apis lucorum Linnaeus, 1761. Fauna Suecica, Ed. 2, p. 425. d.

Apis cryptarum Fabricius, 1775. System. Ent., p. 379. °.

Bombus moderatus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 109. °.

Bombus terrestris var. schmiedeknechti Verhoeff, 1892. Berlin. Ent. Ztschr. 36: 205.

Terrestribombus'lucorum form magnus Vogt, 1911. Gesell. Naturf. Freunde, Sitzber. p. 56.
Oe

Bombus jacobsoni Skorikov, 1912. Rev. Russ. Ent. 12: 610.

Taxonomy: Milliron, 1971. Ent. Soc. Canada, Mem. 82: 45-50, pl. X, map. 1 (synonymy,
redescription, geogr. and floral records). —Loken, 1973. Norsk Ent. Tidsskr. 20: 40-46, fig.
54 (synonymy, tax. characters, color variation).

lucorum patagiatus Nylander. Northeast. Siberia, Kamchatka Peninsula, east to Alaska

Peninsula (Aleutian Range).

Bombus patagiatus Nylander, 1848. Notiser Sallskapet Fauna Fenn., Forhandl. 1: 234. ?
(or possibly worker).

Bombus viduus Erichson, 1851. In Middendorff, Reise in den Aussersten Norden und
Osten Sibiriens 2: 65.

Bombus albocinctus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 397. °.

Bombus florilegus Panfilov, 1956. Zool. Zhur. 35: 1334.

Taxonomy: Milliron, 1971. Ent. Soc. Canada, Mem. 82: 51 (synonymy, tax. characters, geogr.
range).
terricola terricola Kirby. N. S. to Fla., west to B. C., Mont. and S. Dak. Ecology: Colonies are
frequently usurped by queens of Bombus affinis Cress. Parasite: Psithyrus ashtoni
(Cress.), P. insularis (Sm.), P. suckleyi (Greene).
Bombus terricola Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 273. 2.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 81-83, fig. 17 (redescription,
geogr. records). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 519-520, fig. 133,
table 18 (redescription, geogr. and floral records). —Milliron, 1971. Ent. Soc. Canada, Mem.
82: 52-58, pl. X, map 2 (redescription, geogr. and floral records).

Biology: Plath, 1922. Psyche 29: 192-193 (nest). —Plath, 1927. Psyche 34: 122-124 (nest,
parasite). —Plath, 1927. Ent. Soc. Amer., Ann. 20: 188, 191 (hibernation, parasite). —Plath,
1934. Bumblebees and their ways, p. 138 (life history, parasite).

Morphology: Heinrich, 1972. Jour. Comp. Physiol. 77: 49-64 (energetics of temperature
regulation and foraging).
terricola occidentalis Greene. Alaska south to north. Calif., Nev., Ariz., N. Mex. and S. Dak.

Parasite: Psithyrus fernaldae Franklin, P. insularis (Sm.), P. suckleyi (Greene).
Predator: Philanthus bicinctus (Mickel).

Bombus occidentalis Greene, 1858. Lyc. Nat. Hist. N. Y., Ann. 7: 12. 2, 6.

Bombus modestus Smith, 1861. Jour. of Ent. 1: 153. 9.

Bombus proximus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 98. 9.

Bombus howardi Cresson, 1863. Ent. Soc. Phila., Proc. 2: 99. 6.

Bombus perixanthus Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 389. d.

Bombus mckayi Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 125. 2.

Bombus proximus var. coloradensis Titus, 1902. Canad. Ent. 34: 38. °.

Bombus nigroscutatus Franklin, 1908. In Fletcher and Gibson, Ent. Soc. Ontario, Ann.
Rpt. 39: 111. Nomen nudum.

Superfamily APOIDEA 2195

Bombus occidentalis nigroscutatus Franklin, 1912. Amer. Ent. Soc., Trans. 38: 269, 271. 2,
Oi or

Bremus franklini Frison, 1921. Ent. News 32: 147. °.

Bremus terricola var. severini Frison, 1926. Amer. Ent. Soc., Trans. 52: 139. °.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 71-81, figs. 14, 15
(redescriptions, geogr. records, as occidentalis occidentalis, 0. nigroscutatus, franklini).
—Thorp, 1970. Pan-Pacifie Ent. 46: 177-180 (type locality and Ariz. records of franklin‘).
—Milliron, 1971. Ent. Soc. Canada, Mem. 82: 58-67, map 3 (synonymy, redescription, geogr.
and floral records).

Biology: Plath, 1922. Psyche 29: 192 (nest). —Armitage, 1965. Kans. Ent. Soc., Jour. 38: 97
(predator). —Eshelman and Plowright, 1972. Canad. Ent. 104: 389-398 (nest entrance
recognition). —Wellington, 1974. Science 183: 550-551 (ocellar navigation at dusk).

Genus BOMBUS Subgenus FRATERNOBOMBUS Skorikov

Alpigenobombus subg. Fraternobombus Skorikov, 1922. Sta. Region. Protect. Plantes,
Petrograd Bul. 4: 156.
Type-species: Apathus fraternus Smith. Desig. by Frison, 1927.

This subgenus occurs only jn North and Central America.

fraternus (Smith). N. J. to Fla., west to N. Dak., S. Dak., Nebr., Colo. and N. Mex. Predator:
Promachus hinei Bromley.
Apathus fraternus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 385. d.
Bombus scutellaris Cresson, 1863. Ent. Soc. Phila., Proc. 2: 96. 2, d, 9.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 526-527, figs. 131, 132, table
18 (redescription). —Milliron, 1973. Ent. Soc. Canada, Mem. 91: 243-246, pl. XVI, map 36
(redescription, geogr. and floral records).

Biology: Thorp, 1973. Pan-Pacific Ent. 49: 89 (predator).
Genus BOMBUS Subgenus BOMBIAS Robertson

Bombias Robertson, 1903. Amer. Ent. Soc., Trans. 29: 176.
Type-species: Bombias auricomus Robertson. Orig. desig. 4
Nevadensibombus Skorikov, 1922. Sta. Region. Protect. Plantes, Petrograd Bul. 4: 149.
Type-species: Bombus nevadensis Cresson. Desig. by Frison, 1927.

This subgenus is present only in North America.

Biology: Hobbs, 1965. Canad. Ent. 97: 120-128 (ecology).
nevadensis auricomus (Robertson). Ont. to Fla., west to Tex., Okla., Colo., Wyo., Mont. and
south. Canada (Sask., Alta. and B. C.). Parasite: Brachicoma sarcophagina (Twns.),
Physocephala sagittaria (Say), Psithyrus citrinus (Sm.).
Bombias auricomus Robertson, 1903. Amer. Ent. Soc., Trans. 29: 176. 2, d, 9.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 520-521, table 18 (tax. status,
redescription). —Milliron, 1971. Ent. Soc. Canada, Mem. 82: 77-80, map 6 (redescription,
geogr. and floral records). °

Biology: Frison, 1917. Ent. Soc. Amer., Ann. 10: 277-286, pls. 23, 24 (life history, parasites).
—Frison, 1918. Ent. Soc. Amer., Ann. 11: 43-48, pl. 3 (life history, parasite). —Howard,
1918. Ent. News 29: 114-115 (nest). —Plath, 1927. Psyche 34: 127 (nesting habits). —Plath,
1934. Bumblebees and their ways, p. 153 (life history).

nevadensis nevadensis Cresson. Alaska south to Calif., Ariz., N. Mex. and east to Wis.; Mexico

(Hidalgo). Parasite: Physocephala marginata (Say), P. texana (Will.), Psithyrus
insularis (Sm.), P. suckleyi (Greene).

Bombus nevadensis Cresson, 1874. Amer. Ent. Soc., Trans. 5: 102. 2, d.

Bombus improbus Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 186. d.

Bombus nevadensis race cressoni Cockerell, 1899. Ann. and Mag. Nat. Hist. (7) 4: 388. °.

Bombus nevadensis miguelensis Cockerell, 1937. Pan-Pacific Ent. 13: 148. d.

Bombus crotchii semisuffusus Cockerell, 1937. Pan-Pacific Ent. 13: 148. °.

2196 Hymenoptera in America North of Mexico

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 44-49, 61, fig. 10, map 5
(redescriptions, as nevadensis nevadensis, n. miguelensis, crotchii semisuffusus).
—Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 521-523, figs. 129, 130, table 18
(redescription). —Milliron, 1971. Ent. Soc. Canada, Mem. 82: 73-77, pl. XI, map 5
(synonymy, redescription, geogr. and floral records).

Genus BOMBUS Subgenus SEPARATOBOMBUS Frison

Bremus subg. Separatobombus Frison, 1927. Amer. Ent. Soc., Trans. 53: 64.
Type-species: Apis griseocollis Degeer. Orig. desig. (=Bombus separatus Cresson).

This subgenus is endemic in North America.

griseocollis (Degeer). Que. south to Fla., west to B. C., Wash., Oreg. and north. Calif.
Apis griseocollis Degeer, 1773. Mem. Serv. Hist. Insectes, v. 3, p. 576.
(?)virginica Olivier, 1789. Encycl. Meth., v. 4, p. 66.
(?)virginica Fabricius, 1793. Ent. System., v. 2, p. 318.
(?)virginicus Fabricius, 1804. Systema Piezatorum, p. 346.
Bombus separatus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 165. 2, 6, ¥.
Bombus mormonorum Franklin, 1911. Amer. Ent. Soe., Trans. 37: 161. 9, 3, §.
Bremus separatus var. nero Bequaert and Plath, 1925. Mus. Compar. Zool., Bul. 67: 275. &.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Tech. Bul. 40: 50-54, fig. 18, map 6 (synonymy,
redescription, geogr. records). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
527-529, figs. 129, 132, table 18 (synonymy, redescription, geogr. range and floral records).
—Milliron, 1973. Ent. Soc. Canada, Mem. 91: 247-252, pl. XVI, fig. 37 (synonymy,
redescription, geogr. and floral records).

Biology: Plath, 1927. Psyche 34: 125-127 (life history, as separatus). —Plath, 1927. Ent. Soc.
Amer., Ann. 20: 190 (hibernation, as separatus). —Plath, 1934. Bumblebees and their
ways, p. 155 (life history, as separatus). —Eshelman and Plowright, 1972. Canad. Ent.
104: 389-398 (nest entrance recognition).

Morphology: Stephen and Koontz, 1973. Melanderia 13: 13-29, 50 figs. (developmental
changes in preadult stages).

morrisoni Cresson. B. C. to Calif., east to S. Dak., Nebr., Colo. and N. Mex. Parasite:
Monodontomerus montivagus Ashm.
Bombus morrisoni Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 1838. 2, d, 9.
Bombus morrisoni var. umbrosus Friese, 1931. Konowia 10: 301.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 54-57, fig. 12, map 6
(redescription, geogr. records). —Milliron, 1973. Ent. Soc. Canada, Mem. 91: 252-256, pl.
XVI, map 38 (synonymy, redescription, geogr. and floral records).

Biology: Bohart and Knowlton, 1952. Econ. Ent., Jour. 45: 890-891 (yearly population
fluctuations).

Genus BOMBUS Subgenus CROTCHIIBOMBUS Franklin

Bombus subg. Crotchiibombus Franklin, 1954. Amer. Ent. Soc., Trans. 80: 51.
Type-species: Bombus crotchii Cresson. Orig. desig.

Endemic in western North America.

crotchii Cresson. Calif.; Mexico (Baja California).
Bombus crotchii Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 184. 9.
Bombus nigrocinctus Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 342. 2.
Bremus crotchii var. nigricaudus Frison, 1927. Calif. Acad. Sci., Proc. (4) 16: 375. @.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 57-61, fig. 15, map 7
(synonymy, redescription, geogr. records). — Milliron, 1973. Ent. Soc. Canada, Mem. 91:
256-258, pl. XVI, map 39 (synonymy, redescription, geogr. records).

Genus BOMBUS Subgenus CULLUMANOBOMBUS Vogt
Bombus subg. Cullwmanobombus Vogt, 1911. Gesell. Naturf. Freunde, Sitzber. p. 57.

Superfamily APOIDEA 2197

Type-species: Apis culluwmanus Kirby. Desig. by Frison, 1927.
Bremus subg. Rufocinctobombus Frison, 1927. Amer. Ent. Soe., Trans. 53: 78, pl. xvii, fig.
9
Type-species: Bombus rufocinctus Cresson. Monotypic.

Species of this subgenus are found in the Palaearctic, Nearctic and northern Neotropical Re-
gions.

Biology: Hobbs, 1965. Canad. Ent. 97: 1293-1302 (ecology).
rufocinctus Cresson. N.S., N. B. and Que., west to B. C., south to Calif., Ariz., N. Mex., Kans.,

Minn., Ill., Mich., N. Y., Vt. and Maine; Mexico (Distrito Federal, Hidalgo, Mexico,
Michoacan, Morelos and Sonora). Parasite: Physocephala texana (Will.), Psithyrus
fernaldae Franklin, P. insularis (Sm.), P. suckleyi (Greene). Predator: Philanthus
bicinctus (Mickel).

Bombus rufocinctus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 106. 6, 9.

Bombus iridis Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 390. @.

Bombus prunellae Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 391. 9.

Bombus iridis var. phaceliae Cockerell, 1906. Canad. Ent. 38: 160. °.

Bombus rufocinctus var. astragali Cockerell, 1907. Entomologist 40: 97. 2.

Bombus hyperboreus var. albertensis Cockerell, 1909. Canad. Ent. 41: 36. 8.

Bombus (Bombias) mexicensis Franklin, 1911. Amer. Ent. Soc., Trans. 37: 163. 2.

Bombus henshawi Franklin, 1913. Amer. Ent. Soc., Trans. 38: 446. °.

Bombus rufocinctus var. castoris Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 15: 537. d.

Bremus rufocinctus var. sladeni Frison, 1926. Amer. Ent. Soc., Trans. 52: 138. 6. Nomen
nudum.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 62-70, figs. 3, 11, table 1
(synonymy, redescription, variation in color pattern, geogr. records). —Thorp, 1962.
Pan-Pacific Ent. 38: 24-25 (synonymy, geogr. records from and near the type locality of
henshawi). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 529-530, fig. 132, table
18 (redescription). —Milliron, 1973. Ent. Soc. Canada, Mem. 91: 317-326, pl. XIX, map 34
(synonymy, redescription, geogr. and floral records).

Biology: Armitage, 1965. Kans. Ent. Soc., Jour. 38: 97 (predator).

Genus BOMBUS Subgenus PYROBOMBUS Dalla Torre

Bombus subg. Pyrobombus Dalla Torre, 1880. Der Naturhistoriker 2: 40.
Type-species: Apis hypnorum Linnaeus. Monotypic.
Bombus subg. Pyrrhobombus Dalla Torre, 1882. Naturw.-med. Ver. Innsbruck, Ber. 12: 28.
Emendation of Pyrobombus Dalla Torre.
Bombus subg. Poecilobombus Dalla Torre, 1882. Naturw.-med. Ver. Innsbruck, Ber. 12: 23.
Type-species: Bombus sitkensis Nylander. Desig. by Sandhouse, 1943.
Bombus subg. Pratobombus Vogt, 1911. Gesell. Naturf. Freunde, Sitzber., p. 49.
Type-species: Apis pratorum Linnaeus. Desig. by Frison, 1927.
Bombus subg. Hypnorubombus Quilis Perez, 1927. Lab. Hist. Nat. Valencia, Trabhs. 16: 19.
Lapsus.
Bombus subg. Hypnorobombus Quilis Perez, 1927. Lab. Hist. Nat. Valencia, Trabhs. 16: 97.
Type-species: Apis hypnorum Linnaeus. Monotypic. (~Bombus hypnorum

(Linnaeus)).
Bombus subg. Lapponicobombus Quilis Perez, 1927. Lab. Hist. Nat. Valencia, Trabhs. 16:
19, 22, 63.

Type-species: Apis lapponica Fabricius. Desig. by Milliron, 1961. (=Bombus
lapponicus (Fabricius)).
This is a large assemblage of species which occur principally in the Holarctic Region, but a
few species extend into Central America and into the East Indies.
Biology: Hobbs, 1967. Canad. Ent. 99: 1271-1292 (ecology of south. Alta. spp.).
bifarius bifarius Cresson. B. C., Oreg. (Steens Mts.), Calif. (Sierra Nevada Mts.), Idaho, Utah,

Colo. Predator: Philanthus bicinctus (Mickel).
Bombus bifarius Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 185. 2, %.

2198 Hymenoptera in America North of Mexico

Bombus vancouverensis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 187. d.

Bombus cooleyi Morrill, 1903. Canad. Ent. 35: 222.

Bombus edwardsii var. kenoyeri Cockerell, 1915. Ann. and Mag. Nat. Hist. (8) 16: 483. @.
Bombus bifarius var. arctostaphyli Cockerell, 1930. Ann. and Mag. Nat. Hist. (10) 5: 405.

é.
Bombus edwardsii var. fuscifrons Swenk, 1938. Pan-Pacific Ent. 14: 30. 2, &.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 139-145, figs. 5, 29, map 18
(synonymy, redescription, tax. status, variation in color pattern, geogr. records).

Biology: Armitage, 1965. Kans. Ent. Soc., Jour. 38: 97 (predator).
bifarius nearcticus Handlirsch. Alaska and Yukon, south to Calif. (Sierra Nevada Mts.) and
Utah. Parasite: Psithyrus insularis (Sm.), Volucella bombylans (L.).
Bombus nearcticus Handlirsch, 1888. K. K. Naturhist. Hofmus., Ann. 3: 243. 2, 6, &.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 139-141, 145-148, figs. 5, 29,
map 18 (tax. status, redescription, variation in color pattern, geogr. records).
bimaculatus Cresson. Ont. and Maine, south to Fla., west to IIl., Kans., Okla. and Miss.
Parasite: Brachicoma sarcophagina (Twns.).
Bombus bimaculatus Cresson, 1863. Ent. Soc. Phila., Proce. 2: 92. 6.
Bombus ridingsii Cresson, 1878. Acad. Nat. Sci. Phila., Proc., p. 182. 2, &.
Bremus bimaculatus var. ahenus Bequaert and Plath, 1925. Mus. Compar. Zool., Bul. 67:

2D ONG) é
Bremus bimaculatus var. arboreti Bequaert and Plath, 1925. Mus. Compar. Zool., Bul. 67:
216" er
Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 531-532, table 18
(redescription).

Biology: Plath, 1922. Psyche 29: 194-195 (life history). —Plath, 1927. Ent. Soc. Amer., Ann.
20: 189 (hibernation). —Frison, 1928. Ent. Amer. (n. s.) 8: 159-223 (life history). —Plath,
1934. Bumblebees and their ways, p. 141 (life history). —Townsend, 1936. Ent. Soc. Wash.,
Proc. 38: 92-98 (parasites).

caliginosus (Frison). Wash., Oreg. and Calif. Ecology: Generally inhabits more coastal areas,
but the known southernmost populations occur in the San Jacinto Mts. of south. Calif;
the only known nest was located in an old bird nest some 15 feet above ground.
Bremus caliginosus Frison, 1927. Calif. Acad. Sci., Proc. (4) 16: 376. 6.
Bremus caliginosus var. tardus Frison, 1927. Calif. Acad. Sci., Proc. (4) 16: 380. d.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 100-104, fig. 20, map 13
(synonymy, redescription, geogr. records).

Biology: Pitelka, 1954. Pan-Pacific Ent. 30: 220 (nest).
centralis (Cresson). B. C. and Alta. south to Calif., Ariz. and N. Mex. Predator: Philanthus
bicinctus (Mickel).
Bombus centralis Cresson, 1864. Ent. Soc. Phila., Proc. 3: 41. 2.
Bombus juxtus Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 187. 2.
Bombus monardae Cockerell and Porter, 1899. Ann. and Mag. Nat. Hist. (7) 4: 387. 9.
Bremus centralis var. fucatus Frison, 1929. Amer. Ent. Soc., Trans. 55: 107. @.
Bremus centralis var. stolidus Frison, 1929. Amer. Ent. Soc., Trans. 55: 107. @.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 95-100, fig. 19, map 12
(synonymy, redescription, color variation of vestiture, geogr. records).

Biology: Armitage, 1965. Kans. Ent. Soc., Jour. 38: 97 (predator).
cockerelli Franklin. N. Mex., Utah.
Bombus cockerelli Franklin, 1913. Amer. Ent. Soc., Trans. 38: 356. 2, %.
edwardsii Cresson. Oreg., Calif., Nev. (Douglas and Washoe Counties).
Bombus edwardsii Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 184. 2, 8.
Bombus fernaldi Franklin, 1911. Amer. Ent. Soc., Trans. 37: 157. 2, 6, 8.
Bombus lapponicus var. insularis Friese, 1924. Deut. Ent. Ztschr., p. 487. °.

Superfamily APOIDEA 2199

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 109-113, fig. 22, map 10
(redescription, geogr. and floral records).

Biology: Linsley and Michener, 1942. Pan-Pacific Ent. 18: 29 (nest, vernal flights).

Morphology: Heinrich, 1974. Kans. Ent. Soc., Jour. 47: 396-404, 3 figs. (pheromone induced
brooding behavior).
flavifrons dimidiatus Ashmead. South. B. C. to Calif.
Bombus dimidiatus Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 129. 2, §.
Bombus ambiguus Franklin, 1911. Amer. Ent. Soc., Trans. 37: 159. 2, 9.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 85-87, 92-95, map 11
(synonymy, tax. status, redescription, geogr. records).
flavifrons flavifrons Cresson. Alaska south to Calif., Idaho and Utah. Parasite: Psithyrus
insularis (Smith). Predator: Philanthus bicinctus (Mickel).
Bombus flavifrons Cresson, 1863. Ent. Soc. Phila., Proce. 2: 105. 2, 6, %.
Bombus alaskensis Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 128. 2, &.
Bombus flavifrons var. veganus Cockerell, 1903. Amer. Nat. 37: 891. ¢.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 85-92, fig. 18, map. 11
(redescription, geogr. records).

Biology: Sladen, 1915. Canad. Ent. 47: 84 (parasite). —Armitage, 1965. Kans. Ent. Soc., Jour.
38: 97 (predator).

Morphology: Milliron, 1962. Brooklyn Ent. Soc., Bul. 57: 45-46 (gynandromorph).
frigidus Smith. Alaska and N. W. T., south at high elevations to Colo.
Bombus frigidus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 399. @, 6.
Bombus carriei Greene, 1860. Lye. Nat. Hist. N. Y., Ann. 7: 170. @.
Bombus couperi Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 185. 2.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 124-126, fig. 25 (synonymy,
redescription, geogr. range).
frigidus var. alboanalis Franklin. Man., Alaska.
Bombus alboanalis Franklin, 1913. Amer. Ent. Soc., Trans. 38: 385. 2, §.
huntii Greene. B. C. and Alta., south to Calif., Nev., Utah and N. Mex.
Bombus huntii Greene, 1860. Lye. Nat. Hist. N. Y., Ann. 7: 172. 9.
Bombus rufosuffusus Cockerell, 1905. Ent. News 16: 271. °.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 135-139, fig. 28, map 17
(synonymy, redescription, geogr. records).

Biology: Medler, 1959. Ent. News 70: 179-182 (nest).
impatiens Cresson. Ont. and Maine, south to Fla., west to Mich., Ill., Kans. and Miss. Parasite:
Psithyrus citrinus (Smith). Predator: Mallophora orcina (Wied.).
Bombus impatiens Cresson, 1863. Ent. Soc. Phila., Proc. 2: 90. d.
Bombus impatiens var. deayi Chandler, 1956 (1955). Ind. Acad. Sci., Proc. 65: 116. °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 532-533, fig. 183, table 18
(synonymy, redescription, floral records).

Biology: Plath, 1922. Psyche 29: 195-197 (life history). —Plath, 1927. Ent. Soc. Amer., Ann.
20: 187-188, 189 (hibernation). —Frison, 1929. Brooklyn Ent. Soe., Bul. 24: 261-285
(bionomics). —Fattig, 1933. Canad. Ent. 65: 119-120 (asilid predation). —Plath, 1934.
Bumblebees and their ways, p. 144 (life history). —Townsend, 1951. Ent. News 62: 115-116
(hibernation).

melanopygus Nylander. Alaska, south to north. Calif., Idaho and Colo. Predator: Philanthus
bicinctus (Mickel).
Bombus melanopyge Nylander, 1848. Notiser Sallskapet Fauna Fenn., Forhandl. 1: 236. d
2Bombus menestriesii Radoszkowski, 1859. Soc. Nat. Moscou, Bul. 32: 483. 9, d.
Bombus lacustris Cresson, 1863. Ent. Soe. Phila., Proc. 2: 108. 2, d, &.
Bremus melanopygus var. washingtonensis Frison, 1926. Amer. Ent. Soc., Trans. 52: 138.
Oe tele

2200 Hymenoptera in America North of Mexico

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 132-135, fig. 27, map 16
(synonymy, redescription, geogr. records).

Biology: Armitage, 1965. Kans. Ent. Soc., Jour. 38: 97 (predator).
mixtus Cresson. Alaska, south to Calif., Idaho and Colo. Predator: Philanthus bicinctus
(Mickel).
Bombus mixtus Cresson, 1878. Acad. Nat. Sci. Phila., Proc., p. 186. 2, 8.
Bremus edwardsii var. russulus Frison, 1927. Calif. Acad. Sci., Proc. (4) 16: 374. %.

Taxonomy: Stephen, 1957. Oreg. Expt. Sta. Tech. Bul. 40: 113-120, fig. 23, map 15 (synonymy,
redescription, geogr. records).
perplexus Cresson. Alaska to Maine, south to Wis., Ill. and Fla., ?Alta. Parasite: ?Psithyrus
fernaldae Franklin.
Bombus perplexus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 91. 3.
Bombus hudsonicus Cresson, 1863. Ent. Soe. Phila., Proc. 2: 92. 6.
Apathus dorsalis Provancher, 1888. Addit. Corr. Faune Ent. Canada, Hym., p. 343. °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 533-534, table 18 (synonymy,
redescription, geogr. range, floral records).

Biology: Plath, 1927. Psyche 34: 124-125 (nest). —Plath, 1927. Ent. Soc. Amer., Ann. 20: 189

(hibernation). —Plath, 1934. Bumblebees and Their ways, p. 146 (life history).
pleuralis Nylander. Rocky Mountain States, B. C., N. W. T., Yukon, Alaska.

Bombus pleuralis Nylander, 1848. Notiser Sallskapet Fauna Fenn., Forhandl. 1: 231. 2, 6.
pleuralis var. clarus (Frison). B. C., N. W. T., Alaska.

Bremus pleuralis var. clarus Frison, 1926. Amer. Ent. Soc., Trans. 52: 139. 2, 6, 9.
sandersoni Franklin. Ont. to Newfoundland, south to Tenn. and N. C.

Bombus vagans sandersoni Franklin, 1913. Amer. Ent. Soc., Trans. 38: 353. °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 534-535 (tax. status,
redescription).
sitkensis Nylander. Alaska, south to Calif., Idaho, Mont. and Wyo.
Bombus sitkensis Nylander, 1848. Notiser Sallskapet Fauna Fenn., Forhandl. 1: 235. 2, 6.
Bombus oregonensis Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 185. d.
Bombus mixtuosus Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 128. 2, 8.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 120-124, fig. 24, map 14
(redescription, geogr. records).
sylvicola Kirby. Alaska east to Newfoundland, south on the principal cordillera of west. U.S.
(Cascade, Sierra Nevada, Great Basin and Rocky Mts.) to Calif., Nev., Utah and N. Mex.
Bombus sylvicola Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 272.
Bombus gelidus Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 184. ?.
Bremus sylvicola var. sculleni Frison, 1929. Amer. Ent. Soc., Trans. 55: 108. 9.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 126-130, fig. 26, map 16
(synonymy, redescription, variation, geogr. records). —Thorp, 1962. Pan-Pacific Ent. 38:
21-23 (tax. status, geogr. records, variation in colorational pattern). —Milliron, 1971. Ent.
Soe. Canada, Mem. 82: 42 (tax. status). —Loken, 1973. Norsk Ent. Tidsskr. 20: 76 (tax.
status). =

sylvicola var. johanseni Sladen. N. W. T., Baffin Land.
Bombus sylvicola var. johanseni Sladen, 1919. Rpt. Canad. Arctic Exped. 1913-18, v. 3, p.
30g. 2, %.
sylvicola var. lutzi (Frison). Ariz.
Bremus sylvicola var. lutzi Frison, 1923. Amer. Ent. Soc., Trans. 48: 309. °.

Taxonomy: Thorp, 1970. Pan-Pacifie Ent. 46: 177, 180 (validity of Ariz. record).
ternarius Say. Yukon east to N. S., south to Ga., Mich., Kans., Mont. and B. C. Parasite:
Psithyrus insularis (Smith).
Bombus ternarius Say, 1837. Boston Jour. Nat. Hist. 1: 414. ¢.
?Bombus ornatus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 398. 2, 6, &.

Superfamily APOIDEA 2201

Bombus ternarius var. expallidus Cockerell, 1916. Mich. Univ. Mus. Zool., Occas. Papers
Zoo were

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 149-150, fig. 30, map 17
(redescription, variation in color of vestiture, geogr. records). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 536-537, table 18 (synonymy, redescription ).

Biology: Plath, 1927. Psyche 34: 125 (life history). —Plath, 1927. Ent. Soc. Amer., Ann. 20:
189-190 (hibernation). —Plath, 1934. Bumblebees and their ways, p. 149 (life history).
—Craig, 1953. Canad. Ent. 85: 311-312 (parasite).

vagans bolsteri Franklin. Newfoundland.
Bombus bolsteri Franklin, 1913. Amer. Ent. Soc., Trans. 38: 357. 2, 9.
vagans vagans Smith. B. C. east to N. S., south to Ga., Tenn., S. Dak., Mont., Idaho and Wash.
Parasite: Brachicoma sarcophagina (Twns.), Psithyrus citrinus (Smith), Spherularia
bombi Dufour.
Bombus vagans Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 399. °.
Bombus consimilis Cresson, 1864. Ent. Soe. Phila., Proc. 3: 41. °.
Bremus vagans var. coctus Bequaert and Plath, 1925. Mus. Compar. Zool., Bul. 67: 276. °,
3.
Bremus vagans var. helenae Frison, 1929. Amer. Ent. Soc., Trans. 55: 110. 2, d.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 104-106, fig. 21, map 14
(redescription, geogr. records). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152:
537-538 (synonymy, redescription, floral records).

Biology: Plath, 1922. Psyche 29: 197-198 (life history). —Plath, 1927. Ent. Soc. Amer., Ann.
20: 190 (hibernation). —Frison, 1930. Brooklyn Ent. Soc., Bul. 25: 109-122 (life history).
—Plath, 1934. Bumblebees and their ways, p. 151 (life history).

Morphology: Heinrich, 1972. Science 175: 185-187, 2 figs. (temperature regulation).
vandykei (Frison). Wash. to south. Calif.
Bremus flavifrons var. vandykei Frison, 1927. Calif. Acad. Sci., Proc. (4) 16: 375. 8.
Pyrobombus (Pyrobombus) cascadensis Milliron, 1970. Canad. Ent. 102: 382. 2, 9. N. syn.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 106-109. $, 2 (tax. characters,
tax. status, geogr. records). —Thorp, 1969. Pan-Pacific Ent. 45: 87-96. 2, %, d (tax. status,
redescription, key to related spp., geogr. and floral records).

vosnesenskii Radoszkowski. B. C. south to Calif., Nev. (Washoe County); Mexico (Baja
California).
Bombus vosnesenskii Radoszkowski, 1862. Soc. Nat. Moscou, Bul. 35: 589. 2, ¢.
Bombus columbicus Dalla Torre, 1890. Wien. Ent. Ztg. 9: 139.

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1089, fig. 254 (larva). —Stephen, 1957.
Oreg. Agr. Expt. Sta. Tech. Bul. 40: 151-155, fig. 31, map 13 (redescription, geogr. records).

Biology: Hicks, 1929. Pan-Pacific Ent. 5: 97-100 (nest).
Morphology: Heinrich, 1972. Nature 239: 223-225 (physiology of brood incubation).
—Heinrich, 1974. Jour. Comp. Physiol. 88: 129-140 (brood incubation). —Heinrich, 1974.

Kans. Ent. Soc., Jour. 47: 396-404, 3 figs. (pheromone induced brooding-behavior).
—Heinrich, 1975. Jour. Comp. Physiol. 96: 155-166, 9 figs. (thermoregulation).

Genus BOMBUS Subgenus ALPINOBOMBUS Skorikov

Alpinobombus Skorikov, 1914. Rev. Russe d’Ent. 14: 122.
' Type-species: Apis alpinus Linnaeus. Desig. by Frison, 1927.

Members of this subgenus occur in the Alps, Arctic Eurasia and America, Greenland and the
principal cordillera of western North America (Rocky Mountains, Cascades, Sierra Nevada Mts.
and some of the Great Basin Mts.). Some of the species are circumpolar in distribution and at
least one of the species (Bombus hyperboreus Schonherr) apparently produces no workers when
it usurps the nests of other species.

Taxonomy: Loken, 1973. Norsk Ent. Tidsskr. 20: 94-118, figs. (Scandanavian spp.).

2202 Hymenoptera in America North of Mexico

Biology: Hobbs, 1964. Canad. Ent. 96: 1465-1470 (ecology of south. Alta spp.). —Milliron and
Oliver, 1966. Canad. Ent. 98: 207-213, 6 figs. (usurpation). —Richards, 1973. Quaest. Ent. 9:
115-157, 29 figs., 10 tables (life history).

balteatus Dahlbom. Holarctic; Arctic Alaska and Canada, south on principal cordillera of west.

N. Amer. to Calif. (Sierra Nevada Mts. and White Mts.) and N. Mex. (Truchas Peak),
? Ariz. (Patagonia Mts.).

Bombus balteatus Dahlbom, 1832. Bombi Scand., p. 36. °.

Bombus nivalis Dahlbom, 1832. Bombi Scand., p. 40. 9.

Bombus tricolor Dahlbom, 1832. Bombi Seand, p. 41. °.

Bombus kirbiellus Curtis, 1834. Deser. insects brought home by Comdr. James Clark Ross
second voyage, App. Nat. Hist., p. 62. 2, gd, 9.

Bombus kirbyellus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 397. Emend.

Bombus putnami Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 185. 9.

Psithyrus kodiakensis Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 130. ¢.

Bombus kirbyellus var. pyrrhopygus Friese, 1902. In Romer and Schaudinn’s Fauna
Arctica 2: 495.

Bombus kirbyellus var. tristis Sparre-Schneider In Friese, 1902. In Romer and
Schaudinn’s Fauna Arctica 2: 495. Preocce.

Bombus atrifasciatus Morrill, 1903. Canad. Ent. 35: 224. 2.

Bombus kirbyellus var. lysholmi Friese, 1905 (1904). Mus. Zool. Acad. Imp. Sci. St.
Petersbourg, Ann. 9: 519.

?Bombus kirbyellus var. friesei Skorikov, 1908 (1907). Russ. Ent. Obozr. 7: 111.

Bombus kirbyellus var. cinctus Friese, 1911. Deut. Ent. Ztschr. 4: 456. Preocc.

Bombus kirbyellus var. cinctellus Friese, 1911. Deut. Ent. Ztschr. 4: 456.

Bombus kirbyellus var. similis Friese, 1911. Deut. Ent. Ztschr. 4: 684. N. name.

Alpinobombus kirbyellus appropinquans Skorikov, 1914. Russ. Ent. Obozr. 14: 123.

Alpinobombus kirbyellus var. subcollaris Skorikov, 1914. Russ. Ent. Obozr. 14: 123.

Alpinobombus kirbyellus var. gmelini Skorikov, 1914. Russ. Ent. Obozr. 14: 124.

Bombus kirbyellus var. semijaensis Friese, 1923. In Norwegian Exped. Novaya Zemlya
1921, Rpt. Sci. Results, 14: 4.

Bremus kirbyellus var. alexanderi Frison, 1923. Amer. Ent. Soc., Trans. 48: 308. °.

Bremus kirbyellus var. arizonensis Frison, 1923. Amer. Ent. Soc., Trans. 48: 309. °.

Taxonomy: Thorp, 1962. Pan-Pacifie Ent. 38: 23-24 (tax. characters, geogr. records in Calif.).
—Thorp, 1970. Pan-Pacifie Ent. 46: 177 (validity of Ariz. records). —Milliron, 1973. Ent.
Soc. Canada, Mem. 89: 85-89, 94-107, figs. 8, 9 (synonymy, redescription, geogr. records, as
balteatus, kirbyellus, and polaris in part). —Loken, 1973. Norsk Ent. Tidsskr. 20: 105-114,
figs. 26, 69, 70, tables VI-IX (synonymy, tax. characters, tax. status, variation in color of
vestiture).

Biology: Hobbs, 1964. Canad. Ent. 96: 1465-1470, 6 figs. (life history). —Loken, 1973. Norsk

Ent. Tidsskr. 20: 112-114 (life history).

hyperboreus Schonherr. Holarctic (cireumpolar); Arctic Alaska and Canada (Yukon and N. W.

T), Greenland. Ecology: Usurps colonies of Bombus polaris Curtis, B. jonellus (Kirby)
and possibly other species either as a casual or a facultative inquiline.

?Apis arctica Quensel in Acerbi, 1802. Travels through Sweden, Finland and Lapland to
the North Cape in the years 1798 and 1799, v. 2, p. 253, pl. 1, fig. 7.

Bombus hyperboreus Schonherr, 1809. Svenska Vetensk. Akad., Handl. 30: 57. °.

Bombus hyperboreus var. natvigi Richards, 1931 (1927). Tromso Mus. Arshefter 50, no. 6,
195 Gh Cc

Bombus hyperboreus var. vulpinus Friese, 1935. Skr. Svalbard Ishavet 65: 4.

Alpinibombus(!) hyperboreus eskimo Skorikov, 1937. Ent. Meddel. 20: 57. ¢.

Alpinibombus(!) hyperboreus eskimo mod. henrikseni Skorikov, 1937. Ent. Meddel. 20: 57.
é.

Taxonomy: Richards, 1931 (1927). Tromso Mus. Arshefter 50, no. 6, p. 10 (tax. status).
—Milliron, 1973. Ent. Soc. Canada, Mem. 89: 89-94, pl. XII, map 7 (synonymy,
redescription, geogr. records). —Loken, 1973. Norsk Ent. Tidsskr. 20: 114-118, figs. 27, 71
(synonymy, nomenclature).

Superfamily APOIDEA 2203

Biology: Milliron and Oliver, 1966. Canad. Ent. 98: 207-213, 6 figs. (nests, nesting habits,
usurpation). —Richards, 1973. Quaest. Ent. 9: 115-157, 29 figs., 10 tables (nesting habits,
colony development, floral relationships, nest-parasitism, adaptation to arctic
environment). —Loken, 1973. Norsk. Ent. Tidsskr. 20: 117-118 (usurpation).

polaris polaris Curtis. Holarctic (cireumpolar); Arctic Alaska, Canada, Greenland and parts of

Arctic Eurasia. Ecology: Colonies are sometimes usurped by queens of Bombus
hyperboreus Schonherr. Other subspecies, such as Bombus polaris diabolicus Friese,
occur in parts of Arctic Eurasia.

Bombus arcticus Kirby, 1821. Account of animals seen by northern expedition within
Arctic Circle, p. 216. 2, ¢. Preoce.

Bombus polaris Curtis, 1834. Deser. insects brought home by Comdr. James Clark Ross
second voyage, App. Nat. Hist., p. 62. 2, 6.

Bombus Groenlandicus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 393. 2.

Bombus kincaidii Cockerell, 1898. Ann. and Mag. Nat. Hist. (7) 2: 324. 2, 6.

Alpinibombus(!) arcticus mod. natvigioides Skorikov, 1937. Ent. Meddel. 20: 56, 57. Nomen
nudum.

Taxonomy: Yarrow, 1955. Ann. and Mag. Nat. Hist. (12) 8: 152 (tax. status, as arcticus var
polaris). —Milliron, 1973. Ent. Soc. Canada, Mem. 89: 98-107, pl. XII, map 9 (synonymy,
redescription, geogr. records, in part, includes balteatus in part). —Loken, 1973. Norsk
Ent. Tidsskr. 20: 101-104, figs. 25, 68 (tax. status, nomenclature, synonymy, geogr. range,
as arcticus).

Biology: Milliron and Oliver, 1966. Canad. Ent. 98: 207-213, 6 figs. (nest, nesting habits,
usurpation). —Ri¢hards, 1973. Quaest. Ent. 9: 115-157, 29 figs., 10 tables (nesting habits,
artificial domiciles, colony development, floral relationships, usurpation, adaptation to
arctic environment). —Loken, 1973. Norsk Ent. Tidsskr. 20: 104 (nests).

strenuus Cresson. Alaska, Yukon, N. W. T., south to B. C., ?Alta.
Bombus strenuus Cresson, 1863. Ent. Soe. Phila., Proc. 2: 102. 2, d.
Bombus neoboreus Sladen, 1919. Rpt. Canad. Arctic Exped. 1913-18, v. 3, p. 28g. 2, d, 9.
Bombus (Alpinobombus) hyperboreus clydensis Yarrow, 1955. Ann. and Mag. Nat. Hist.
G2)I8 F150 Oa

Taxonomy: Milliron, 1973. Ent. Soc. Canada, Mem. 89: 108-111, pl. XII, map 10 (synonymy,
redescription, geogr. records).

Genus BOMBUS Subgenus SUBTERRANEOBOMBUS Vogt

Bombus subg. Subterraneobombus Vogt, 1911. Gesell. Naturf. Freunde, Sitzber., p. 62.
Type-species: Apis subterraneus Linnaeus. Desig. by Frison, 1927.

Species of this subgenus occur in Eurasia as far south as the Himalayas and also are found in
western North America north of Mexico.

Biology: Hobbs, 1966. Canad. Ent. 98: 288-294 (ecology of south. Alta. spp.).
appositus Cresson. B. C. east to Sask., south to N. Mex., Ariz. and Calif. (Cascades and Sierra
Nevada Mts.). Parasite: Melittobia chalybii Ashm., Physocephala texana (Will.),
Psithyrus fernaldae Franklin, P. insularis (Sm.), P. suckleyi (Greene).
Bombus appositus Cresson, 1878. Acad. Nat. Sci. Phila., Proc. p. 183. 2, 5, %.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 22-26, fig. 6, map 1
(redescription, color variation of vestiture, geogr. records). —Milliron, 1973. Ent. Soc.
Canada, Mem. 89: 113-117, pl. XII, map 11 (redescription, geogr. records).

borealis Kirby. South. Canada from N. S. to Alta. and north. U. S. from Maine to N. J., west to
N. D. and S. Dak.
Bombus borealis Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 272.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 522-523, fig. 130, table 18
(redescription, geogr. range). —Milliron, 1973. Ent. Soc. Canada, Mem. 89: 117-120, pl. 12,
map 12 (redescription, geogr. records).

Biology: Plath, 1922. Psyche 29: 193-194 (life history). —Plath, 1927. Ent. Soc. Amer., Ann.
20: 189 (hibernation).

2204 Hymenoptera in America North of Mexico

Taxonomy: Plath, 1934. Bumblebees and their ways, p. 160 (life history).

Genus BOMBUS Subgenus FERVIDOBOMBUS Skorikov

Fervidobombus Skorikov, 1922. Sta. Region. Protect. Plantes, Petrograd Bul. 4: 153.
Type-species: Apis fervida Fabricius. Desig. by Frison, 1927.

This subgenus is found only in the New World and contains representatives in North and
South America.

Biology: Hobbs, 1966. Canad. Ent. 98: 33-39 (ecology of south. Alta. spp.).
californicus Smith. B. C. and Alta., south to Calif., Ariz. and N. Mex.; Mexieo (Baja California
and Sonora). Parasite: Psithyrus insularis (Sm.), P. suckleyi (Greene).
Bombus californicus Smith, 1854. Cat. Hym. Brit. Mus., v. 2. p. 400. @.
Bombus dubius Cresson, 1863. Ent. Soc. Phila., Proc. 2: 97. 3.
Bombus consanguineus Handlirsch, 1888. K. K. Naturhist. Hofmus., Ann. 3: 239. 2, 3, 8.
Bombus neglectulus Ashmead, 1902. Wash. Acad. Sci., Proc. 4: 124. 2, 8.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 32-39, figs.»2, 8, map 3
(redescription, variation in color patterns, geogr. records, as californicus californicus and
c. consanguineus). —Milliron, 1973. Ent. Soc. Canada, Mem. 89: 155-160, map 17 (tax.
status, synonymy, redescription, geogr. records, as fervidus californicus).

fervidus fervidus (Fabricius). Que. and N. B. south to Ga., west to B. C., Wash., Oreg. and

Calif.; Mexico (Chihuahua). Parasite: Brachicoma sarcophagina (Twns.), Melittobia
chalybii Ashm., Spherularia bombi Dufour. Predator: Philanthus bicinctus (Mickel).
Another subspecies, Bombus fervidus sonomae Howard, occurs in Mexico.

Apis fervida Fabricius, 1798. Sup. Ent. System., p. 274. %.

Apis alata Fabricius, 1798. Sup. Ent. System., p. 274.

Bombus elatus Fabricius, 1804. Systema Piezatorum, p. 352. Emend.

Bombus fervidus var. dorsalis Cresson, 1879. Amer. Ent. Soc., Trans. 7: 230. °.

Bombus nevadensis race aztecus Cockerell, 1899. Ann. and Mag. Nat. Hist. (7) 4: 389. 2.

Bombus fervidus var. umbraticollis Friese, 1931. Konowia 10: 301.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 27-32, figs. 2, 7, map 2
(redescription, variation in color patterns, geogr. records). —Mitchell, 1962. N. C. Agr.
Expt. Sta. Tech. Bul. 152: 523-524, fig. 130, table 18 (redescription, geogr. range, floral
records). —Medler, 1965. XIIth Internat]. Congr. Ent., Proc. 1964: 388-389 (size variation
in worker caste). —Milliron, 1973 Ent. Soc. Canada, Mem. 89: 147-155, pl. XIII, map 16
(synonymy, tax. status, redescription, geogr. and floral records).

Biology: Fairchild and Barrett, 1906. Ent. Soc. Wash., Proc. 8: 13-14, 1 pl. (copulation).
—Plath, 1922. Psyche 29: 180-187 (defense). —Plath, 1922. Psyche 29: 198-200 (life history).
—Plath, 1927. Ent. Soc. Amer., Ann. 20: 191 (hibernation). —Bequaert, 1932. Brooklyn Ent.
Soc., Bul. 27: 151 (arboreal nest). —Plath, 1934. Bumblebees and their ways, p. 166 (life
history, parasite). —Townsend, 1936. Ent. Soc. Wash., Proc. 38: 92-98 (parasite).
—Armitage, 1965. Kans. Ent. Soc., Jour. 38: 97 (predator). —Milliron, 1967. Canad. Ent. 99:
1321-1332 (artificially induced hibernation).

pennsylvanicus pennsylvanicus (Degeer). Que. and Ont., south to Fla., west to Minn., S. Dak.,

Nebr., Colo. and N. Mex.; Mexico and possibly Central America. Parasite: Brachicoma
sarcophagina (Twns.), Physocephala sagittaria (Say), Psithyrus insularis (Sm.), P.
variabilis (Cress.), Spherularia bombi Dufour. Predator: Mallophora bomboides (Wied.).

Apis pensylvanica Degeer, 1773. Mem. Serv. Hist. des Ins., v. 3, p. 575. 9.

Apis americanorum Fabricius, 1775. Systema Ent., p. 380.

Apis antiguensis Fabricius, 1775. Systema Ent., p. 380. °.

?Apis nidulans Fabricius, 1798. Sup. Ent. System., p. 274. 6.

Bombus pallidus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 92. 9.

Bombus pennsylvanicus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 94. Emend.

Psithyrus cevalliae Cockerell, 1899. Entomologist 32: 157. 3.

Bombus americana Howard, 1901. The insect book, pl. 1, figs. 30, 31. Lapsus calami.

Bombus titust Ashmead, 1902. Ent. News 13: 50. 6.

Bombus pennsylvanicus var. umbratus Friese, 1931. Konowia 10: 301.

Superfamily APOIDEA 2205

Taxonomy: Michener, 1953. Kans. Univ. Sci. Bul. 35: 1087, figs. 248-253 (larva, as
americanorum). —Moure, 1960. Studia Ent. 3: 151-152 (synonymy, notes on type of
antiguensis). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 524-526, figs. 130, 131,
table 18 (synonymy, redescription, geogr. range, floral records). —Milliron, 1973. Ent. Soe.
Canada, Mem. 89: 190-196, pl. XIII, map 25 (synonymy, tax. status, redescription, geogr.
and floral records).

Biology: Frison, 1921. Canad. Ent. 53: 100 (parasite). —Rau, 1922. Acad. Sci. St. Louis, Trans.
24: 40 (life history). —Rau, 1924. Ent. Soe. Amer., Ann. 17: 368-381 (hibernation, as
americanorum). —Plath, 1927. Ent. Soe. Amer., Ann. 20: 190 (hibernation, as
americanorum). —Frison, 1930. Ent. Soc. Amer., Ann. 23: 644-665 (life history, as
americanorum). —Rau, 1934. Acad. Sci. St. Louis, Trans. 28: 223 (life history, as
americanorum). —Plath, 1934. Bumblebees and their ways, p. 163 (life history, as
americanorum). —Rau, 1937. Brooklyn Ent. Soc., Bul. 32: 61 (nest-founding, as
americanorum). —Brower, Brower and Westcott, 1960. Amer. Nat. 94: 343-355 (mimicry
experimentation, as americanorwm). —Milliron, 1967. Canad. Ent. 99: 1330-1331
(hibernation).

pennsylvanicus sonorus Say. Tex., west to Calif.; Mexico. Parasite: Physocephala burgessi
(Will.), P. terana (Will.).
Bombus sonorus Say, 1837. Boston Jour. Nat. Hist. 1: 413. 9.
Bombus sonorus flavodorsalis Franklin, 1913. Amer. Ent. Soc., Trans. 38: 409. 9, 9.

Taxonomy: Stephen, 1957. Oreg. Agr. Expt. Sta. Tech. Bul. 40: 40-43, fig. 9, map. 4
(synonymy, redescription, geogr. records, as sonorus). —Milliron, 1973. Ent. Soc. Canada,
Mem. 89: 196-201, map 26 (tax. status, redescription, geogr. and floral records).

Biology: Ryckman, 1953. Pan-Pacifie Ent. 29: 144 (parasite). —Hurd and Linsley, 1975.
Smithsn. Contrib. Zool. 193: 48 (floral relationships).

UNPLACED TAXON IN BOMBUS

Bombus praticola Kirby, 1837. Fauna Bor.-Amer., v. 4, p. 274. This may be the correct name
for Bombus pleuralis Nylander.

Genus PSITHYRUS Lepeletier

Psithyrus Lepeletier, 1832. Soc. Ent. France, Ann. 1: 372.
Type-species: Apis rupestris Fabricius. Desig. by Curtis, 1833.
Apathus Newman, 1835. Ent. Mag. 2: 404, footnote.
Type-species: Apis rupestris Fabricius. Autobasic with Psithyrus; name proposed
to replace Psithyrus, thought to be preoccupied by Psithyros Hubner, 1819.
Psithyrus subg. Ashtonipsithyrus Frison, 1927. Amer. Ent. Soc., Trans. 53: 69.
Type-species: Apathus ashtoni Cresson. Orig. desig.
Psithyrus subg. Fernaldaepsithyrus Frison, 1927. Amer. Ent. Soc., Trans. 53: 70.
Type-species: Psithyrus fernaldae Franklin. Orig. desig.
Psithyrus subg. Eopsithyrus Popov, 1931. Eos 7: 134.
Type-species: Psithyrus tibetanus (Morawitz). Orig. desig.
Psithyrus subg. Metapsithyrus Popov, 1931. Eos 7: 135.
Type-species: Psithyrus campestris (Panzer). Orig. desig.
Psithyrus subg. Allopsithyrus Popov, 1931. Eos 7: 136.
Type-species: Psithyrus barbatellus (Kirby). Orig. desig.
Psithyrus subg. Ceratopsithyrus Pittioni, 1949. Eos 25: 271.
Type-species: Psithyrus klapperichi Pittioni. Orig. desig. and Monotypice.
The genus Psithyrus contains a few species of obligate social parasites on the members of the
genus Bombus. They closely resemble the species of Bombus, but lack the worker caste, and the
queens lack pollen-collecting baskets on the hind legs.

Taxonomy: Frison, 1919. Ill. State Acad. Sci., Trans. 12: 157-165 (Ill. spp.). —Frison, 1923.
Amer. Ent. Soc., Trans. 48: 307-326 (synonymy, tax. characters, geogr. records). —Frison,
1926. Amer. Ent. Soc., Trans. 52: 129-145 (synonymy, tax. characters, geogr. records).
—Frison, 1927. Amer. Ent. Soc., Trans. 53: 51-78, pls. XVI-XVII (systematic relationships

2206 Hymenoptera in America North of Mexico

of spp. in America north of Mexico). —Milliron, 1961. Kans. Ent. Soc., Jour. 34: 59-60
(subgeneric synonymy). —Medler, 1962. Canad. Ent. 94: 444-447, 1 fig., 3 tables
(morphometric study). —Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 538-544, fig.
134, table 18 (east. U.S. spp.).

Biology: Sladen, 1899. Ent. Monthly Mag. 35: 230-234 (habits, host). —Sladen, 1915. Canad.
Ent. 47: 84 (habits, host). —Frison, 1921. Canad. Ent. 53: 100-101 (hosts). Plath, 1922. Biol.
Bul. 43: 23-44, pl. I (life history, hosts). Frison, 1926. Ent. Soc. Amer., Ann. 19: 205-221, pl.
XVII, fig. 3b (life history, hosts).

—Plath, 1927. Brooklyn Ent. Soc., 22: 121-125 (life history, enemy of Apis mellifera).
—Plath, 1934. Bumblebees and their ways, 201 pp., New York, Macmillan (life history,
hosts).
ashtoni (Cresson). P. E. I. west to Sask., south to N. Dak., Minn., Wis., Mich., Ohio, W. Va. and
Va. Host: Bombus affinis Sm., B. terricola Kby.
Apathus ashtoni Cresson, 1864. Ent. Soc. Phila., Proc. 3: 42. °.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 539-540, table 18
(redescription).

Biology: Plath, 1922. Psyche, 29: 191 (hibernation, hosts). —Plath, 1934. Bumblebees and
their ways, p. 169 (habits, hosts).
citrinus (Smith). P. E. I. and N. B., south to Fla. and Ala., west to S. Dak. and N. Dak. Host:

Bombus impatiens (Cr.), B. nevadensis awricomus (Robt.), B. vagans (Sm.); also
occasionally attempts to invade hives of Apis mellifera Linn. For years Psithyrus
laboriosus (Fabricius) was considered to be the correct name for this species; however,
an examination of the type specimen (see Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul.
152: 333) has demonstrated that laboriosus is a member of the genus Emphoropsis in
the Anthophoridae.

Apathus citrinus Smith, 1854. Cat. Hym. Brit. Mus., v. 2, p. 385. ¢.

Apathus contiguus Cresson, 1863. Ent. Soc. Phila., Proc. 2: 112. d.

Taxonomy: Mitchell, 1962. N. C. Agr. Expt. Sta. Tech. Bul. 152: 540-541, table 18
(redescription).

Biology: Frison, 1921. Canad. Ent. 53: 100-101 (habits, hosts, as laboriosus). —Bequaert and
Plath, 1925. Mus. Compar. Zool., Bul. 67: 268-272 (life history, host, as laboriosus).
—Plath, 1927. Ent. Soc. Amer., Ann. 20: 191 (hibernation, as laboriosus). —Plath, 1927.
Brooklyn Ent. Soc., Bul. 29: 121-125 (inquiline in hives of Apis mellifera Linn.). —Plath,
1934. Bumblebees and their ways, p. 175 (hosts, habits).

fernaldae Franklin. Alaska and Canada, south to N. C. and Tenn. in east. U. S. and Colo. and
Calif. in west. U. S. Host: Bombus appositus Cress., B. perplerus Cress.?, B. rufocinctus
Cress., B. terricola occidentalis Greene.
Psithyrus fernaldae Franklin, 1911. Amer. Ent. Soc., Trans. 37: 164. ?.
Psithyrus tricolor Franklin, 1911. Amer. Ent. Soc., Trans. 37: 167. ¢.
Psithyrus wheeleri Bequaert and Plath, 1925. Mus. Compar. Zool., Bul. 67: 265. 2, d.

Taxonomy: Frison, 1923. Amer. Ent. Soc., Trans. 48: 321 (synonymy). —Frison, 1926. Amer.
Ent. Soc., Trans. 52: 145 (tax. status of wheeleri). —Mitchell, 1962. N. C. Agr. Expt. Sta.
Tech. Bul. 152: 541-542, fig. 134, table 18 (redescription).

Biology: Plath, 1927. Ent. Soc. Amer., Ann. 20: 191 (hibernation). —Plath, 1934. Bumblebees
and their ways, p. 171 (habits, hosts).

Morphology: Milliron, 1961. Brooklyn Ent. Soc., Bul. 55: 109-113, 2 figs. (gynandromorph).
insularis (Smith). Canada, south to Calif., Ariz. (Oak Creek Canyon), N. Mex., Nebr. (Sioux
County), N. Y. (Ithaca) and N. H. (Durham), ?Alaska (Berg Bay). Host: Bombus
appositus Cress., B. bifarius nearcticus Handlirsch, B. californicus Sm., B. flavifrons
Cress., B. nevadensis nevadensis Cress., B. pennsylvanicus (DeG.), B. rufocinctus
Cress., B. ternarius Say, B. terricola occidentalis Greene, B. t. terricola Kby.
Bombus interruptus Greene, 1858. Lye. Nat. Hist. N. Y., Ann. 7: 11. 2. Preoce.
Apathus insularis Smith, 1861. Jour. of Ent. 1: 155. @.

Superfamily APOIDEA 2207

Psithyrus consultus Franklin, 1913. Amer. Ent. Soc., Trans. 38: 459. 3.
Psithyrus crawfordi Franklin, 1913. Amer. Ent. Soc., Trans. 38: 464. 2, ¢.

Taxonomy: Frison, 1923. Amer. Ent. Soc., Trans. 48: 321-322 (synonymy). —Mitchell, 1962. N.
C. Agr. Expt. Sta. Tech. Bul. 152: 542-543, table 18 (redescription).

Biology: Sladen, 1915. Canad. Ent. 47: 84 (host). —Plath, 1927. Ent. Soc. Amer., Ann. 20: 191
(hibernation, host). —Plath, 1934. Bumblebees and their ways, p. 173 (life history, hosts).
—Craig, 1953. Canad. Ent. 85: 311-312 (host).

suckleyi (Greene). Alaska, south to Calif., Utah and Colo. Host: Bombus appositus Cress., B.
californicus Sm., B. nevadensis nevadensis Cress., B. rufocinctus Cress., B. terricola
occidentalis Greene, B. t. terricola Kby.

Bombus suckleyi Greene, 1860. Lye. Nat. Hist. N. Y., Ann. 7: 169. ¢.
Psithyrus latitarsus Morrill, 1903. Canad. Ent. 35: 224. 9.

Taxonomy: Frison. 1926. Amer. Ent. Soc., Trans. 52: 144 (synonymy).
variabilis (Cresson). Ohio south to Fla., west to N. Dak., S. Dak., Nebr. Kan., Okla., Tex. and
N. Mex.; Mexico (Orizaba). Host: Bombus pennsylvanicus (Deg.).
Apathus variabilis Cresson, 1872. Amer. Ent. Soc., Trans. 4: 284. 2, d.
Psithyrus bicolor Franklin, 1913. Amer. Ent. Soc., Trans. 38: 460. 9.

Taxonomy: Frison, 1923. Amer. Ent. Soc., Trans. 48: 322 (synonymy). —Mitchell, 1962. N. C.
Agr. Expt. Sta. Tech. Bul. 152: 543-544, fig. 134, table 18 (redescription).

Biology: Frison, 1916. Brooklyn Ent. Soc., Bul. 11: 46-47 (habits, host). —Frison, 1921. Canad.
Ent. 53: 100 (host). —Webb, 1961. North Central Branch, Ent. Soc. Amer., Proc. 16: 16
(habits).

SUBFAMILY APINAE

The subfamily contains two tribes, the Meliponini and the Apini of which only the latter is
represented in America north of Mexico by the introduced European honeybee. The members of
this subfamily are all highly eusocial and live in perennial colonies.

TRIBE APINI

The tribe includes only the genus Apis which prior to the activities of man was indigenous to
Africa, Eurasia, Japan, Formosa, the Philippines and Indonesia. One of its species, the common
honeybee (Apis mellifera Linnaeus) has been successfully introduced into the Western Hemi-
sphere and many areas of the Old World beyond its original range (Africa, Europe, and western
Asia).

Taxonomy: Maa, 1953. Treubia 21: 525-640 (classification). —Ruttner, 1968. Jn Chauvin,

Traite de biologie de l’abeille, Paris, Masson et cie 1: 27-44 (review of included spp. and
races).

Biology: Michener, 1974. The Social Behavior of the Bees, chapter 30: 347-366 (natural
history).

Morphology: Cruz-Landim, 1974. N. Y. Ent. Soc., Jour. 71: 2-31 (evolution of wax and scent
glands).

Genus APIS Linnaeus

Apis Linnaeus, 1758. Syst. Nat., Ed. 10, p. 343.
Type-species: Apis mellifera Linnaeus. Desig. by Latreille, 1810. (=A pis mellifica
Linnaeus).
Apicula Rafinesque, 1814. Principles fondamentaux de somiologie, Palermo, p. 29. N. name,
proposed unnecessarily to replace Apis Linnaeus.
Apiarus Rafinesque, 1815. Analyse Nature ou Tabl. Univers, Palermo, p. 123. N. name,
proposed unnecessarily to replace Apis Linnaeus.
Megapis Ashmead, 1904. Ent. Soc. Wash., Proc. 6: 120.
Type-species: Apis dorsata Fabricius. Orig. desig.
Micrapis Ashmead, 1904. Ent. Soc. Wash., Proc. 6: 122.

2208 Hymenoptera in America North of Mexico

Type-species: Apis florea Fabricius. Monotypic and orig. desig.
Sigmatapis Maa, 1953. Treubia 21: 556.
Type-species: Apis cerana Fabricius. Orig. desig.

mellifera Linnaeus. Worldwide. Introduced in Western Hemisphere. Represented in the Old

World by numerous subspecies, several of which have been introduced into North
America with the result that much interbreeding has occurred among them. The
escaped bees of the woodlands are largely A. mellifera mellifera (German bee) while
the beekeepers’ bees are largely A. mellifera ligustica (Italian bee). A. mellifera
remipes (Caucasian bee) is also utilized by some beekeepers. For convenience these
forms are listed below in the synonymy.

Apis mellifera Linnaeus, 1758. Syst. Nat., Ed. 10, p. 576.

Apis mellifica Linnaeus, 1761. Fauna Suecica, Ed. 2, p. 421. N. name, proposed
unnecessarily to replace mellifera.

Apis mellifica var. Ligustica Spinola, 1806. Insectorum Liguriae, v. 1, p. 35.

Apis mellifica var. remipes Gerstaecker, 1862. Geog. Verbr. d. Honigbiene, p. 61.

Taxonomy: Alpatov, 1929. Quart. Rev. Biol. 4: 1-58 (biometric studies). —Michener, 1953.
Kans. Univ. Sci. Bul. 35: 1094, figs. 275-280 (larva). —Torchio and Torchio, 1975. Utah
State Agr. Expt. Sta., Research Rpt. 20: 1-36, 78 figs. (larvae).

Biology: Buttel-Reepen, 1903. Die stammesgeschichtliche Entstehung des Bienenstaates, 167
pp., Leipzig, G. Thieme. —Buttel-Reepen, 1915. Leben und Wesen der Bienen, 300 pp.,
Braunschweig, Vieweg und Sohn. —Philip and Vansell, 1932. Calif. Agr. Expt. Sta., Cire.
62: 1-27 (pollination of deciduous fruits). —Butler, 1949. The honeybee, 436 pp., Oxford,
Clarendon Press. —Percival, 1950. New Phytol. 49: 40-63 (pollen collection). —Frisch, 1950.
Bees, their vision, chemical senses and language, 119 pp., Ithaca, New York, Cornell Univ.
Press. —Wykes, 1952. Jour. Expt. Biol. 29: 511-519 (nectar preferences). —Frisch, 1953.
Aus dem Leben der Bienen, 180 pp., Revised ed., Berlin, Springer-Verlag. —Ribbands,
1953. The Behavior and Social Life of Honeybees, 352 pp., Bee Research Assn. Ltd.
(republished in 1964, by Dover Publ., Inc., New York). —Butler, 1954. The World of the
Honeybee, 226 pp., London, Collins. —Frisch, 1955. The Dancing Bees, 183 pp., New York,
Harcourt, Brace and Co. —Smith, 1960. Beekeeping in the tropics, 265 pp., London,
Longmans. —Lindauer, 1961. Communication among social bees, 143 pp., Cambridge,
Massachusetts, Harvard Univ. Press. —Singh, 1962. Beekeeping in India, 275 pp., New
Delhi, India Council of Agr. Research. —Grout, 1963. The hive and the honeybee, 652 pp.,
Revised, Hamilton, Illinois, Dadant and Sons. —Zander, 1964. Das Leben der bienen,
(Handbook der Bienenkunde, vol. 4), 6th edition, Stuttgart, K. Weiss. —Frisch, 1965.
Tanzsprache und Orientierung der Bienen, 566 pp., Berlin, Springer-Verlag. (republished
in 1967 in English by Belknap Press, Harvard University Press). —Ordetx and Espina,
1966. La apicultura en las tropicos, 412 pp., Mexico City, B. Trucco. —Frisch, 1967. The
dance language and orientation of bees, 566 pp., Cambridge, Massachusetts, Harvard Univ.
Press. —Chauvin, 1968. Traite de biologie de l’abeille, vols. I-V, Paris, Masson et Cie.

— Wenner, 1971. The bee language controversy, 109 pp., Educational Programs
Improvement Corporation, Boulder, Colo. —Michener, 1975. Ann. Rev. Ent. 20: 399-416
(Brazilian bee problem). —Edrich, 1977. Anim. Behaviour 25: 342-363 (interaction of light
and gravity in the orientation of the waggle dance).

Morphology: Stellwaag, 1910. Ztschr. Wiss. Zool. 95: 518-550, 6 figs., 2 pls. (structure and
mechanics of flight apparatus). —Frisch, 1915. Zool. Jahrb. abt. Zool. Physiol. 35: 1-182
(form and color senses). —Snodgrass, 1925. Anatomy and Physiology of the honeybee, 327
pp., New York, McGraw Hill Co. —Kuhn, 1927. Ztschr. Vergleich. Physiol. 5: 762-800 (color
vision). —Trojan, 1930. Ztschr. Morph. Okol. Tiere 19: 678-685 (Dufour’s gland). — Hertz,
1939. Jour. Expt. Biol. 16: 1-8 (color vision). —Snodgrass, 1942. Smithsn. Mise. Coll. 103:
1-120, 32 figs. (skeleto-muscular mechanisms). —Snodgrass, 1956. Anatomy of the
honeybee, 334 pp., Ithaca, New York, Cornell Univ. Press. —Simpson, 1960. Jour. Ins.
Physiol. 4: 107-121 (function of salivary glands). —Barbier and Lederer, 1960. Acad. Sci.
Paris, Compt. Rend. 250: 4467-4469 (chemical structure of royal jelly). —Barbier, Lederer
and Nomura, 1960. Acad. Sci. Paris, Compt. Rend. 251: 1133-1135 (synthesis of substances
in royal jelly). —Barbier and Pain, 1960. Acad. Sci. Paris, Compt. Rend. 250: 3740

Superfamily APOIDEA = 2209

(mandibular gland secretions). —Daly, 1964. Calif. Univ. Pubs. Ent. 39: 1-77, 54 figs.
(skeleto-muscular morphogenesis of thorax and wings). — Youseff, 1971. Smithsn. Contrib.

Zool. 99: 1-54, 10 figs., 5 tables (topography of cephalic musculature and nervous system).
—Cruz-Landim, 1974. Brasil. Biol., Rev. 34: 105-118, 5 figs. (cardiac musculature).
—Cruz-Landim, 1975. Ciencia e Cultura 27: 278, 3 figs., 2 tables (extracellular crystals).

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