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PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of
WASHINGTON

Volume 84

OFFICERS FOR THE YEAR 1982

President Margaret S. Collins
President-elect Manya B. Stoetzel
Recording Secretary Thomas E. Wallenmaier
Corresponding Secretary Ralph E. Harbach
Treasurer F. Christian Thompson
Editor David R. Smith
Custodian John F. Carroll
Program Committee Chairman Jay C. Shaffer
Membership Committee Chairman Joyce A. Utmar
Delegate to the Washington Academy of Sciences Margaret S. Collins
Hospitality Chairman Helen Sollers-Riedel

Published by the Society
WASHINGTON, D.C.
1982

TABLE OF CONTENTS, VOLUME 84
ADAMSKI, D. and T. M. PETERS—Axillary structure of the Tortricidae (Lepidop-

ADAMSKI, D.—See PETERS, T. M.

AGAFITEI, N. J—See SELANDER, R. B.

ALM, S. R. and F. E. KURCZEWSKI—Antennal sensilla and setae of Anoplius
fenebrosus. (Cresson) (Etymenoptera: Pompilidae) 22s... 0. ses ise oe ee eee ens

BLANCHARD, A. and J. G. FRANCLEMONT—A new species of Zale Hiibner from
Texas and New Mexico (Lepidoptera: Noctuidae: Catocalinae) ..................5.

BLANCHARD, A. and J. G. FRANCLEMONT—Marilopteryx carancahua, a new
genus and new species from east Texas (Lepidoptera: Noctuidae: Hadeninae) ......

397

ii PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

BLANCHARD, A. and E. C. KNUDSON—A new species of Symmetrischema
Povolny (Lepidoptera: Gelechiidae) from Texas ............. ee eeeee esse ence eee

BROWN, R. L.—Notes on Gretchena: A new species and the synonymy of Gwendolina
(Lepidoptera: Tortricidae): - s0,.2 s +1 a:chg gees ottes ios coke aS cei s wojme oe ++ oracarelenrteiepee

BURNS, J. M.—Lychnuchoides frappenda from central Mexico joins lunus and zweifeli
in a /unus group of Atrytonopsis (Lepidoptera: Hesperiidae: Hesperiinae) ..........

CALABRESE, D. M. and P. TALLERICO—Chromosome study in males of Nearctic
species of Gerris Fabricius and Limnoporus Stal (Hemiptera: Heteroptera: Ger-
litt (3) Coton API RCE PRO OC REO ae DONO oo nila i Don erinted OaTeD MEME oa eb! n coe

CALABRESE, D. M.—See DUARTE, J.

CAPRILES, J. M.—See HENRY, T. J.

CWIKLA, P. S. and P. H. FREYTAG—Three new leafhoppers (Homoptera: Cica-
dellidac)ifrom Cocos Island! iss = x. :<cxsustevsveacusucncushenevedsptey ler ote ete tene) ayeise) scetslions arever oRaeaens

DARLING, D. C. and N. F. JOHNSON—Egg mortality in the eastern tent caterpillar,
Malacosoma americanum (Lepidoptera: Lasiocampidae): The role of accessory
glandssecretions:and egg mass Shape? <n a tercverh Pee ai ehe ele ete cle ols oe o.0 ons nce Sle cme ners

DELONG, D. M.—New Central and South American leafhoppers of the *‘Bahita”’
group (Homoptera: Cicadellidae: Deltocephalinae) >..-......-...-..--+-+s+e2-e-ee

DELONG, D. M.—New species of Xestocephalinae (Homoptera: Cicadellidae) from
Mexicosbanamas Peru. and: Brazil) e227 accl-emccosecnn soe cs aie on. oe eee

DELONG, D. M.—Some new Neotropical leafhoppers of the subfamilies Iassinae
ands Deltocephalinae (Homoptera: Gicadellidae)io>.- 4. -4250050. 222 200s) eee

DUARTE, J. and D. M. CALABRESE—Is the binomen Lygaeus kalmii Stal (Hemip-
tera: Heteroptera: Lygaeidae) applied to sibling species? ......................6:-

EADS, R. B., G. C. SMITH, and G. O. MAUPIN—A CO, platform trap for taking
adult DermacentoriandersonucAcati-slxodidae)) asses aero eee oe oon eee ne

FAIRCHILD, G. B.—See WILKERSON, R. C.

FAIRCHILD, G. B.—See WILKERSON, R. C.

FARRIER, M. H.—See SHERRON, D. A.

FISHER, E. M. and H. A. HESPENHEIDE—Taxonomy and ethology of a new
Central American species of the robber fly genus Glaphyropyga (Diptera: Asilidae)

FISHER, T. W.—See ORTH, R. E.

FOOTE, B. A.—Biology and immature stages of Setacera atrovirens, a grazer of
floating. algal mats (Diptera: Ephydridae) fo: ais dacs b eS va cos: wae el eeeeenee

FRANCLEMONT, J. G—See BLANCHARD, A.

FRANCLEMONT, J. G.—See BLANCHARD, A.

FREYTAG, P. H.—See CWIKLA, P. S.

GIBSON, L. P.—New species of Urosigalphus (Hymenoptera: Braconidae) from
ONC a cai said ns igpnys S's audho' Ade Seo Agden Sa aie sd IT a ce

GIBSON, L. P.—New species of Urosigalphus (Hymenoptera: Braconidae) from
SOMONE IA ASNT GEA faa ice cx piece a, be'S 0! ws, ig aeeneie gS acer tae dear BAe Ce ee

GILES, F. E. and W. W. WIRTH—Three new species of Serromyia (Diptera:
Ceratopogonidae) from Sri Lanka, with new records and a key to the Oriental and
AUST AN AN SPECIES: lich. igs vic. c' be 0' «cha k ey eee eae Acimeke ee ae ee

GILES, F. E. and W. W. WIRTH—New species and new collection records of
Ceratoporonidae (Diptera) from) Sri Gama gc oo5 a6 Ysein civis Behe visita oo we Ae ee ee

GLICK, J. I. and G. R. MULLEN—A new species of biting midge of the genus
Alluaudomyia Kieffer (Diptera: Ceratopogonidae) from the southeastern United
BEAR os ARE clot iin By Sony ep sie’ ch ale v. 6 eid gg ROE este aI ENG, See ae

GORDH, G. and B. A. HAWKINS—Tetrastichus cecidobroter (Hymenoptera: Euloph-

535

632

448

184

3)

610

301

342

716

828

97

167

440

822

539

VOLUME 84, NUMBER 4

idae), a new phytophagous species developing within the galls of Asphondylia (Dip-
tera: Cecidomyiidae) on Atriplex (Chenopodiaceae) in southern California ..........

GORDON, R. D.—An old world species of Scymnus (Pullus) established in Pennsyl-
vania and New York (Coleoptera: Coccinelidae) .................22e--02e eee eeeee

GORDON, R. D.—Two new species of Nephaspis Casey (Coleoptera: Coccinellidae)
frommlnntaadrand: Colombia aero ee ere rte inte ote eters ics 2 oelereeeiGheetsl olay

GRISSELL, E. E.—See SCHAUFF, M. E.

HARRIS, S. C., P. K. LAGO, and R. W. HOLZENTHAL—An annotated checklist
of the caddisflies (Trichoptera) of Mississippi and southeastern Louisiana. Part II:
Rhy acophvloidear ray eyes ii. G12 aSi.re Aarcintgie ore mater teks fae mee ace atalee olste s wihan = aaaalalsin| aya

HARRIS, S. C.—See HOLZENTHAL, R. W.

HARRIS, S. €.—See LAGO, P. K.

HAWKINS, B. A.—See GORDH, G.

HEITZMAN, R. L.—Descriptions of the mature larva and pupa of Hypomecis umbro-
Sarian (Elubnen)) (eepidopteras Geometridae) sec ete ie etait eines eee rele tet lh

HENRY, C. S.—Reproductive and calling behavior in two closely related, sympatric
lacewing species, Chrysopa oculata and C. chi (Neuroptera: Chrysopidae) .........

HENRY, T. J.—The onion plant bug genus Labopidicola (Hemiptera: Miridae):
Economic implications, taxonomic review, and description of a new species ........

HENRY, T. J.—New synonymies and a new combination in the North American Miridae
MHentiptera) crac ec ews sole es ose ved scl cape me leis seiner tel gona Sita olerden ety erly

HENRY, T. J. and J. M. CAPRILES—The four “‘ocelli’’ of the isometopine genus
Isometocoris Carvalho and Sailer (Hemiptera: Miridae) ................--.2+-eeee-

HEPPNER, J. B.—Synopsis of the Glyphipterigidae (Lepidoptera: Copromorphoidea)
DEANE SOULE 4 4 oreo, Mal eicaciok Rio EIB SCR canoe eains higue a aninodo wontons 7am te

HEPPNER, J. B.—Synopsis of the Hilarographini (Lepidoptera: Tortricidae) of the
WHO otis aie be ee ns Bec ae a nice alo aie cil Acer theca aCIMOI bio OO GD. c.0 DOwS OBS

HESPENHEIDE, H. A.—See FISHER, E. M.

HOEBEKE, E. R. and A. G. WHEELER, JR.—Rhopalus (Brachycarenus) tigrinus,
recently established in North America, with a key to the genera and species of
Rhopalidae in eastern North America (Hemiptera: Heteroptera) ..........-.-....+.

HOEBEKE, E. R.—See WHEELER, A. G., JR.

HOLZENTHAL, R. W., S. C. HARRIS, and P. K. LAGO—An annotated checklist
of the caddisflies (Trichoptera) of Mississippi and southeastern Louisiana. Part III:
Himmnep hiloid earanGacOnGlsiOmSirc- 1 ite -toiey= etre ones ole epee teen dreac die e= eh) ote tehe tote det tel 2

HOLZENTHAL, R. W.—See HARRIS, S. C.

HOLZENTHAL, R. W.—See LAGO, P. K.

HUDSON, A.—Evidence for reproductive isolation between Xestia adela Franclemont
and Xestia dolosa Franclemont (Lepidoptera: Noctuidae) ................0ee eee eue

HUNG, A. C. F.—Chromosome and isozyme studies in Trichogramma (Hymenoptera:
MTICHOPrAMMAtIGAE): sce. So ees RUSS SES IS Pe RAIS, HIE MEN, | Sows

JOHNSON, M. D.—See LAGER, T. M.

JOHNSON, N. F.—See DARLING, D. C.

KELLEY, R. W. and J. C. MORSE—A key to the females of the genus Oxyethira
(Trichoptera: Hydroptilidae) from the southern United States ...............-..05.

KINGSOLVER, J. M.—Taxonomic studies in the genus Rhipibruchus Bridwell (Coleop-
tera: Bruchidae), with descriptions of four new species ................50.s0ceeeee

KINGSOLVER, J. M.—Conicobruchus albopubens (Pic) (Coleoptera: Bruchidae) and
its host Cyamopsis tetragonoloba (L.) (Leguminosae), with the designation of a
ISRO TOA nid ao aie nie aang DOO oor © Suh SE SOs GA Cer RO Cig aon Sen Ie OL iog Sok aae

ill

509

213

513

iV PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

KIRCHNER, R. F.—A new Allocapnia from West Virginia (Plecoptera: Capniidae) ...

KIRCHNER, R. F.—See KONDRATIEFF, B. C.

KITTLE, P. D.—Two new species of water striders of the genus Trepobates Uhler
(elemiptera: Gerridae) soc. Mes ce ater emt letee aeie ae tekst: baci aalels alee tee netate

KNUDSON, E. C.—See BLANCHARD, A.

KONDRATIEFF, B. C. and R. F. KIRCHNER—Notes on the winter stonefly genus
Allocapnia (Plecoptera: Capniidae) cree «cite ts ot cia tte eye) See ttt ee tian

KONDRATIEFF, B. C. and J. R. VOSHELL, JR.—The Perlodinae of Virginia, USA
(Plecoptera: Perlodidac) iter Rie Rr Nite el Sis slncior etal chaise +, = aoe

KORMILEV, N. A.—Six new species of Neotropical Aradidae (Hemiptera) ..........

KURCZEWSKI, F. E.—An additional study on the nesting behaviors of species
ob Miscophus (Hymenoptera? Sphecidae))-,. .. occ... asec icistel le ne sie ce eer eita anette

KURCZEWSKI, F. E. and D. J. PECKHAM—Nesting behavior of Lyroda subita
(Say): (Ely menopterassSphecidae)itz% 2.002 eaSo: ein veal eee esi. tn ele eee

KURCZEWSKI, F. E.—See ALM, S. R.

KURCZEWSKI, F. E.—See O'BRIEN, M. F.

LAGER, T. M., M. D. JOHNSON, and W. P. MCCAFFERTY—tThe mayflies of north-
eastern Minnesota (Ephemeroptera) ic... ace stone aie soe nse Sica Biche Sid s vlote cle ee

LAGO, P. K., R. W. HOLZENTHAL, and S. C. HARRIS—An annotated checklist
of the caddisflies (Trichoptera) of Mississippi and southeastern Louisiana. Part I:
Introduction and Hydropsychoidea = 6 Cr ciss cetera cette elotelote oto iets <forec «ore

LAGO, P. K.—See HARRIS, S. C.

LAGO, P. K.—See HOLZENTHAL, R. W.

LAMBDIN, P. L.—See MILLER, G. L.

LAVIGNE, R. J.—Ethology of Neoitamus vittipes (Diptera: Asilidae) in South Aus-
Lea FEN Sores SC eRe bee nor ear ta, Oa a ee RE CIO RECT. Sc

LAVIGNE, R. J.—Notes on the ethology of Neoscleropogon elongatus (Diptera:
Asilidae) in), SouthtAus traliates.crsccyaeedte orate ee ae eee isis ices oe eee

McCAFFERTY, W. P.—See LAGER, T. M.

MAIER, C. T.—Abundance and distribution of the seventeen-year periodical cicada,
Magicicada septendecim (Linnaeus) Hemiptera: Cicadidae—brood II), in Connec-
TLCELE Ha, cee ERR e ES EO a as eee tr a eo MeO PE rd tio cer hae 2 eee ee

MAIER, C. T.—Larval habitats and mate-seeking sites of flower flies (Diptera: Syrph-
AGS ME RISTATINAL)) «0.2555. cc oie eve Reale An choles Rote NO ee Ae bd sav oo SRE ee ee ore

MARSH, P. M.—Two new species of Heterospilus (Hymenoptera: Braconidae) from
Mexico being introduced against the cotton boll weevil, Anthonomus grandis (Cole-
optera?Cureulionidae) (ais 04. Rasen. RIN oe Bsa aoa Se ae oe ee

MARSH, P. M.—See STARY, P.

MATHIS, W. N.—Description of a new species of Nocticanace Malloch (Diptera:
Canacidae) from Sri Lanka with notes on two related species ..................05-

MATTA, J. F.—The bionomics of two species of Hydrochara (Coleoptera: Hydrophil-
idae) with descriptions of their larvae

MAUPIN, G. O.—See EADS, R. B.

MAYOR, A. J.—A review of the genus Chaetocoelus LeConte (Coleoptera: Mala-
CUNGAE WN +s leek Wee Aaa Uo ase RRS HS Shae end cpa te A eee

MORSE, J. C.—See KELLEY, R. W.

MULLEN, G. R.—See GLICK, J. I.

MILLER, G. L. and P. L. LAMBDIN—Hemerobius stigma Stephens (Neuroptera:
Hemerobiidae): External morphology of the ge... .. ss. 20150. occauectses Seo

786

157

240

761
480

67

149

729

495

617

742

430

603

849

42]

461

475

VOLUME 84, NUMBER 4

NORTON, R. A.—Arborichthonius n. gen., an unusual enarthronote soil mite (Acarina:
Oribaten) roms Ontarowe nici eee eae eee Peel 5 Sef B i SP we
O'BRIEN, M. F. and F. E. KURCZEWSKI—Further observations on the ethology
of Alysson conicus Provancher (Hymenoptera: Sphecidae) ........................
ORTH, R. E.—Five new species of Pherbellia Robineau-Desvoidy, subgenus Oxytaenia
Sacksirom North) America (Diptera:iSciomyzidae)i sa. cee: eee eos vieeinle> avela eile a
ORTH, R. E. and T. W. FISHER—A new species of Tetanocera Dumeéril from Colo-
ragor(Dipteray SClOMy ZiGde)) i hi At: SE, Ale ae Ieee ie eI Sie alo eres
PECKHAM, D. J—See KURCZEWSKI, F. E.
PETERS, T. M. and D. ADAMSKI—A description of the larva of Dixella nova (Walker)
(Diptera eDixidae) i vecusctmd sidesenaea «Kona ton ete Gees Gomismtin cit eden sk SAMS
PETERS, T. M.—See ADAMSKI, D.
PINTO, J. D.—The phenology of the plant bugs (Hemiptera: Miridae) associated with
Ceanothus crassifolius in a chaparral community of southern California ............
PINTO, J. D—New synonymies, lectotype designations, and other notes on North
AmMencanvEprcaural (Coleoptera; Meloidae)i hemes weenie seer ee nee eee oc
REICHART, C. V.—An addition to the genus Anisops of Australia (Hemiptera:
INOTOMECIIGAE)! HAO DSES oe cen ns See acre cet TERMI ERRYS domes Mes ees hod cite nck
ROLSTON, L. H.—A revision of Euschistus Dallas subgenus Lycipta Stal (Hemiptera:
Pentatomidae) eitort..caceee et otter ae Scrais sie eet ci stench rere cae iar oree sree exe
ROSS, M. H.—See SHERRON, D. A.
ROTH, L. M.—Ovoviviparity in the blattellid cockroach, Symploce bimaculata (Ger-
Staccken) (Dictyoptera: Blattariay Blattellidac) mcm ateeee eeetie eee meee ce ae see ee
SCARBROUGH, A. G.—Coexistence in two species of Holcocephala (Diptera: Asilidae)
invas\ianvland habitat bredatonysbehavion. ase Jes aaetene eee coe ee erie:
SCHAUFF, M. E. and E. E. GRISSELL—Nomenclatural notes on Polynema (Hyme-
noptera: Mymaridae), with description of a new species ..............0ceeeeee cece
SELANDER, R. B.—Larval development in blister beetles of the genus Linsleya
(Coleoptera Meloidae) rt. Bek See AS Pe PFs ean erated sroleere shades coma teeta ths
SELANDER, R. B.—Sexual behavior, bionomics, and first-instar larvae of the Lauta
and Diversicornis groups of Epicauta (Coleoptera: Meloidae) .....................
SELANDER, R. B. and N. J. AGAFITEI—First-instar larvae of the Uniforma Group
eithe genus ‘Epicauta (Coleoptera: Meloidae): ........2.<....<,..« esate epalnees ase ek
SHERRON, D. A., C. G. WRIGHT, M. H. ROSS, and M. H. FARRIER—Density,
fecundity, homogeneity, and embryonic development of German cockroach (Blattella
germanica (L.)) populations in kitchens of varying degrees of sanitation (Dictyoptera:
ES ALTE MT AG) |x, oy See uss ht es Set robe tay hoot etrec Mee eats ea es Ses ee See Tae
SMITH, C. F.—A key to the species of Hyalomyzus (Homoptera: Aphididae) in North
pamerica, with the: description of a New Species )x.22k.8s sae ween een se ele Ante
SMITH, D. R.—Smyphyta (Hymenoptera) of Sri Lanka ................0cce cece eee
SMITH, G. C.—See EADS, R. B.
STARY, P. and P. M. MARSH—A new species of Trioxys (Hymenoptera: Aphidiidae)
NTC UON Cd PCCAIMADING o octiea casts +.«5 descdne eo eet Oe Cee ER eae IS oe te ere
STARK, B. P. and K. W. STEWART—Oconoperla, a new genus of North American
Rerlodinae; (ElecopteraveenOdidae)i...e shoes ct told aceite cee cient ene
STEINER, W. E., JR.—Poecilocrypticus formicophilus Gebien, a South American
beetle established in the United States (Coleoptera: Tenebrionidae) ................
STEWART, K. W.—See STARK, B. P.
STEYSKAL, G. C.—A second species of Ceratitis (Diptera: Tephritidae) adventive in
RBC UMN SEI ere rarities a Lie Eee ees Sak Me Oe oe elie aon, eee:

521

349

376

325
117

746

vi PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

STIMMEL, J. F.—Seasonal history of the white peach scale, Pseudaulacaspis penta-
gona (Targ.-Tozz.) (Homoptera: Diaspididae), in northeastern Pennsylvania ........ 128

TALLERICO, P.—See CALABRESE, D. M.

THORPE, K. W.—Six Trichogramma (Hymenoptera: Trichogrammatidae) species

associated with a Maryland cornfield, with description of a new species ............ 16
THRELFALL, W.—Ectoparasites (Mallophaga, Acarina) from the double-crested cor-

morant(Phalacrocorax auritus) yin Florida cer we sate le ett eves cls tole o> heel eee ale 369
TODD, E. L.—The noctuoid moths of the Antilles—part II (Lepidoptera: Arctiidae:

BETIGOPINAE) bee ockessisy vol ovens ¢ Scehetsls Sis Sia sysie ete Cheer ee eC Ret tepeiane oer 315
VALLEY, K. R.—A new Liriomyza mining leaflets of black locust (Diptera: Agro-

BINV AUAE Ni aac 2 orto ele eieagas wie eee & in occa ple 6 abelnr = piapiapinnrs Guayelaaanasiaig Aa ee 781

VOSHELL, J. H., JR—See KONDRATIEFF, B. C.
WEBB, D. W.—Smittia lasiops (Malloch): A redescription of the adults with a descrip-

tion of the immature stages (Diptera: Chironomidae) ...................2sseceeee- 468
WHEELER, A. G., JR.—Coccobaphes sanguinarius and Lygocoris vitticollis (Hemip-

tera: Miridae): Seasonal history and description of fifth-instar, with notes on other

Mura s associated Gwithimaple: 3 5\sc.stye = sts eis: alee ie la isies ahi snewele a 177
WHEELER, A. G., JR.—Clanoneurum americanum (Diptera: Ephydridae), a leafminer
of theglittorallchenopod) Suaedailinéarisana sey ce Pe etree elie exes eevee ee 297

WHEELER, A. G., JR. and E. R. HOEBEKE—Psallus variabilis (Fallén) and P.
albipennis (Fallén), two European plant bugs established in North America, with
notes on taxonomic changes (Hemiptera: Heteroptera: Miridae) ................... 690

WHEELER, A. G., JR.—See HOEBEKE, E. R.

WHITEHEAD, D. R.—Foods of Caulophilus spp., particularly the broadnosed grain
weevil, C. oryzae (Gyllenhal), based on interception records (Coleoptera: Curculion-

[dae ts GOSSOnINAe) es ee Se eee ie oe ker na oh eee 81
WILKERSON, R. C. and G. B. FAIRCHILD—Two new species of Esenbeckia (Dip-
teras Tabanidae)| fromaMexicowtts2:.cee. Seer enrich ee Saco See 489

WILKERSON, R. C. and G. B. FAIRCHILD—Five new species of Diachlorus
(Diptera: Tabanidae) from South America with a revised key to species and new

Cavers golele) 1 | eed AT Ser aeo oc aa One UGC Ton OOEROmCcnootmoeere sco ncn sr 636
WIRTH, W. W.—The cacao-pollinating midges of the Forcipomyia argenteola group
(Diptera: Ceratopogonidae) az .sac cc seressc esate ene aeretaeters tems oon aa siacxale Aer ae 568

WIRTH, W. W.—See GILES, F. E.

WIRTH, W. W.—See GILES, F. E.

WOJTOWICZ, J. A.—Description of the larva and female of Pycnopsyche flavata
(Banks) with comparative notes on the ecology of P. flavata and P. gentilis
(McEachlan)i(@inchopterasleimnephilidac) mercer: os eine sie eaie eels eel eee 304

WRIGHT, C. G.—See SHERRON, D. A.

NOTES

BLANK, D. L.—See SHUBECK, P. P.
CALABRESE, D. M.—Hybridization between Gerris alacris Hussey and Gerris

comatus Drake and Hottes (Hemiptera: Heteroptera: Gerridae) in nature........... 209
GROGAN, W. L., JR.—New records of mites (Acari: Digamasellidae; Erythraeidae)

phoretic on biting midges (Diptera: Ceratopogonidae) ..................cceeee cece 211
HEPPNER, J. B—Change of name of a North American Ypsolopha (Lepidoptera:

OCD) i ee ee ae hn eee, Seay ener Rare akan ee waiae rs We 602

VOLUME 84, NUMBER 4 Vii

KINGSOLVER, J. M.—A 2000+-year-old beetle (Coleoptera: Dermestidae) .......... 390
KISSINGER, D. G.—Insect label production using a personal computer ............. 855
RIBBLE, D. W.—The status of Nomia foxii Dalla Torre (Hymenoptera: Halictidae) ... 208
RUSSELL, L. M.—The habits and appearance of a rare mealybug, Eurycoccus

blanchardii (King and Cockerell) (Homoptera: Coccoidea: Pseudococcidae) ........ 410
SHUBECK, P. P. and D. L. BLANK—Silphids attracted to mammal carrion at

C@heltenhams)Marylandi(Coleoptera: Silphidae) “22... -.-.e6e>c eck «2 soe tee ae 409
STAINES, C. L., JR.—Distributional records of Platypodidae (Coleoptera) in Mary-

ETI RRR PEI ote oie CO eee eee Ren OI Paci icc he renee orgie bar, rari aioe cae ice AE ORE 858
STEYSKAL, G. C.—An Oriental fly found in Trinidad (Diptera: Platystomatidae) ..... 859
VARSHNEY, R. K.—Ferrisia lobdellae, a new name for Ferrisia setosa (Lobdell,

1950) (@Hlomopterakeseudococcidae) mame he oe ocleciie is tein inle > ie oeisie ieteve ile let ae tle 857

Book REVIEWS
BATRA, S. W. T.—The Plant Protection Discipline, Problems and Possible Develop-

MEN tA STRATE RIES we cRP Sr Pcp lene che Teton ele cata. 5, Gos0015 (nederatos Made asepegs ca eytayee e's ea Negoneneatte dieters 412
BATRA, S. W. T.—Systematics of the Colletidae Based on Mature Larvae with
Phenetic Analysis of Apoid Larvae (Hymenoptera: Apoidea) ..................2-0. 413
GRISSELL, E. E.—Taxonomic Studies of the Encyrtidae with the Description of New
SpeclestandtasNew Genus ve coc actrees ss fechas stcuiec cps netche ONe eRe eee pean neler cate fe 652
NICKLE, D. A.—The North American Grasshoppers. Volume I, Acrididae: Gompho-
Ceninacrand A CHGINAC. fecrgachac Auccsheress Gots he eit er Kove UTR eee eee ae tee 861
NICKLE, D. A.—Faunal Affinities, Systematics and Bionomics of the Orthoptera of
thesGalifornia Channel Wslandsy <i. .ttocnce. cc's oe oe see cic e ce Spiele Ses ave Wiehe, sietereqele veel 862
POOLE, R. W.—The World of the Tent-Makers. A Natural History of the Eastern
fentsCatenpil karaay..6. 8c aeek sits PAS tals cee ccalend Sep ote ckeboeestictece, sree ona ek eoiteteers 651
OBITUARY
Luella Walkley Muesebeck—MUESEBECK, C. F. W. ............... sc ceecsssnaee 864

INDEX TO NEw TAXA, VOLUME 84

Acari borea Orth (Pherbellia), 28
Arborichthonius Norton, 86 calatheae Wirth (Forcipomyia), 573
styosetosus Norton (Arborichthonius), 86 californica Orth (Pherbellia), 25
dryas Fisher (Glaphyropyga), 717

Coleoptera habecki Wilkerson and Fairchild (Diachlorus),
atratus Kingsolver (Rhipibruchus), 675 646
bicolor Gordon (Nephaspis), 335 heppneri Wilkerson and Fairchild (Dia-
namolica Gordon (Nephaspis), 335 chlorus), 647

oedipygus Kingsolver (Rhipibruchus), 668 heveli Giles and Wirth (Serromyia), 442
psephenopygus Kingsolver (Rhipibruchus), — hirsutipalpus Wilkerson and Fairchild (Esen-

678 beckia (Esenbeckia)), 492
rugicollis Kingsolver (Rhipibruchus), 666 letica Wilkerson and Fairchild (Diachlorus),
; 641
PB leucotibialis Wilkerson and Fairchild (Dia-
arnaudi Orth and Fisher (Tetanocera), 685 chlorus), 645

biclausa Wilkerson and Fairchild (Esenbeckia — maculipennis Giles and Wirth (Serromyia),
(Ricardoa)), 490 444

Vill PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

marthae Orth (Pherbellia), 28

orientalis Giles and Wirth (Parabezzia), 823

paludum Orth (Pherbellia), 31

punctata Giles and Wirth (Serromyia), 446

robiniae Valley (Liriomyza), 781

schizonyx Giles and Wirth (Atrichopogon),
825

taprobane Mathis (Nocticanace), 421

trevori Wilkerson and Fairchild (Diachlorus),

644

ursilacus Orth (Pherbellia), 33

variegata Glick and Mullen (Alluaudomyia),
540

youngi Wirth (Forcipomyia (Forcipomyia)),
579

Hemiptera

aterrimus Kormilev (Aneurus), 482
ayersi Reichart (Anisops), 366
brailovskyi Kormilev (Aneurus), 480
carri Kittle (Trepobates), 157
cepula Henry (Labopidicola), 8
guanacastensis Kormilev (Mezira), 487
hondurensis Kormilev (Mezira), 485
machadus Rolston (Euschistus (Lycipta)),

284
piliger Kormilev (Notapictinus), 483
polhemi Kittle (Trepobates), 161
proxima Kormilev (Mezira), 486

Homoptera

aesta DeLong (Krisna), 610

Angubahita DeLong, 185

arta DeLong (Angubahita), 185

atrata DeLong (Frequenamia), 190

biflavus Cwikla and Freytag (Scaphytopius
(Cloanthanus)), 632

bilineatus DeLong (Portanus), 391

blockeri DeLong (Scaroidana), 611

canolaterus Cwikla and Freytag (Chloro-
tettix), 634

caudatus DeLong (Portanus), 392

confusa DeLong (Perubahita), 184

dubius DeLong (Xestocephalus), 394

hamatus DeLong (Tenucephalus), 611

incisus DeLong (Mendozellus), 615

lamina DeLong (Parabahita), 187

mitchellensis Smith (Hyalomyzus), 325

murrayae Cwikla and Freytag (Jdona), 635

particula DeLong (Frequenamia), 188

patula DeLong (Megabahita), 187

serratus DeLong (Mendozellus), 613

similis DeLong (Xestocephalus), 395

spinosus DeLong (Portanus), 392

tarandus DeLong (Mendozellus), 615

thalla DeLong (Parabahita), 188

variabilis DeLong (Xestocephalus), 393

Hymenoptera

alius Gibson (Urosigalphus (Urosigalphus)),
170

annulatus Marsh (Heterospilus), 850

cecidobroter Gordh and Hawkins (Tetra-
stichus), 426

diversus Gibson (Urosigalphus (Microuro-
sigalphus)), 167

ema Schauff and Grissell (Polynema), 530

flexus Gibson (Urosigalphus (Neouro-
sigalphus)), 172

marylandense Thorpe (Trichogramma), 18

megalopus Marsh (Heterospilus), 854

meridianus Gibson (Urosigalphus (Neouro-
sigalphus)), 100

minuta Smith (Nesoselandria), 125

monelliopsis Stary and Marsh (Trioxys (Tri-
oxys)), 726

porteri Gibson
sigalphus)), 174

surinamensis Gibson (Urosigalphus (Neo-
urosigalphus)), 175

yucatanensis Gibson (Urosigalphus (Neo-
urosigalphus)), 98

(Urosigalphus (Neouro-

Lepidoptera

carancahua Blanchard and Franclemont
(Marilopteryx), 271

chisosensis Blanchard and Franclemont
(Zale), 134

flinti Todd (Stenognatha), 316

kendallorum Blanchard and Knudson (Sym-
metrischema), 628

Marilopteryx Franclemont, 270

obsoletana Brown (Gretchena), 596

Plecoptera
frumi Kirchner (Allocapnia), 786
Oconoperla Stark and Stewart, 746

weaveri Stark and Stewart (Oconoperla),
750

VOL. 84 JANUARY 1982 NO. 1
; (ISSN 0013-8797)

"PROCEEDINGS

of the

INTOMOLOGICAL SOCIETY
ot WASHINGTON

DEPARTMENT OF ENTOMOLOGY
> SMITHSONIAN INSTITUTION
that WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY
J

CONTENTS
BLANCHARD, A. and J. G. FRANCLEMONT—A new species of Zale Hiibner from

Texas and New Mexico (Lepidoptera: Noctuidae: Catocalinae) ................. 134

DELONG, D. M.—New Central and South American leafhoppers of the **Bahita’* group
saomoptera: Cicadellidac: Deltocephalinac) .....--.-5..... 2.000 c5 eres tv neneus 184

GIBSON, L. P.—New species of Urosigalphus (Hymenoptera: Braconidae) from

GIBSON, L. P.—New species of Urosigalphus (Hymenoptera: Braconidae) from South
PUNRRCURIEEN Oreare e hne e t oy, BSin e St acdc as hid aie e oe Ce nas SoM a eee

) HEITZMAN, R. L.—Descriptions of the mature larva and pupa of Hypomecis umbro-
Mame ubrer). (Lepidoptera: Geometridae) « .). o <0. one =< ss nie peo nc Ses xe alan ess 111

_HENRY, C. S.—Reproductive and calling behavior in two closely related, sympatric
lacewing species, Chrysopa oculata and C. chi (Neuroptera: Chrysopidae) ....... 191
HENRY, T. J.—The onion plant bug genus Labopidicola (Hemiptera: Miridae): Eco-
nomic implications, taxonomic review, and description of a new species ......... |
HEPPNER, J. B.—Synopsis of the Glyphipterigidae (Lepidoptera: Copromorphoidea)
PREM Sree SS 2S Sr tre use Sani de ah es © ho we CS ke ie oan ae

KITTLE, P. D.—Two new species of water striders of the genus Trepobates Uhler
USSR RR ERS ch CRCERIRIAE Do ca eee ola One ein aie hea ach is oie at dle ek ete ei NE ca oi cll u 0 157

-KURCZEWSKI, F. E.—An additional study on the nesting behaviors of species of
REINER CLIVINCHOMCTA: SPHECIGAG) 620-08 oa cates aero oits ce Ca ce eens 67

KURCZEWSKI, F. E. and D. J. PECKHAM—Nesting behavior of Lyroda subita
EMME SUI CUCEMERE SDRECIOAR) Vo... J Gia oh wv dic sh aslvn aeieies aw + ds ie. wd Deitans

(Continued on back cover)

THE

ENTOMOLOGICAL SOCIETY

OF WASHINGTON

ORGANIZED MARCH 12, 1884

OFFICERS FOR 1982

MARGARET S. COLLINS, President
MANYA B. STOETZEL, President-Elect

RALPH E. HARBACH, Corresponding Secretary
F. CHRISTIAN THOMPSON, Treasurer

DavipD R. Smitu, Editor
Publications Committee
E. Eric GRISSELL

GEORGE C. STEYSKAL
JOHN M. KINGSOLVER WAYNE N. MATHIS

THOMAS E. WALLENMAIER

Honorary President
C. F. W. MUESEBECK

Honorary Members

FREDERICK W. Poos ASHLEY B. GURNEY RAYMOND A. St. GEORGE

All correspondence concernin

g Society business should be mailed to the appropriate officer at the following address: Entomologic:
Society of Washington, %

Department of Entomology, Smithsonian Institution, Washington, D.C. 20560.

MEETINGS. —Regular meetings of the Society are held in the Ecology Theater, Natural History Building, Smithsonian Institution, ¢
the first Thursday of each month from October to June, inclusive, at 8 P.M. Minutes of meetings are published regularly in th
Proceedings.

MEMBERSHIP. —Members shall be persons who have demonstrated interest in the science of entomology. Annual dues for membe
are $10.00 (U.S. currency) of which $9.00 is for a subscription to the Proceedings of the Entomological Society of Washington for on
year.

PROCEEDINGS. —Published quarterly beginning with January by the Society at Washington, D.C. Members in good standing recei
the Proceedings of the Entomological Society of Washington. Nonmember subscriptions are $18.00 per year, domestic, and $20.00 p
year, foreign (U.S. currency), payable in advance. All remittances should be made payable to The Entomological Society of Was
ington.

The Society does not exchange its publications for those of other societies.

Please see p. 183 of the January 1981 issue regarding preparation of manuscripts.

STATEMENT OF OWNERSHIP
Title of Publication: Proceedings of the Entomological Society of Washington.

Frequency of Issue: Quarterly (January, April, July, October).

Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Washington, % Departmer
of Entomology, Smithsonian Institution, 10th and Constitution NW, Washington, D.C. 20560.

Editor: Dr. David R. Smith, Systematic Entomology Laboratory, % U.S. National Museum NHB 168, Washington, D.C. 20560.
Managing Editor and Known Bondholders or other Security Holders: none.

This issue was mailed 16 February 1982
Second Class Postage Paid at Washington, D.C. and additional mailing office.

PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA

HELEN SOLLERS-RIEDEL, Hospitality Chairma}
Jay C. SHAFFER, Program Chairma\}

THOMAS E. WALLENMAIER, Recording Secretary Joyce A. UTMAR, Membership Chairma}y
JOHN F. CARROLL, Custodia

Jack E. Lipes, Delegate, Wash. Acad. Sci

|

/
|
|
|

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 1-15

THE ONION PLANT BUG GENUS LABOPIDICOLA (HEMIPTERA:
MIRIDAE): ECONOMIC IMPLICATIONS, TAXONOMIC
REVIEW, AND DESCRIPTION OF A NEW SPECIES

THOMAS J. HENRY

Systematic Entomology Laboratory, I[BIII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—The onion plant bug genus Labopidicola Kelton is reviewed,
and L. cepula from Texas is described as new. New distribution records
and known records for all species are given, and a summary of the economic
literature and speculation on pest potential are presented. Figures of male
genitalia of all species and the adult male of L. idahoensis and a key to the
species are provided to facilitate recognition.

Until recently all species of Labopidicola Kelton were placed in Labo-
pidea Uhler, a larger western genus. Knight (1923) described the first
species, allii, from Missouri where it was common on wild garlic. He later
recognized ainsliei from Iowa and planifrons from South Dakota (Knight,
1928) but gave no host data. Johnston (1930) described geminatus from a
long series collected on Allium canadense L. in Texas, and Knight (1968)
added the Sth species of this genus, idahoensis, again without a host as-
sociation. Kelton (1979) in his revision of the genus Labopidea discovered

that the above species were not congeneric with Uhler’s type-species, chlor-

—————Erreane eas

_ iza Uhler; hence, he erected Labopidicola to accommodate them and made

idahoensis the type of the genus. Tate (1940) first associated ainsliei with
Allium spp., and, more recently, Kelton (1979) reported this host relation-
ship for idahoensis and planifrons.

In this paper I summarize the economic literature, speculate on the pest
potential of the species in the genus, redescribe the known taxa, describe
one new species, figure the adult male of idahoensis and male genitalia of
all species, provide new state records for three species and known distri-
butions for all species, and give the first comprehensive key to separate
species of the genus.

The following abbreviations used are for institutions cited in this paper:
AMNH, American Museum of Natural History, New York; ORSU, Oregon
State University, Corvallis; PDA, Pennsylvania Department of Agriculture,

2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Harrisburg; PSU, Pennsylvania State University, University Park; TAM,
Texas A&M University, College Station; USNM, National Museum of Nat-
ural History, Washington, D.C.

ECONOMIC IMPLICATIONS

All species in this genus are restricted to feeding on onion and garlic,
Allium spp. There are numerous records of at least two species damaging
cultivated onions. The most frequently mentioned species, L. allii, has been
given the common name “‘‘onion plant bug’’ because it often becomes a
serious pest (Froeschner, 1949). Glenn (1923) provided the first life history
study in his paper on ‘‘The onion capsid, Orthotylus translucens Tucker’
(later recognized as L. allii by Henry and Wheeler, 1979). Glenn observed
this species as early as 1915 and noted that onion tops often were killed,
and that the mirids were sometimes so abundant that they covered the
ground as one walked through the fields.

Stiles (1937) recorded L. allii as a new pest for Oklahoma, and Smith and
Kelly (1937, 1938) and Bryson (1937a, 1937b, 1939) reported allii damaging
onions throughout Kansas. From 1934 to 1936 allii became ‘‘exceedingly
abundant’? in Missouri, and many plantings of Bermuda onions were de-
stroyed (Knight, 1941). Drake (1933, 1936, 1937a, 1937b, 1940), Haseman
(1941), and Smith et al. (1946) provided numerous other records of L. allii
causing injury. Populations as high as 15-20 bugs per plant in over 200 acres
of onions have been observed in Texas (U.S. Dep. Agric., 1955). The light
to medium damage reported in several Texas counties (U.S. Dep. Agric.,
1957) apparently is the most recent record of onion plant bugs as pests.
Damage to Moraine locust (a cultivar of honeylocust, Gleditsia triacanthos
L.) by L. ainsliei in North Carolina (U.S. Dep. Agric., 1959, 1960b) should
be attributed to Diaphnocoris chlorionis (Say), a species restricted to hon-
eylocust (Wheeler and Henry, 1976). The reports of Labopidea spp. on
sugar beets in Utah (Knowlton, 1933) and winter wheat in Montana (U.S.
Dep. Agric., 1960a) have not been examined but probably represent mis-
identifications (or true Labopidea spp.) or accidental host records.

Other than Westcott’s (1973) listing of L. allii as a pest on onions and my
new Pennsylvania record (Henry, 1977a), I am not aware of any recent
interest in these mirids as pests or of any additional reports of damage. It
is not clear why onion plant bugs apparently no longer reach damaging
levels. It may be that current cultivation practices and the use of more
modern pesticides or, possibly, the introduction of resistant varieties have
helped eliminate much of the threat to cultivated onions. Early remedies
like whale oil soap mixed with water (Glenn, 1923) and 2% sulphur mixed
with derris or diatomaceous earth (Drake, 1938) were said to give some
control, but it may be that simple cultural methods actually have minimized
the importance of onion bugs. Glenn (1923) noted that where fields were

VOLUME 84, NUMBER 1 3

burned over or plowed under, few mirids survived to the following year,
and Knight (1941), after establishing that L. allii overwintered as eggs de-
posited in stems, suggested that eliminating old stems and destroying nearby
wild Allium spp. would control onion plant bugs.

It is likely that these early season, univoltine species are unable to reinfest
onions successfully as many populations consist of a high proportion of
brachypterous females; this loss of flight would prevent easy migration of
large numbers of bugs from wild Allium spp. back into cultivated fields.

While the onion bugs have not attracted much attention in recent years,
the potential exists for members of this genus to reach damaging numbers
in commercial plantings and home gardens. If large reservoir patches of wild
Allium spp. are present and if poor sanitation practices are followed, onion
plant bugs could again become a major concern in the United States.

Labopidicola Kelton

Labopidicola Kelton, 1979: 757. Type-species: Labopidea idahoensis
Knight.

Diagnosis.—Generally small to medium-sized mirids, length 3.30-—4.70
mm in macropterous male and female, 2.75—3.20 mm in brachypterous fe-
male; body coloration yellow to bluish green, pubescence erect, suberect,
and recumbent silvery and brown to fuscous setae (darker setae often be-
coming bristle-like), intermixed with recumbent sericeous setae, especially
on head and pronotum. Head broad, distance across eyes wider than an-
terior margin of pronotum; vertex wide, nearly 3x the dorsal width of an
eye, base with a distinct carina reaching across to posterior angles of each
eye; rostrum short, stout, not reaching beyond apices of procoxae or middle
of sternum; pronotum trapeziform, posterior angles strongly rounded, basal
margin straight to weakly concave, middle of anterior margin often weakly
sinuate, calli distinct, weakly concave, area just posterior sunken or de-
pressed; mesoscutum broadly exposed, scutellum equilateral; hemelytra
uniformly opaque green; membrane translucent to opaque, usually fumate
or tinged with brown, totally brachypterous forms without hemelytral mem-
brane common (brachyptery has not been observed in males); venter and
legs uniformly greenish to testaceous. Male genitalia: Left paramere
C-shaped with left side often slender and curved in toward base; right par-
amere generally globose with a slender spine or arm extending anteriorly
when viewed in situ; aedeagus with 2-3 spiculi, ductus seminis thickened,
tapering apically, secondary gonopore usually subapical, middle of basal
margin of genital segment with a group of 2 or 3 irregularly shaped processes
that are hidden when parameres are in position.

Remarks.—Labopidicola are recognized by the overall green color, the
broad vertex with a distinct transverse basal carina, the short, thickened

4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

rostrum (Fig. 7) that does not reach beyond the middle of the sternum, the
two types of setae on the dorsum, and the male genitalia.

Labopidicola keys to Labopidea in Knight’s (1941: 81; 1968: 93) general
faunal studies and my (Henry, 1977b: 606) key to the green Orthotylinae of
the eastern United States, but species of Labopidicola can be distinguished
from Labopidea by the more distinct basal carina on the vertex, the tylus
or clypeus clearly visible from above, the shorter, stouter rostrum, and the
lack of fuscous markings on the head, calli, and antennae. Kelton (1980)
provided a key to separate Labopidicola from other genera in the Canadian
Prairie Provinces.

KEY TO SPECIES OF LABOPIDICOLA!

1. Hemelytra clothed with pale and/or silvery and erect, brown to fus-

cOus*setae (Setae may. become: bristle-like)y sia. 0.5 ich clo vee 2
— Hemelytra clothed with only pale or silvery setae, at most, a few
darker setae scattered on basal areas of coria and clavi ............ 4

2. Small species, length in macropterous male less than 3.80 mm; dor-
sum clothed with both long, erect, rather fine, pale and brown to
fuscous setae; length of 4th antennal segment much greater than
length of Ist antennal segment; male genitalia (Fig. 4); Tex. ......

TEAR. LANs DRT He SUT ETE ae pa geminatus (Johnston)

— Larger species, length in macropterous male averaging 4.00 mm or
more; dorsum clothed with numerous, rather short, dark, bristle-like
setae; length of 4th antennal segment subequal to length of Ist seg-
MIEMO PAs SRR SEK, DCS Te Se SE OR eet ee 3

3. Length of 2nd antennal segment greater than basal width of prono-
tum by more than the dorsal width of 2 eyes; male genitalia (Fig. 5);
Ida rGolis BC .AltassSacskeyMan® : stiiaren. idahoensis (Knight)

— Length of 2nd antennal segment greater than basal width of prono-
tum by less than the dorsal width of | eye; male genitalia (Fig. 3);
PRR ES asst PRUNE eS eas, eee) Deed © cepula, new species

4. Dorsum clothed only with short, recumbent pale setae; male geni-
taliar(rie ol); lly dace Mich. sPasTenieek tick dec ainsliei (Knight)

— Dorsum clothed with short, recumbent, or suberect, and long erect

PUDESEENCE WA Sosy eee Le see Oe) wae ae ee 5

Length of 3rd antennal segment almost equal to basal width of

pronotum in male; length of pronotum and width of vertex subequal:

male genitalia (Fig. 2); Utah east to Pa. and N.C. ...... allii (Knight)

‘n

' This key works best with well-preserved, unrubbed males. In specimens having the dorsal
pubescence clearly disturbed, only male genitalia will accurately separate species; brachyp-
terous females are best identified by their association with males.

VOLUME 84, NUMBER | 5

Length of 3rd antennal segment clearly less than basal width of
pronotum in male; length of pronotum less than width of vertex;
male genitalia (Fig. 6); Ia., S.D.; Man., Sask. .... planifrons (Knight)

Labopidicola ainsliei (Knight)
Fig. |

Labopidea ainsliei Knight, 1928: 235; Knight, 1941: 105; Froeschner, 1949:
171; Carvalho, 1958: 81.
Labopidicola ainsliei: Kelton, 1979: 757.

Diagnosis.—Male length 3.60 mm (range of 5 specimens 3.56—3.76 mm,
¥ = 3.63 mm), width 1.20 mm. General color yellow to yellowish green;
dorsum only with short, recumbent, pale, simple setae, sparsely intermixed
with silvery sericeous pubescence. Head: Width 0.86 mm, vertex 0.50 mm.
Rostrum: Length 0.62 mm, reaching apices of procoxae. Antenna: Uni-
formly yellowish or testaceous, segments III and IV more brownish; I,
length 0.32 mm; IJ, 1.12 mm; HI, 0.80 mm; IV, 0.54 mm. Pronotum: Length
0.42 mm, basal width 0.92 mm. Hemelytron: Uniformly yellowish to yel-
lowish green, with short, recumbent simple setae, intermixed with a few
silvery sericeous setae; membrane translucent, shaded with brown. Venter
and legs yellowish or pale yellowish green. Male genitalia: Fig. 1.

Macropterous female.—Length 3.32—3.52 mm (x* = 3.42 mm for 2 speci-
mens). All specimens available are very teneral and not measurable beyond
lengths. Very similar to male in color and general form.

Brachypterous female.—Length 2.76 mm, with apices of hemelytra reach-
ing apex of abdomen, width 1.20 mm. Head: Width 0.86 mm, vertex 0.50
mm. Rostrum: Length 0.70 mm, reaching apices of procoxae. Antenna: I,
Length 0.30 mm; II, 1.06 mm; III, 0.90 mm; IV, 0.42 mm. Pronotum: Length
0.40 mm, basal width 0.90 mm.

Specimens examined.—IOWA: Holotype 6, Sioux City, June 25, 1925,
C. N. Ainslie coll. (USNM); 5 od, 1 macropterous 2, Ames, June 8-10,
1929, H. H. Knight and H. M. Harris colls., on onion (USNM); 3 6, 2
macropterous 2°, Ames, June 1, 1933-34, on wild onion (TAM). MICHI-
GAN: 2 6, 2 macropterous @, 2 brachypterous ° , May 1942, E. I. McDaniel
coll., on chives (USNM). PENNSYLVANIA: | ¢, Howard, June 18, 1978,
V. Haas coll., from light trap (PSU). TENNESSEE: 2 6, Hamilton Co.,°
May 10, 1942, Turner coll., in light trap at edge of peach orchard (USNM).

Remarks.—Labopidicola ainsliei is best recognized by the short, recum-
bent, pale setae on the hemelytra and by the male genitalia. The right par-
amere (Fig. Id) has the apex rather broadened; the left paramere is strongly
C-shaped.

This species, previously known only from Illinois and lowa (Knight,
1941), is now recorded from Michigan, Pennsylvania, and Tennessee.

6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Labopidicola allii (Knight)
Fig. 2

Orthotylus translucens Glenn (nec Tucker), 1923: 79.

Labopidea allii Knight, 1923: 31; Knight, 1941: 105; Froeschner, 1949: 171;
Carvalho, 1958: 81; Henry, 1977a: 417.

Labopidicola allii Kelton, 1979: 757.

Diagnosis.—Male, length 3.67 mm (range of 15 specimens 3.60-4.32 mm,
¥ = 3.76 mm), width 1.36 mm. General color green to bluish green, thickly
clothed with erect and semierect simple setae, intermixed with silvery, se-
riceous pubescence, especially on head and pronotum. Head: Width 0.78
mm, vertex 0.48 mm, thickly clothed with silvery sericeous setae, especially
along margins of eyes, median line, and on tylus. Rostrum: Length 0.70
mm, reaching apices of procoxae. Antenna: Green to testaceous, segment
II testaceous, tinged with green, III and IV brownish; I, length, 0.38 mm;
II, 1.16 mm; III, 1.04 mm; IV, 0.46 mm. Pronotum: Length 0.48 mm, basal
width 1.00 mm, with scattered sericeous setae, lateral and anterior margins
and median line with distinct rows of silvery sericeous setae. Hemelytron:
Uniformly green, thickly clothed with erect and semierect pale setae, inter-
mixed with a few scattered sericeous setae; membrane translucent brown,
veins green. Venter green; legs green with apical halves of tibiae and all of
metafemur becoming more testaceous. Male genitalia: Fig. 2.

Macropterous female.—Length 4.28 mm (range of 15 specimens 4.08-4.64
mm, *« = 4.32 mm), width 1.48 mm. Head: Width 0.92 mm, vertex 0.56 mm.
Rostrum: Length 0.84 mm, reaching apices of procoxae. Antenna: I, Length
0.38 mm; II, 1.20 mm; III, 0.94 mm; IV, 0.36 mm. Pronotum: Length 0.50
mm, basal width 1.18 mm.

Brachypterous female.—Length 3.25 mm (range of 5 specimens 2.84—3.40
mm, < = 3.08 mm), length to apex of wings 2.92 mm (range of 5 specimens
2.60-3.14 mm, * = 2.80 mm), wings reaching anterior margin of 8th abdom-
inal tergum. Head: Width 0.94 mm, vertex 0.60 mm. Rostrum: Length 0.78
mm. Antenna: I, Length 0.32 mm; I, 1.02 mm; III, 0.92 mm; IV, 0.50 mm.
Pronotum: Length 0.40 mm, basal width 0.96 mm.

Specimens examined.—ARKANSAS: 1 macropterous 9, Washington
Co., May 29, 1964, C. E. McCou coll. (USNM). ILLINOIS: 1 6, 1 mac-
ropterous 2, Olney, June 1915, on onion (USNM); | ¢, Urbana, 1923, P.
A. Glenn coll. (USNM). INDIANA: 2 macropterous 2, 1 brachypterous

Lex

Figs. 1-3. Male genitalia. 1, Labopidicola ainsliei. 2, L. allii. 3, L. cepula. a. Left para-
mere, lateral aspect in situ. b, Left paramere, lateral aspect. b’, Left paramere, inside lateral
aspect. c, Left paramere, dorsal aspect. d, Right paramere, lateral aspect. e, Ventral process.
f, Aedeagus.

VOLUME 84, NUMBER 1

8 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2, Posey Co., June 6, 1931, on onions (USNM). IOWA: 2 macropterous
2, Ames, June 8-10, 1929, H. H. Knight coll. (USNM); 1 ¢, 3 macropterous
2, Ames, May 8, 1933-34, on onions (TAM); | ¢, 6 macropterous 2°,
Bloomfield, May 5, 1936, F. Andre coll. (USNM); | ¢, | macropterous
2, Bloomfield, June 15, 1937, F. Andre coll. (USNM); 2 macropterous @,
2 brachypterous 2, Little Rock, July 2, 1897 (USNM); | d, Pleasant Valley,
May 16, 1930, on onions (USNM); | brachypterous 9, 6 mi. NW Ledyard,
July 7, 1928, G. D. Hendrickson coll. (USNM). KANSAS: | 3, 3 macrop-
terous 2, Manhattan, May 6, 1936, R. H. Painter coll. (\USNM). KEN-
TUCKY: | 3, 1 macropterous °, Lexington, May 10, 1938, L. H. Town-
send coll., on onions (TAM). MICHIGAN: | brachypterous °, Rochester,
May 28, 1942 (USNM). MISSOURI: 2 macropterous @ (including holotype),
6 Sth-instar nymphs, Jackson, Apr. 1921, A. C. Burrill coll., on wild garlic
(USNM). NORTH CAROLINA: 2 brachypterous 2, Union Co., Monroe,
May 25, 1980, A. G. Wheeler, Jr. coll., on Allium vineale L. (PDA). OKLA-
HOMA: 3 6, 3 macropterous 2, Chickasha, Apr. 1928, M. W. Shakleford
coll. (USNM). PENNSYLVANIA: 2 ¢, 13 macropterous ?, Dauphin Co.,
Clark’s Valley, May 19, 1977, T. J. Henry coll., on Allium sp. (PDA,
USNM); | ¢, Howard, Centre Co., June 1978, V. Haas coll., at light (PSU).
UTAH: 2 6, Promontory, May 25, 1952, G. F. Knowlton coll. (USNM).
VIRGINIA: 7 3,3 2, King George Co., Rt. 206 just E of Stafford Co. line,
May 8, 1981, A. G. Wheeler, Jr. coll., on Allium sp. WEST VIRGINIA: 4
6,9 macropterous 2, Grant Co., Shale Barrens, Rt. 28, 1.2 mi. W Peters-
burg, May 29, 1978, T. J. Henry and A. G. Wheeler, Jr. colls., on Allium
sp. (PDA, USNM).

Remarks.—Labopidicola allii is best separated from other Labopidicola
species by the thickly pubescent dorsum that lacks darker setae and by the
left paramere that is broadened laterally forming a mitten-shaped pattern.

Labopidicola allii is the most common and widespread member of the
genus. This species, previously known from Illionois, Indiana, lowa, Kan-
sas, Missouri, Oklahoma (Kelton, 1979), and Pennsylvania (Henry, 1977a),
is now recorded from Arkansas, Kentucky, Michigan, Utah, Virginia, and
West Virginia.

Labopidicola cepula Henry, New Species
Fig. 3

Diagnosis.—Holotype male, length 4.36 mm (length of male paratype 4.28
mm), width ca. 1.36 mm (wings slightly spread). General color yellow green;
pronotum, scutellum, and hemelytra with erect and semierect dark bristle-
like setae, intermixed with recumbent, silvery sericeous setae. Head: Width
0.82 mm, vertex 0.50 mm. Rostrum: Length 0.78 mm, reaching middle of
sternum. Antenna: Testaceous; I, length 0.42 mm; II, 1.18 mm; III, 0.98
mm; 1V, 0.42 mm. Pronotum: Length 0.46 mm, basal width 1.12 mm. Hem-

VOLUME 84, NUMBER 1 9

elytron: Uniformly yellowish green; membrane translucent brown. Venter
and legs yellowish to yellowish green. Male genitalia: Fig. 3.

Allotype.—Macropterous female: Length 4.08 mm (range of 6 specimens
3.88-4.12 mm, « = 4.02 mm), width 1.56 mm. Head: Width 0.88 mm, ver-
tex, 0.58 mm. Rostrum: Length 0.84 mm. Antenna: I, Length 0.36 mm; II,
1.20 mm; III, 0.90 mm; IV, 0.42 mm. Pronotum: Length 0.48 mm, basal
width 1.14 mm.

Brachypterous female.—Length 3.56 mm (range of 3 specimens 3.56—3.68
mm, <« = 3.61 mm), wing tips reaching 9th abdominal tergum, width 1.48
mm. Head: Width 0.88 mm, vertex 0.56 mm. Rostrum: Length 0.82 mm.
Antenna: I, Length 0.42 mm; II, 1.16 mm; III, 0.90 mm; IV, 0.46 mm.
Pronotum: Length 0.46 mm, basal width 1.06 mm.

Type-material.—Holotype ¢, Bastrop Co., Texas, Bastrop State Park,
May 8, 1966, J. C. Schaffner coll., taken on Allium sp. (USNM type no.
76881). Allotype female, same data as holotype (USNM). Paratypes: 1 6,
5 macropterous 2, 4 brachypterous 2, same data as holotype (TAM,
USNM).

Remarks.—Labopidicola cepula resembles allii in male genitalia, but is
most similar to idahoensis in dorsal pubescence. It is interesting that a male
and female of geminatus were also collected at Bastrop Co., Texas, Bastrop
State Park (May 8, 1966 and May 5, 1968 by Dr. Schaffner).

Etymology.—The specific name cepula is from the Latin feminine dimin-
utive for onion.

Labopidicola geminatus (Johnston)
Fig. 4
Labopidea geminata Johnston, 1930: 298; Carvalho, 1958: 82.
Labopidicola geminata: Kelton, 1979: 758.

Diagnosis.—Male, length 3.52 mm (range of 9 specimens 3.12—3.76 mm,
¥ = 3.52 mm), width 1.16 mm. General color yellowish to pale green; dor-
sum with numerous scattered silvery setae, intermixed with erect, brown
to fuscous setae. Head: Width 0.74 mm, vertex 0.40 mm, thickly clothed
with silvery sericeous setae, with distinct rows forming along inside margins
of eyes and on median line, vertex scattered with a few long, erect darker
setae. Rostrum: Length 0.60 mm, reaching apices of procoxae. Antenna:
Segment I green, II testaceous, II] and IV brown; I, length 0.28 mm; II,
1.00 mm; III, 0.78 mm; IV, 0.42 mm. Pronotum: Length 0.42 mm, basal
width 0.94 mm, thickly clothed with silvery, sericeous setae, especially
along median line and on anterior margin, intermixed with erect darker
setae; scutellum and mesoscutum with scattered silvery setae and a single
row of these setae along median line. Hemelytron: Green with erect pale
and brown to fuscous setae, intermixed with recumbent silvery setae; mem-

10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

brane fumate. Venter yellowish green; legs testaceous to yellowish green.
Male genitalia: Fig. 4.

Macropterous female.—Length 3.76 mm (range of 6 specimens 3.44—3.84
mm, <* = 3.64 mm), width 1.36 mm. Head: Width 0.80 mm; vertex 0.52
mm. Rostrum: Length 0.70 mm, reaching just beyond apices of procoxae
or to middle of mesosternum. Antenna: I, Length 0.32 mm; II, 0.88 mm;
III, 0.72 mm; IV, 0.44 mm. Pronotum: Length 0.42 mm, basal width 1.02 mm.

Brachypterous female.—Length 3.08 mm (range of 5 specimens 3.04-3.28
mm, + = 3.08 mm), length to apices of wings 2.44 mm (range 2.44—2.68 mm,
¥ = 2.56 mm), apices of wings reaching anterior margin of 7th abdominal
tergum. Head: Width 0.80 mm, vertex 0.52 mm. Rostrum: Length 0.72 mm.
Antenna: 1, Length 0.30 mm; II, 0.94 mm; III, 0.80 mm; IV, 0.44 mm.
Pronotum: Length 0.36 mm, basal width 0.90 mm.

Specimens examined.—TEXAS: 3 d, Bosque Co., 2 miles W Iredell,
Apr. 24, 1970, J. C. Schaffner coll. (TAM); | ¢, 4 brachypterous 2, Brazos
Co., Minter Springs, Apr. 12, 1970, V. V. Board coll. (TAM); 3 6, Brazos
Co., College Station, Apr. 6, 1964, H. R. Burke coll., at light (TAM); 2
36, Brazos Co., College Station, Apr. 15, 1978, T. J. Henry, J. C. Schaffner,
R. T. Schuh colls., taken on Allium sp. (USNM); 8 6, Burnett Co., Inks
Lake St. Park, Apr. 25-28, 1965-1968, J. C. Schaffner coll. (TAM); 7 3, 6
macropterous °, Dimmit Co., Jan. 31, 1946, H. T. Hibbs coll. (TAM,
USNM); 18 3, 4 macropterous 2, 8 brachypterous 2, Gonzales Co., Pal-
metto State Park, Mar. 26—May 4, 1964-70, V. V. Board, H. R. Burke, J.
C. Schaffner colls. (TAM, USNM); 1 6, 1 macropterous 2, 3 brachypterous
2 (paratypes), Grimes Co., Huntsville, Apr. 4, 1928, H. G. Johnston coll.
(USNM); 1 36, Hunt Co., Apr. 23, 1954, C. F. Garner coll. (USNM).

Remarks.—Labopidicola geminatus was described as the ‘‘southern
twin” of planifrons, but actually is closer to idahoensis based on the pres-
ence of darkened setae on the dorsum. It differs from idahoensis in the
longer dorsal pubescence and proportionately shorter second antennal seg-
ment, and by the male genitalia.

The name geminata is here amended to the masculine geminatus. Ac-
cording to George Steyskal, Cooperating Scientist of the Systematic Ento-
mology Laboratory, USDA, because Kelton (1979) did not provide the
meaning of Labopidicola, and because words ending in -icola form nouns
of common gender, this genus must be considered masculine.

Labopidicola idahoensis (Knight)
a (ee Pe Ae

Labopidea idahoensis Knight, 1968: 97.
Labopidicola idahoensis: Kelton, 1979: 757; Kelton, 1980: 245.

VOLUME 84, NUMBER 1 11

Figs. 4-6. Male genitalia. 4, Labopidicola geminatus. 5, L. idahoensis. 6, L. planifrons.
a, Left paramere, lateral aspect in situ. b, Left paramere, lateral aspect. c, Left paramere,
dorsal aspect. d, Right paramere, lateral aspect. e, Ventral process. f, Aedeagus.

12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Diagnosis.—Male, length 4.02 mm (range of 15 specimens 3.80-4.28 mm,
¥ = 4.05 mm), width 1.28 mm. General color yellowish green; dorsum with
scattered silvery, sericeous setae and rather short, bristle-like fuscous setae.
Head: Width 0.86 mm, vertex 0.52 mm. Rostrum: Length 0.76 mm, reach-
ing just past apices of procoxae to middle of mesosternum. Antenna: Green-
ish yellow, segments III and IV becoming brown; I, length 0.44 mm; II,
1.38 mm; III, 0.88 mm; IV, 0.48 mm. Pronotum: Length 0.46 mm, basal
width 1.00 mm, with recumbent, silvery, sericeous setae, thickly intermixed
with fuscous bristle-like setae. Hemelytron: Yellowish green with numer-
ous, short, fuscous bristle-like setae intermixed with a few simple pale setae;
membrane smoky translucent to fumate. Venter and legs yellowish green.
Male genitalia: Fig. 5.

Macropterous female.—Not examined.

Brachypterous female.—Length 3.96 mm (range for 15 specimens 3.36—
4.00 mm, < = 3.76 mm), length to apices of wings 2.96 mm (range for 15
specimens 2.76—3.20 mm, ¥ = 2.98 mm), width 1.32 mm. Head: Width 0.94
mm, vertex 0.60 mm. Rostrum: Length 0.80 mm, reaching just beyond
procoxae to middle of mesosternum. Antenna: I, Length 0.44 mm; II, 1.32 mm;
III, 0.96 mm; IV, 0.46 mm. Pronotum: Length 0.42 mm, basal width 0.96
mm.

Specimens examined.—IDAHO: holotype 6, | brachypterous 2°, Ge-
nesse, alt. 2700’, May 5, 1937, V. E. Nygren coll. (USNM). OREGON: 27
3d, 32 brachypterous ¢, Clatsop Co., Saddle Mt. State Park, June 14, 1979,
R. T. Schuh coll., on Allium sp. (AMNH, USNM). UTAH: 2 brachypterous
2, Cache Jct., June 11, 1903, no coll. data (USNM); | brachypterous 2°,
Ogden, June 9, 1927, G. F. Knowlton coll. (USNM).

Remarks.—Labopidicola idahoensis is most similar to geminatus in ap-
pearance but can be separated from it by the shorter, more thickly set, dark,
bristle-like setae found on the dorsum (Fig. 6) and other characters given
in the key.

Labopidicola idahoensis, previously known from Idaho (Knight, 1968)
and Colorado, British Columbia, Alberta, Saskatchewan, and Manitoba
(Kelton, 1979), is now recorded from Oregon and Utah.

Labopidicola planifrons (Knight)
Fig. 6
Labopidea planifrons Knight, 1928: 234; Knight, 1941: 105; Carvalho, 1958:
82.
Labopidicola planifrons: Kelton, 1979: 754; Kelton, 1980: 246.
Diagnosis.—Male, length 4.04 mm, width ca. 1.36 mm. General color pale

greenish, posterior /2 of pronotum, hemelytral margins, setal punctures and
legs tinged with blue green, dorsum clothed with erect and suberect, silvery,

VOLUME 84, NUMBER I 13

Figs. 7, 8. Labopidicola idahoensis. 7, Dorsal habitus of adult male. 8, Lateral aspect of
head and pronotum.

14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

simple setae, sparsely intermixed with silvery sericeous pubescence. Head:
Width 0.84 mm, vertex 0.52 mm. Rostrum: Length 0.70 mm, reaching just
beyond procoxae to middle of sternum. Antenna: Segment I greenish, II
and III testaceous; I, length 0.36 mm; II, 1.00 mm; III, 0.88 mm; IV, broken.
Pronotum: Length 0.44 mm, basal width 1.02 mm. Hemelytron: Uniformly
pale green, tinged along margins and on setal punctures with blue green;
membrane translucent brownish. Venter and legs pale greenish. Male gen-
italia: Fig. 6.

Remarks.—I have examined only the holotype of planifrons. It is nearest
to allii in general color and pubescence but can be separated by the char-
acters given in the key. This is the only species with a strongly recurved
hook on the right arm of the left paramere (Fig. 6c).

Labopidicola planifrons was described from South Dakota and later re-
ported from Iowa, Manitoba, and Saskatchewan (Kelton, 1979).

ACKNOWLEDGMENTS

I thank the following people and their institutions for kindly lending spec-
imens: K. C. Kim (PSU), J. C. Schaffner (TAM), R. T. Schuh (AMNH),
M. A. Schwartz (ORSU), and A. G. Wheeler, Jr. (PDA). I also thank George
Steyskal (SEL) for his expert interpretation of the gender for Labopidicola.

LITERATURE CITED

Bryson, H. R. 1937a. A plant bug (Labopidea allii Knight). U.S. Dep. Agric. Insect Pest
Surv. Bull. 17: 186.

— . 1937b. A plant bug (Labopidea allii Knight). U.S. Dep. Agric. Insect Pest Surv. Bull.

17: 245.

. 1939. Onion plant bug (Labopidea allii Knight). U.S. Dep. Agric. Insect Pest Surv.

Bull. 19: 151.

Carvalho, J. C. M. 1958. Catalogue of the Miridae of the world. Part III. Subfamily Ortho-
tylinae. Arq. Mus. Nac., Rio de J. 47: 1-161.

Drake, C. J. 1933. Biology and control of onion insects. Report of agriculture research for the
year ending June 30, 1933. Agric. Exp. Stn., lowa State Coll. Agric. Mech. Arts 1933:

71-72.

. 1936. A mirid (Labopidea allii Knight). U.S. Dept. Agric. Insect Pest Surv. Bull. 16:
193.

. 1937a. A plant bug (Labopidea allii Knight). U.S. Dep. Agric. Insect Pest Surv. Bull.
17: 186.

—. 1937b. Biology and control of onion insects. Report of agriculture research for the
year ending June 30, 1937. Part I. Agric. Exp. Stn., lowa State Coll. Agric. Mech. Arts
1937: 141-142.

—. 1938. Biology and control of onion insects. Report on agriculture research for the

year ending June 30, 1938. Part I. Agric. Exp. Stn., Iowa State Coll. Agric. Mech. Arts

1938: 135.

1940. Biology and control of onion insects particularly as vectors in yellow dwarf
disease. Report on agriculture research for year ending June 30, 1940. Part II. Agric.
Exp. Stn., lowa State Coll. Agric. Mech. Arts 1940: 151.

VOLUME 84, NUMBER 1 15

Froeschner, R. C. 1949. Contribution to a synopsis of the Hemiptera of Missouri. Pt. 1V. Am.
Midl. Nat. 42: 123-188.
Glenn, P. A. 1923. The onion capsid, Orthotylus translucens Tucker. J. Econ. Entomol. 16:

79-81.

Haseman, L. 1941. A mirid (Labopidea allii Knight). U.S. Dep. Agric. Insect Pest Surv. Bull.
21; 190.

Henry, T. J. 1977a. A mirid (Labopidea allii). U.S. Dep. Agric. Coop. Plant Pest Rep. 2(24):
417.

. 1977b. Orthotylus nassatus, a European plant bug new to North America (Heterop-

tera: Miridae). U.S. Dep. Agric. Coop. Plant Pest Rep. 2(31): 605-608.

Henry, T. J. and A. G. Wheeler, Jr. 1979. Orthotylus translucens: Taxonomic status and
correction of published misidentifications (Hemiptera: Miridae). Proc. Entomol. Soc.
Wash. 81: 60-63.

Johnston, H. G. 1930. Four new species of Miridae from Texas (Hemiptera). Bull. Brooklyn
Entomol. Soc. 25: 295-300.

Kelton, L. A. 1979. Labopidea Uhler in North America with descriptions of a new species
and new genus (Heteroptera: Miridae). Can. Entomol. 111: 753-758.

———. 1980. The plant bugs of the prairie provinces of Canada. Heteroptera: Miridae. Bio-
syst. Res. Inst., Ottawa. 408 pp.

Knight, H. H. 1923. A new species of Labopidea on garlic (Heteroptera-Miridae). Bull.

Brooklyn Entomol. Soc. 18: 31.

. 1928. New species of Labopidea and Macrotyloides (Hemiptera, Miridae). Can. Ento-

mol. 60: 233-236.

—. 1941. The plant bugs, or Miridae of Illinois. Ill. Nat. Hist. Surv. Bull. 22, 234 pp.

—. 1968. Taxonomic review: Miridae of the Nevada Test Site and the western United
States. Brigham Young Univ. Sci. Bull. 9(3): 282 pp.

Knowlton, G. F. 1933. Notes on Utah Heteroptera. Entomol. News 44: 261-264.

Smith, R. C., G. A. Dean, and E. G. Kelly. 1946. The fifteenth or 1945 annual insect population
summary of Kansas. J. Kans. Entomol. Soc. 19: 37-52.

Smith, R. C. and E. G. Kelly. 1937. The sixth annual insect population summary of Kansas
covering the year 1936. J. Kans. Entomol. Soc. 10: 113-132.

—. 1938. The seventh annual insect population summary of Kansas, covering the year
1937. J. Kans. Entomol. Soc. 11: 54-76.

Stiles, C. F. 1937. A plant bug (Labopidea allii Knight). U.S. Dep. Agric. Insect Pest Surv.
Bull. 17: 186.

Tate, H. D. 1940. Insect vectors of yellow dwarf, a virus disease of onion. Iowa State Coll.
J. Sci. 14: 267-294.

U.S. Dep. Agric. 1955. A mirid (Labopidea allii). U.S. Dep. Agric. Coop. Econ. Insect Rep.

iets mirid (Labopidea allii). U.S. Dep. Agric. Coop. Econ. Insect Rep. 8(13):

‘i

ane A mirid (Labopidea ainsliei). U.S. Dep. Agric. Coop. Econ. Insect Rep. 9(25):
eyed A mirid (Labopidea sp.). U.S. Dep. Agric. Coop. Econ. Insect Rep. 10(31):
Une A mirid (Labopidea ainsliei). U.S. Dep. Agric. Coop. Econ. Insect Rep.

10(42): 987.

Westcott, C. 1973. The gardener’s bug book. 4th ed. Doubleday and Co., Inc. 689 pp.

Wheeler, A. G., Jr. and T. J. Henry. 1976. Biology of the honeylocust plant bug, Diaphnocoris
chlorionis, and other mirids associated with ornamental honeylocust. Ann. Entomol.
Soc. Am. 69: 1095-1104.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 16-22

SIX TRICHOGRAMMA (HYMENOPTERA: TRICHOGRAMMATIDAE)
SPECIES ASSOCIATED WITH A MARYLAND CORNFIELD,
WITH DESCRIPTION OF A NEW SPECIES‘?

KEVIN W. THORPE

Beneficial Insect Introduction Laboratory, IIBIII, Agric. Res. Serv.,
USDA, Bldg. 417, BARC-East, Beltsville, Maryland 20705; Maryland Cen-
ter for Systematic Entomology, Department of Entomology, University of
Maryland, College Park, Maryland 20742.

Abstract.—Trichogramma species occurring along the edge of a 3 ha
stand of field corn in the Beltsville Agricultural Research Center, Beltsville,
Maryland, were surveyed during the 1980 season by maintenance of He-
liothis virescens (F.) egg cloths in ten locations. A total of 1931 eggs from
79 egg cloths was parasitized by Trichogramma spp. Six species of Tricho-
gramma were collected. These species were, in descending order of abun-
dance, 7. minutum Riley, T. pretiosum Riley, T. exiguum Pinto and Platner,
T. parkeri Nagarkatti and 7. marylandense, n. sp., and T. retorridum (Gi-
rault). Trichogramma marylandense is described and illustrated.

Trichogramma wasps have been mass reared and released against insect
pests for over 70 years (see Ridgway et al., 1981, for a review). Some recent
examples of successful biological control of pests with Trichogramma are
reported by Oatman and Platner (1978) on tomato, Parker (1971) on cabbage,
and Ables et al. (1979) on cotton. However, in spite of recent advances in
Trichogramma taxonomy (Nagarkatti and Nagaraja, 1971; Pinto et al.,
1978), which have made possible the accurate identification of most of the
known North American species, little effort has been made to determine the
endemic species of Trichogramma in agricultural areas. Surveys of Tricho-
gramma species are now needed to obtain information on their distribution,

' Scientific Article No. A2970. Contribution No. 6030 of the Md. Agric. Exp. Stn., College
Park.

* From a dissertation to be submitted to the Graduate School, University of Maryland, by
the author in partial fulfillment of the requirements for the Ph.D. degree in Entomology.

3’ Mention of a commercial product in this paper does not constitute an endorsement of this
product by the USDA.

VOLUME 84, NUMBER 1 17

biology, and ecology so that their potential for use as biological control
agents can be evaluated.

During 1980 and 1981 I surveyed the species of Trichogramma occurring
in and around a field of corn at the Beltsville Agricultural Research Center,
Beltsville, Maryland. My objectives were to determine what species were
present and to obtain information on their relative abundance.

MATERIALS AND METHODS

In 1980 I surveyed the Trichogramma species by placing ten strips of
cloth (except on sampling dates | and 2 when only two cloths were used),
each containing ca. 100 naturally oviposited Heliothis virescens (F.) eggs,
in ten locations around the perimeter of a 3 ha stand of field corn. Four of
the locations were adjacent to a 0.2 ha plot of sweet corn (variety NK 199)
within the larger field. The phenology of the corn was as follows: The field
corn was planted 27 May, tasseled 21 July, was in silk from 29 July to 8
August, and was machine harvested 22 September; the sweet corn plot
contained staggered plantings so that all stages of growth were continuously
present. Sweet corn was planted on 30 May and every week thereafter. The
first planting tasseled on 10 July and was in silk from 18 to 28 July. The
sweet corn was not harvested.

Egg cloths were maintained in the sample area from 7 May to 7 November
1980. The cloths were attached to a 5 cm wire ringed with Tanglefoot® to
limit access to egg predators and then suspended from stakes or branches
at a height of ca. 1 m. The egg cloths were replaced every 3—5 days and
held in the laboratory for emergence of parasitoids. Prior to use all H.
virescens eggs were killed by exposure to ca. 30 krad of gamma radiation.
Identifications were made in collaboration with Carll Goodpasture, formerly
of the Beneficial Insect Introduction Laboratory, I[BIII, ARS, USDA.

In 1981 a 0.2 ha plot of soybeans replaced the 0.2 ha sweet corn plot. The
3 ha field was again planted to field corn. I continued the survey by con-
centrating 45 egg cloths in and around the soybean plot.

The description of the Trichogramma species (except color characters)
is based on 15th-generation adults from a culture started in 1980 and 6th-
generation adults from a culture started in 1981. Both cultures were main-
tained at 20-25°C. The adults were preserved on glass slides in Hoyer’s
medium. The terminology used is that of Pinto et al. (1978). Quantitative
data listed in the description are those of the holotype (male) or allotype
followed (in brackets) by the mean, standard error, and range of that same
character measured on 30 additional specimens. (All characters are not from
the same 30 specimens since all characters are not clearly visible on all
specimens.) Description of color is based on freshly killed 15th-generation
adults from the 1980 culture. Since rearing temperature has been shown to
affect color (Kwangtung Collaborative Research Group of Biological Con-

18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

trol of Rice Pests, 1974), these insects were reared for at least two gener-
ations at 27°C, 70-80% RH, and constant light.

RESULTS AND DISCUSSION

A total of 1931 eggs on 79 egg cloths was parasitized by Trichogramma
in 1980. Dates of parasitization ranged from 12 May (first sample) to 21
October. Six species of Trichogramma were reared from these eggs. These
species were, in descending order of abundance, 7. minutum Riley (36), T.
pretiosum (Riley) (31), 7. exiguum Pinto and Platner (16), 7. parkeri Na-
garkatti (3) and T. marylandense, n. sp. (3), and T. retorridum (Girault) (1).
Figure | shows the number of cloths with eggs parasitized by each species
on each sampling date. Unfortunately, parasitism of eggs on a single cloth
by more than one female cannot be detected unless the females are of dif-
ferent species, since their progeny are otherwise indistinguishable. Trich-
ogramma minutum, T. pretiosum, and T. exiguum were collected much
more frequently than were the other three species and were collected
throughout the entire season, whereas the latter three species were collected
only briefly early in the season. The one exception was the collection of T.
marylandense in September, when parasitism was at a marked peak. Dif-
ferent eggs on the same cloth were parasitized by two different species of
Trichogramma on 11 occasions (T. minutum/T. pretiosum, 5; T. minutum/
T. exiguum, 4; T. pretiosum/T. exiguum, 1; T. pretiosum/T. marylandense,
1). On two occasions individuals of two species emerged from the same egg
(1. minutum ¢ and T. exiguum 3; T. minutum &@ and T. pretiosum @).

The 1981 survey is not yet completed, however specimens of T. mary-
landense collected in 1981 are included in the material examined for the
description. Trichogramma marylandense was collected on 13 July, 20 July,
and 17 August 1981, and sampling is continuing.

One of the objectives of the 1980 study was to determine the relative
abundance of the Trichogramma species in the survey area. However, the
number of females of each species sampled may not reflect their abundance.
Differences in host acceptability and microhabitat preference could affect
sample size. The discovery of six sympatric Trichogramma species all at-
tacking the same host (in this case an artificially provided host) should serve
to emphasize the need for basic ecological studies of Trichogramma species.
An understanding of such specific characteristics as habitat and host pref-
erence is essential to biological control programs involving species of this
genus.

Trichogramma marylandense Thorpe, New Species
Figs. 2-4

Type-data.—Holotype adult ¢ on slide with two other specimens under
separate coverslips; holotype nearest right margin of slide with right label

VOLUME 84, NUMBER 1 19

oy

I T. minutum 8 T. marylandense ]
7 ? T. pretiosum § T. parkeri
[ 4
$ T. exiguum 2 T. retorridum
a 6
o
Do
o
ie Sif |
aN
7)
wo
i 4} J
oO
Q
—
= |
= 3 i |
i S =
° | i
= 2 : 4
4 :
z =
Tr 2 . Ne LE . 4
He Nees Ne : RN
ES Nese Ne H ES
: N NN BUEN Ae EE R
fe) Nes. = ES, EL RS ENED

12 30 9 20 1 11 21 1 hia 22 2 12 23 4 14 24 qT
May June July August September October
Collection date

Fig. 1. Number of cloths with Heliothis virescens eggs parasitized by Trichogramma spp.
on each collection date, Beltsville, Maryland, 1980.

“Trichogramma marylandense Thorpe, Beltsville, Md. 12-4/l, mounted 31
XII 80, F15 lab culture, ex. Heliothis virescens egg cloth, coll. 26 VI 80, K.
Thorpe, Hoyers.’’ Allotype adult 2 on slide with two other specimens under
separate coverslips, same label; allotype center specimen. There are 145
paratypes (80 3, 65 2), 94 of which (50 6, 44 2) are on 32 slides all with
same data as holotype. Remaining 51 paratypes (30 ¢, 21 2) are on 17 slides
with right label ‘‘Trichogramma marylandense Thorpe, Beltsville, Md. 20-
3-2, mounted | IX 81, F6 lab culture, ex. Heliothis virescens egg cloth, coll.
13 VII 81, K. Thorpe, Hoyers.’’ Holotype (USNM type no. 100087) and
allotype are deposited in the collection of the National Museum of Natural
History, Washington, D.C.

Holotype male.—Eye and ocelli red; antenna and head yellow; thorax
yellow with black pronotum and mesoscutum; legs yellow except meso- and
metacoxae, metafemur, and 3rd tarsal segments black; wings clear with
smokey patch extending from base to even with distal end of stigma; Ist
abdominal segment yellow, remaining segments black.

Antenna (Fig. 2) with flagellum (excluding annellus) relatively long and
curved, 0.16 as wide as long (0.032 mm x 0.199 mm) [0.17 + 0.002; 0.15—
0.19], 1.21 as long as metatibia (0.165 mm) [1.20 + 0.005; 1.15-1.27], with
2 distinct constrictions, flagellar setae moderately long, tapering gradually

20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2

.o mm
Figs. 2-4. Trichogramma marylandense, holotype male. 2, Antenna. 3, Genital capsule. 4,
Aedeagus.

from base, 46 in number [46.6 + 1.63; 41-55], longest 2.46 as long (0.078
mm) as maximum width of flagellum [2.37 + 0.03; 2.07-2.75].

Forewing with vein tracts well defined, area between 4th and 5th vein
tracts with 34 setae [28.5 + 0.81; 22-38], longest seta on postapical wing
margin 0.12 as long as maximum wing width (0.032 mm and 0.267 mm)
[0.13 + 0.003; 0.10-0.18). Hindwing with posterior vein tract extending 4
to %x the distance of middle tract; anterior tract of 1-4 setae extending less
than 3 distance of middle tract.

Mesoscutellum with anterior setae 0.25 x as long as posterior setae (0.010
mm and (0.039 mm) [0.28 + 0.012; 0.19-0.50].

Genital capsule (Fig. 3) 0.37 as wide as long (0.036 mm x 0.099 mm)
[0.37 + 0.003; 0.33-0.39]; dorsal expansion of gonobase (DEG) shallowly
constricted at base, sides concave, width at base (0.023 mm) 0.64 the
width of genital capsule [0.71 + 0.016; 0.53-0.93]; DEG (0.102 mm long)
extending 1.02 the length of genital capsule [0.97 + 0.006; 0.92-1.03];
chelate structures (CS) (0.095 mm long) extending 0.95 the length of genital
capsule [0.91 + 0.008; 0.82—0.97]; median ventral projection (MVP) (0.090
mm long) long and slender, sharply pointed apically, extending 0.90 the
length of genital capsule [0.89 + 0.004; 0.83-0.92]; aedeagus (Fig. 4) (0.112
mm long) 0.68 as long as metatibia [0.66 + 0.006; 0.58—0.73], 1.12 as

VOLUME 84, NUMBER 1 21

“

long as genital capsule [1.12 + 0.004; 1.08-1.16]; apodemes (0.053 mm long)
comprising 0.48 length of aedeagus [0.50 + 0.009; 0.43-0.61].

Allotype female.—Color as in male except antenna and head yellow shad-
ed with black; mesoscutum yellow.

Forewing area between 4th and Sth vein tracts with 29 setae [28.2 + 0.78;
19-37]; longest seta on postapical wing margin 0.12 as long as maximum
wing width (0.032 mm and 0.267 mm) [0.13 + 0.002; 0.11-0.16].

Ovipositor 0.88 as long as metatibia (0.153 mm and 0.175 mm) [0.88 +
0.008; 0.80—0.97].

Remarks.—Males of 7. marylandense differ from males of most known
North American species of Trichogramma by having a DEG that is greater
than 0.90 the length of the genital capsule. The only other North American
species similar to T. marylandense in this respect are T. brevicapillum Pinto
and Platner, 7. fasciatum (Perkins), T. julianoi Platner and Oatman, T.
maltbyi Nagaraja and Nagarkatti, and 7. retorridum (Girault). The flagellar
setae of T. brevicapillum males are much shorter and stouter than those of
T. marylandense (1.17 longer than the flagellar width in T. brevicapillum
(Pinto et al., 1978) compared to 2.37x longer in 7. marylandense). The
DEG of T. marylandense males is narrower at its apex than the distance
across the gonostyli, which distinguishes it from 7. fasciatum males in
which the DEG is broader at its apex than the distance across the gonostyli
(Pinto et al., 1978). The genital capsule of 7. julianoi is very similar to that
of T. marylandense but can be distinguished by the length of the apodemes
of the aedeagus (0.80 the length of the aedeagus (Platner and Oatman,
1981) compared to 0.50 the length of the aedeagus in 7. marylandense).
Trichogramma maltbyi males differ from T. marylandense males by having
the DEG narrower at its base (ca. % genital capsule width in 7. maltbyi
(Nagaraja and Nagarkatti, 1973) compared with 0.71 in T. marylandense)
and by having longer antennal setae (nearly 4x the flagellar width (Nagaraja
and Nagarkatti, 1973) compared with 2.37 in 7. marylandense). Tricho-
gramma retorridum males can be separated from 7. marylandense males by
their very short, suboval flagellum (0.43 x as wide as long (Pinto et al., 1978)
compared to 0.17 as wide as long in 7. marylandense). The male genital
capsule of 7. marylandense is very similar to that of the Indian species 7.
achaeae Nagaraja and Nagarkatti (1969). However, the latter species has
a minute and inconspicuous MVP, whereas the MVP in 7. marylandense
is long and slender and attains 0.89 the length of the genital capsule. Also,
in T. achaeae the aedeagus and apodemes together attain less than the
length of the genital capsule while in 7. marylandense they attain 1.12 its
length.

Hosts and distribution.—Original specimens of 7. marylandense were
reared from Heliothis virescens eggs artifically placed on the edge of a corn
field in Beltsville, Maryland on 23 June, 26 June, and 12 September 1980,

22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

and from eggs placed in and near a soybean field at the same location on 13
July, 20 July, and 17 August 1981. At present these are the only known
collections of this species. Natural hosts are unknown.

ACKNOWLEDGMENTS

I thank Eric Grissell, Systematic Entomology Laboratory, HBIII, USDA,
and Akey C. F. Hung and David L. Vincent, Beneficial Insect Introduction
Laboratory, I[BIII, USDA, for their critical review of the manuscript.

LITERATURE CITED

Ables, J. R., S. L. Jones, R. K. Morrison, V. S. House, D. L. Bull, L. F. Bouse, and J. B.
Carlton. 1979. New developments in the use of Trichogramma to control lepidopteran
pests of cotton. Proc. Beltwide Cotton Prod. Res. Conf. 3 pp.

Kwangtung Collaborative Research Group of Biological Control of Rice Pests, Kwangtung
Province. 1974. The control of rice leaf roller, Cnaphalocrocis medinalis Guenee by
trichogrammatid egg parasites. (In Chinese with English summary.) Acta Entomol. Sin-
ica 7(3): 269-280.

Nagaraja, H. and S. Nagarkatti. 1969. Three new species of Trichogramma from India. En-

tomophaga 14(4): 393-400.

1973. A key to some new world species of Trichogramma (Hymenoptera: Tricho-
grammatidae), with descriptions of four new species. Proc. Entomol. Soc. Wash. 75(3):
288-297.

Nagarkatti, S. and H. Nagaraja. 1971. Redescriptions of some known species of Trichogram-
ma (Hymenoptera: Trichogrammatidae) showing the importance of the male genitalia
as a diagnostic character. Bull. Entomol. Res. 61: 13-31.

Oatman, E. R. and G. R. Platner. 1978. Effect of mass releases of Trichogramma pretiosum
against lepidopterous pests on processing tomatoes in Southern California, with notes
on host egg population trends. J. Econ. Entomol. 71(6): 896-900.

Parker, F. D. 1971. Management of pest populations by manipulating densities of both hosts
and parasites through periodic releases, pp. 365-376. Jn Huffaker, C. B., ed., Biological
Control. Plenum Press, N.Y. 511 pp.

Pinto, J. D., G. R. Platner, and E. R. Oatman. 1978. Clarification of the identity of several
common species of North American Trichogramma (Hymenoptera: Trichogrammati-
dae). Ann. Entomol. Soc. Am. 71(2): 169-180.

Platner, G. R. and E. R. Oatman. 1981. Description of a new species of Trichogramma
(Hymenoptera: Trichogrammatidae) from New York. Proc. Entomol. Soc. Wash. 83(1):
164-167.

Ridgway, R. L., F. R. Ables, C. Goodpasture, and A. W. Hartstack. 1981. Trichogramma
and its utilization for crop protection in the U.S.A., pp. 41-48. Jn Coulson, J. R., ed.,
Proceedings of the Joint American-Soviet Conference on the Use of Beneficial Organ-
isms in the Control of Crop Pests. Entomol. Soc. Amer., College Park, Md. 62 pp.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 23-37

FIVE NEW SPECIES OF PHERBELLIA ROBINEAU-DESVOIDY,
SUBGENUS OXYTAENIA SACK, FROM NORTH AMERICA
(DIPTERA: SCIOMYZIDAE)

R. E. ORTH

Staff Research Associate, Department of Entomology, Division of Bio-
logical Control, University of California, Riverside, California 92521.

Abstract.—Pherbellia borea, P. californica, P. marthae, P. paludum, and
P. ursilacus are described as new species from North America. Illustrations
and photographs of the copulatory apparatus and geographic distribution
are given for the new species as well as for P. bryanti Steyskal and P.
propages Steyskal. All are members of the P. propages group.

The genus Pherbellia Robineau-Desvoidy is known from all the zoogeo-
graphic realms of the world. Within the family Sciomyzidae Pherbellia con-
stitutes by far the largest genus. Prior to this study there were 32 described
species of Pherbellia from North America north of Mexico. Some of the
species which occur in the Nearctic also occur in the Neotropic and Pa-
laearctic regions.

It seems advisable to recognize Pherbellia as a single genus as did Steys-
kal (1961, 1966) until a more comprehensive taxonomic study can be made.
The genus has been broken into several subgenera. This paper is concerned
only with North American members of the subgenus Oxytaenia Sack.

The type-species of the subgenus Oxytaenia is Pherbellia brunnipes
(Meigen). It was recorded by Melander (1920) and Steyskal (1965) as Hol-
arctic, being found in Wyoming and Idaho, USA, as well as Europe. How-
ever, Steyskal (1966) stated in regard to P. brunnipes **. . . my examination
of European material shows a species different from any I have seen from
this continent, including material from the Melander collection. Of the
American species, it is most closely related to P. bryanti and P. propages
....’ Close examination of illustrations by El’berg (1965) and Rozkosny
(1966) showing European terminalia further confirms that P. brunnipes is
Palaearctic. On the basis of external characters, close scrutiny of a female
specimen from Melander’s collection from Wyoming labeled P. brunnipes
reveals it to be P. prefixa Steyskal.

Presently from North America there are nine species referred to the sub-

24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

genus Oxytaenia, which is now comprised of Pherbellia beatricis, P. bryan-
ti, P. prefixa, P. propages plus the five new species which are herein de-
scribed. All are quite small, rather inconspicuous, dull colored flies.

The new species are separated from Pherbellia propages and P. bryanti.
These new species along with P. propages and P. bryanti are here referred
to as the P. propages group. The male postabdomen of Pherbellia bryanti,
P. propages, and the new species provides the most definitive means of
separation. For identification the postabdomen must first be severed from
the specimen and placed in a 10% solution of potassium hydroxide until the
viscera and muscular tissues have been dissolved. The postabdomen is then
flushed with distilled water and examined under the microscope in a droplet
of glycerine. Staining the prepared section with acid fuchsin will further
intensify subtle membranous structures, thereby facilitating determinations.

The postabdomen of North American Oxytaenia may still be typified as
stated by Steyskal (1966): ‘‘surstyli of anterior and posterior pairs close
together, the posterior pair broad basally, usually with a narrow apical point
more or less directed posteriad, the anterior pair small and lying backward
against the posterior pair; hypandrium with anterior margin transverse, lat-
eral processes consisting of broad anterior lamina, and more or less blade-
like posterior sclerite bearing variously developed apical prongs; phallapo-
deme deeply cleft, Y-shaped, arms bearing clamshell-like pregonite;
aedeagus broad; postgonites massive, black, usually with small but deep
apical emargination giving them the appearance of a crab’s claw; ejaculatory
apodeme narrow-headed, relatively small and short, but with large vesicle
(seminal pump), the vas deferens short and of small diameter.”

The Pherbellia propages group is here considered to be monophyletic.
The adults show scarcely any external differences. However, dissection of
the male terminalia reveals distinct differences in the aedeagi and the pos-
terior process of the hypandria. Lesser differences are seen in the posterior
and anterior surstylus as well as the ejaculatory apodeme and seminal pump.
Some species are allopatric such as P. californica and P. ursilacus as shown
on the locality map (Fig. 34). Four of the species are sympatric at Aklavik,
Northwest Territories (Figs. 33, 34).

The holotypes of both Pherbellia propages and P. bryanti have been
examined and are as illustrated in this paper. Examination of paratypes of
the above species reveals that several species of the P. propages group are
represented. Therefore, the locality maps as they appear in this paper in-
dicate the known distribution for P. propages and P. bryanti, sensu stricto.

The groundwork and stimulus for this study was provided by Steyskal
(1966). In his description of Pherbellia propages he observed and illustrated
differences in the posterior process of the hypandrium. He felt that the
differences might represent subspecies, however he made no mention at
that time of aedeagal differences.

Members of the Pherbellia propages group share the following charac-

VOLUME 84, NUMBER 1 25

ters: Frons with median stripe less than 2% as long as distance from ocellus
to frontal margin; anterior fronto-orbital bristle much shorter than posterior
fronto-orbital bristle; arista with short, fine, closely set hairs; mesopleuron
entirely bare; pteropleuron with all bristles of nearly equal length; sterno-
pleuron usually with 2 very strong bristles and several weak bristles along
the upper surface; wing not patterned, at most with anterior margin and
crossvein clouded; wing length of males, 2.3-3.4 mm; first vein not sur-
passing level of ta, either ending short of it or at the same level; halter pale
yellowish or whitish.

All possess a mesoscutal pattern of intradorsocentral and sublateral
brown stripes. The degree of distinctness seems dependent on the condition
of the insect at the time of capture. In general old battered specimens tend
to loose their distinct stripes. All specimens I have seen in the Pherbellia
ursilacus series show this character indistinctly.

Pherbellia californica Orth, New Species
Figs. 1, 2, 6, 17-20, 34

Holotype male.—Height of head *4 width. Medifacies yellowish to whitish
pruinose, facial grooves subshiny, parafacies and cheeks pruinose yellowish
to whitish respectively. Frons yellowish, slightly narrowed anteriorly. Mid-
frontal stripe extending 2 distance from anterior ocellus to anterior margin
of frons. Ocellar triangle and orbital plates with greyish pruinosity. Orbital
plates tapered anteriorly, extending beyond midfrontal stripe. Orbito-anten-
nal spot lacking; narrow strip of whitish pruinosity along upper orbital mar-
gin. Two pairs of fronto-orbital bristles, anterior pair /% as long; ocellars,
postocellars, and inner and outer verticals well developed. Occiput greyish
pruinose. Short black setae on lower % of cheeks and parafacies on anterior
'Y of frons, between ocellar and postocellar bristles, along outer parts of
orbital plates, and in midcervial patch. Lateral occipital margins with strong-
er setae and bristles. Antennae testaceous, segment 3 elongate oval. Arista
blackish with short hairs. Palpi yellowish, labium and labella yellowish
brown.

Thorax dorsally grey, pruinose, with indistinct brownish longitudinal
stripes.

Mesopleuron grey, pruinose throughout, posterior % brownish. Remain-
der of thoracic surfaces greyish pruinose with some tendency toward brown
or brownish yellow. Mesopleuron bare; pteropleuron with a cluster of bris-
tles of nearly equal size. Sternopleuron with fine short setae over most of
surface, 2 well developed bristles dorsally and well developed bristles ven-
trally. Prosternum bare.

Coxae yellowish white, pruinose. Forefemur and tibia dark brownish in-
fumated, tarsal segments light brown infumated. Mid- and hindlegs entirely
testaceous.

Wing length 3.4 mm. Membrane greyish-yellow hyaline; costal margin

26 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Pherbellia californica, paratype male. Point-mounted. Photo by M. E. Badgley,
University of California, Riverside.

and wing veins testaceous; crossveins brownish, slightly infuscated. No
stump veins; anterior cross vein slightly oblique, first vein not surpassing
the level of anterior cross vein; anal vein reaching wing margin. Halter,
Squama, and squamal ciliae yellowish.

Abdominal segments testaceous, slightly infumated dorsally; andrium tes-
taceous; terminalia as in Fig. 2. Copulatory apparatus as in Figs. 6, 17-20.

Allotype female.—Similar to holotype except for reproductive structures.
Wing length 3.3 mm.

Holotype.—d, California, Mendocino Co., 2 mi N of Willits, 24 April
1967, elevation 1330 feet. T. W. Fisher—R. E. Orth, field notes accession
no. AS-578. National Museum of Natural History.

Allotype.—@, same data as holotype. Deposited with holotype.

Paratypes.—Same locality and collector data as holotype. 12 June 1966,
AS-482 (8 2, 26 5); 24 May 1967, AS-578 (13 2, 36 3); 23 April 1968, AS-
669 (3 2, 3 3); 24 April 1968, AS-671 (2 3d). At Agriculture Canada, Cali-
fornia Academy of Sciences, Cornell University, University of California
at Riverside, and the National Museum of Natural History.

VOLUME 84, NUMBER 1 27

0.2mm

Figs. 2, 3. 2, Pherbellia californica, paratype male; 2 mi N of Willits, Mendocino Co.,
California, 24 May 1967, T. W. Fisher-R. E. Orth; terminalia, sinistral view, inverted; apae =
apical process of aedeagus; as = anterior surstylus; ce = cerci; pph = posterior process of
hypandrium. 3, P. borea, paratype male; Hay River, N.W.T., 10 Sept. 1932, O. Bryant, lot
no. 352; terminalia, sinistral view, inverted; ae = aedeagus; alh = anterior lamina of hypan-
drium; ap = apical prong; ep = epandrium; ps = posterior surstylus.

Other specimens.—In addition to the above I have seen material from the
following localities: California: Humboldt Co., 2 mi S of Orick; Marin Co.,
2 mi NW of Bolinas; .5 mi S of Tomales; Mendocino Co., 7 mi N of Hopland;
Ukiah; Plumas Co., 1 mi E of Chester; .5 mi S of Crescent Mills; Rock
Creek at Hwy. 36; Shasta Co., 5.25 mi NW of Anderson; Siskiyou Co., 3.5
mi S of Fort Jones, Scott Valley. Oregon: Benton Co., Parker Creek,
Mary’s Peak; Hood River Co., .5 mi S of Sherwood Campground, Mt. Hood
National Forest; Marion Co., Breitenbush; Wasco Co., Hwy. 26, | mi S of
Hwy. 216 Jct. Washington: Lewis Co., Adna.

Variation.—Wing length 2.6-3.4 mm in males, 2.8-4.0 mm in females.

Discussion.—This species is known only from California, Oregon and
Washington. It overlaps no other species in the Pherbellia propages group.
It is the only species in the group which shows considerable variation in the
aedeagus. In specimens collected in the general proximity of the type-lo-
cality the apical process of the aedeagus bends back on its self while in
more northern specimens it is opened out more (see Figs. 17, 18). Deviation
of the aedeagus and subtle differences in the posterior process of the hy-
pandrium in material from localities away from the type-locality may rep-
resent a subspecies.

28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pherbellia borea Orth, New Species
Figs. 3, 4, 11=13533

Holotype male.—Similar to Pherbellia californica except as follows. Mid-
and hindfemora infuscated, brown, hindfemur darker than midfemur; tibiae
and tarsi brown, slightly infumated. Abdominal segments brownish grey.
Terminalia as in Fig. 3. Copulatory apparatus as in Figs. 4, 11-13. Wing
length 2.9 mm.

Female.—Not known.

Holotype.—d, Northwest Territories, Aklavik, 25 July 1931, O. Bryant,
lot no. 300. National Museum of Natural History.

Paratypes.—Northwest Territories: Aklavik, 5 August 1930, lot no. 110,
18 Sept. 1930, lot no. 153, 10 Aug. 1931, lot 289, O. Bryant G d)isinay
River, 10 Sept. 1932, lot no. 352, O. Bryant (1 6); Hyndman, 68°5’N,
131°03’W, 30 July 1969, 1 Aug. 1969, G. E. Shewell (3 6, 2 d) respectively.
Wing length 2.6-3.0 mm. At Agriculture Canada, California Academy of
Sciences, University of California at Riverside, and the National Museum
of Natural History.

Discussion.—Distribution of this species is the most northern of the Pher-
bellia propages group. The southern-most collection site is Hay River,
Northwest Territories approximately 61° north latitude. This species name
is from the Latin adjective borea—northern.

Pherbellia marthae Orth, New Species
Figs. 7, 21-23, 33

Holotype male.—Similar to Pherbellia californica except as follows. Mid-
and hindlegs brown, slightly infuscated, hindfemur darker than midfemur.
Abdominal segments brownish grey. Copulatory apparatus as in Figs. 7,
21-23. Wing length 3.2 mm.

Female.—Not known.

Holotype.—d , Churchill, Manitoba, 2-9 Aug. 1937, D. G. Denning. Na-
tional Museum of Natural History.

Paratypes.—Alberta: Cascade Trail, Banff, 10 July 1968, elevation 5000
feet, H. J. Teskey (1 6); Cooking Lake, 20 June 1937, 25 July 1937, F. O.
Morrison (2 3); 1.5 mi W of Lacombe, 23 May 1962, C. O. Berg (2 3);
Slave Lake, 6 June 1966, G. E. Shewell (1 3). British Columbia: Ketchum
Lake, 58°22'N, 131°45’W, 26 Aug. 1960, W. W. Moss (1 3); Spectacle Lake,
Oliver, 10 June 1959, L. A. Kelton (1 3). Manitoba: Churchill, 2-9 Aug.
1937, D. G. Denning (1 ¢); Herchmer, 10 Aug. 1937, D. G. Denning (1
3); The Pas, 28 May 1930, O. Bryant, lot no. 6 (1 6); Treesbank, 17 Oct.
1915, J. M. Aldrich (1 6). Northwest Territories: Aklavik, 18 June 1931, 22
June: 1931; 24 June 1931, 'O. Bryant; lot nos: 234, 238, 241 :(h 3d 1G
3); Fort Franklin, Great Bear Lake, 22 June 1969, G. E. Shewell (3 3); Lac
Maunoir, N shore, 15 July 1969, 17 July 1969, G. E. Shewell (1 3, 11 3)

VOLUME 84, NUMBER 1 29

10
Figs. 4-10. Aedeagi, sinistral profiles. 4, Pherbellia borea, paratype male; Aklavik,
N.W.T., 18 Sept. 1930, O. Bryant, lot no. 153. 5, P. bryanti; Yellowstone National Park,

+

Gibbon Falls, Wyoming, 27 July 1956, A. L. Melander. 6, P. californica, paratype male; 2 mi
N of Willits, Mendocino Co., California, 24 May 1967, T. W. Fisher-R. E. Orth. 7, P. marthae,
paratype male; Churchill, Manitoba, 2-7 Aug. 1937, D. G. Denning. 8, P. paludum, paratype
male: Gander, Newfoundland, 17 June 1961, C. P. Alexander. 9, P. propages; Long Lake,
Burleigh Co., North Dakota, 4 June 1968, W. W. Wirth. 10, P. ursilacus, paratype male; Bear
Lake, Bear Lake Co., Idaho, 19 July 1959, B. A. Foote.

30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1a
2 13 i 16
0.1mm
Us) 20
18
Figs. 11-20. 11-13, Pherbellia borea, paratype male; Aklavik, N.W.T., 18 Sept. 1930, O.

Bryant, lot no. 153. 14-16, P. bryanti; Yellowstone National Park, Gibbon Falls, Wyoming,
27 July 1956, A. L. Melander. 17-20, P. californica: 17, 19, 20, Paratype male, 2 mi N of
Willits, Mendocino Co., California, 24 May 1967, T. W. Fisher-R. E. Orth; 18, Adna, Wash-

VOLUME 84, NUMBER 1 3]

respectively; Norman Wells, 25 June 1969, G. E. Shewell (3 36). Saskatch-
ewan: Cantyre, 53°23'N, 101°50'W, 12 Sept. 1959, J. R. Vockeroth (4 3);
Quill Lake, 20 May 1968, H. J. Teskey (1 6); Uranium City, 18, 19, 21 June
1962, J. G. Chillcott (1 6, 16,14); Yorkton, 16 Sept. 1959, J. R. Vockeroth
(1 6). Yukon: 58 mi E of Dawson, Gravel Lake, 12 Aug. 1962, P. J. Skitsko
(1 3); La Force Lake, 9 July 1960, J. E. H. Martin (1 ¢); Takhini Hot
Springs, 16 Aug. 1962, R. E. Leech (1 6). Alaska: Fairbanks, 7 June 1948,
R. Sailer (2 5); Goldstream River Valley, 8% mi N of Fairbanks, 29 July
1971, B. A. Foote (1 6); Wonder Lake, McKinley National Park, 25 July
1965, D. Chant (1 6). Montana: 2 mi E of Babb, 22 July 1967, B. A. Foote
(2 3d); 7 mi W of Eureka, 15 July 1965, B. A. Foote (1 6). North Dakota:
McHenry Co., 4 mi N of Upham, 5 June 1969, W. W. Wirth (1 3); Mountrail
Co., Powers Lake, 8 June 1969, W. W. Wirth (2 3). Wing length 2.3-3.2
mm. At Agriculture Canada, Cornell University, University of California at
Riverside, and the National Museum of Natural History.
Discussion.—This species was the most common of the P. propages
group found in collections. It, along with three other species of the group,
has been collected at Aklavik, Northwest Territories. It is a northwestern
and mid-continent species. This species is named for my wife, Martha.

Pherbellia paludum Orth, New Species
Figs. 8, 24-26, 33

Holotype male.—Similar to Pherbellia californica except as follows. Mid-
and hindfemora and tibiae infumated, brown, hindfemur darker than mid-
femur; mid- and hindtarsi light brownish. Abdominal segments brownish
grey. Copulatory apparatus as in Figs. 8, 24-26. Wing length 2.65 mm.

Female.—Not known.

Holotype.—dé, Newfoundland, Woody Point Rd., 10 June 1961, C. P.
Alexander. National Museum of Natural History.

Paratypes.—Alberta: 40 mi W of Edmonton (Wabamun Lake), 23 May
1962. ,€. ©; Bere (vd)3) 1.5. .mi W of .Lacombe,,23 May 1962,.C. O. Bere
(1 36). Newfoundland: Gander, 17 June 1961, C. P. Alexander (1 ¢); Woody
Point Rd., 10 June 1961, C. P. Alexander (1 6). Northwest Territories:
Aklavik, 5 Aug. 1930, lot no. 110, O. Bryant (2 ¢); Aklavik, 18 July 1932,
lot no. 305, O. Bryant (1 ¢). Ontario: Midland, 12 May 1959, J. G. Chillcott
(1 3); 5 mi S of Severn Falls, 4 May 1959, J. G. Chillcott (2 ¢). Alaska:
Fort Richardson, Eagle River Flats, 25 May 1948, E. Lepage (1 ¢); Matan-

—

ington, 10 July 1917, A. L. Melander. 11, 14, 17, 18, Aedeagus sinistral profile. 12, 15, 19,
Posterior process of hypandrium, side view. 13, 16, 20, Posterior process of hypandrium,
posterior oblique view.

32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

28 2Q 31 32

Figs. 21-32. 21-23, Pherbellia marthae, paratype male; Churchill, Manitoba, 2-7 Aug.
1937, D. G. Denning. 24-26, P. paludum, paratype male; Gander, Newfoundland, 17 June
1961, C. P. Alexander. 27-29, P. propages; Long Lake, Burleigh Co., North Dakota, 4 June

VOLUME 84, NUMBER 1 33

uska, 14 Aug. 1952, C. O. Berg (1 6). Idaho: Benewah Co., 1 mi N of
Plummer, 26 May 1959, B. A. Foote (1 6). Michigan: Midland Co., 19 May
1939, R. R. Dreisbach (1 ¢); Osecola, 1 June 1947, R. R. Dreisbach (1
3). Wyoming: Biscuit Basin, Yellowstone National Park, 2 Aug. 1934, A.
L. Melander (1 3). At Agriculture Canada, California Academy of Sciences,
Cornell University, University of California at Riverside, and the National
Museum of Natural History.

Discussion.—Pherbellia paludum has the most widespread distribution of
any species of the P. propages group. However in collections it is relatively
rare. This species is known transcontinentally north of 44°N latitude. This
species name is the genitive pleural of the Latin palus, meaning ‘‘of
marshes.”

Pherbellia ursilacus Orth, New Species
Figs. 10, 30-32, 34

Holotype male.—Similar to Pherbellia californica except as follows. This
specimen, along with the paratypes, shows a tendency toward being greasy.
Coloration tends toward yellowish brown in most areas. Thorax dorsally
cinereous, with only faint longitudinal darker stripes. Copulatory apparatus
as in Figs. 10, 30-32. Wing length 3.2 mm.

Female.—Not known.

Holotype.—d, Bear Lake, Bear Lake Co., Idaho, 19 July 1959, B. A.
Foote. National Museum of Natural History.

Paratypes.—Same data as holotype (18 ¢). Wing length 2.5—3.2 mm. At
Cornell University, University of California at Riverside, and the National
Museum of Natural History.

Discussion.—This species is known only from Bear Lake, Idaho. The
type-locality is described by Foote (1961). The collecting site was a marshy
area approximately four miles west of the town of Mud Lake. It is a flood
plain area poorly drained, with many extensive marshes bordering the Bear
River north of the lake. The collection locality was at the edge of one of
these marshes. Foote states ‘‘The marsh is completely unshaded and sup-
ports arich stand of hydrophilic vegetation composed primarily of numerous
species of sedges (mostly Carex spp.) and rushes. At the time of collection
in mid-July the water depth was less than one foot, and extensive areas
lacked standing water, although the soil was water-logged. Aquatic snails
are abundant and include such common genera as Lymnaea, Physa, Heli-

i
1969, W. W. Wirth. 30-32, P. ursilacus, paratype male; Bear Lake, Bear Lake Co., Idaho, 19
July 1959, B. A. Foote. 21, 24, 27, 30, Aedeagus, sinistral profile. 22, 25, 28, 31, Posterior
process of hypandrium, side view. 23, 26, 29, 32, Posterior process of hypandrium, posterior
oblique view.

3d eras ‘

é

ta
bt

SCALE
ox .
f GOODE BASE MAP SERIES
DEPARTMENT OF GEOGRAPHY
THE UNIVERSITY OF CHICAGO
LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION HENRY M LEPPARD, EDITOR

Fig. 33. Collection sites for Pherbellia borea, P. marthae, and P. paludum.

VOLUME 84, NUMBER 1 35

Taree eg hd

ot ae

i

\

P. bryanti

L eee es

By

P. ursilacus ¥.

P. propages

120 110 100 90

“Geo? tap” MILOMEERS GOODE BASE MAP SERIES
CRPANTAENT OF GEOSAADY
ee uneven

OF CHKAGO
LAMBERT AZIMUTHAL EQUAL-AREA PROJECTION VeNEY ® LEPPARD, EDITOR

Fig. 34. Collection sites for Pherbellia bryanti, P. californica, P. propages, and P. ursi-

36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

soma, and Gyraulis. Many snails are stranded on the wet soil as the water
level recedes during the summer months. A very interesting group of scio-
myzid species was taken including some forms not collected elsewhere in
the state (e.g., Pherbellia obtusa and Sepedon anchista).’’ Under distri-
bution Foote (1961) reports ‘‘Pherbellia obtusa (Fallén).—Recorded from
Bear Lake on July 19 [1959] when approximately 60 adults were taken during
a two-hour search of the marsh.”’ It has since been determined that P.
obscura is Palaearctic. According to Bratt et al. (1969) P. argyra, a holarctic
species, has been until recently misidentified as P. obtusa. | have seen
specimens from Cornell University which are P. argyra and bear the exact
above data. With the above information it appears that Foote’s P. obtusa
was comprised of both P. argyra and P. ursilacus.

This species most closely resembles Pherbellia marthae. This species
name is the latinization of the type-locality; Ursa—bear + lacus—lake; it
is either a noun in apposition or in the genitive case, both of which have
the same form.

Pherbellia bryanti Steyskal
Figs. 5, 14-16, 34

The holotype male was examined and agrees well with Figs. 5, 14-16.
Collection data for the holotype: Aklavik, Northwest Territories, 20 Aug.
1930, O. Bryant, National Museum of Natural History Type no. 67875. Wing
length 2.9 mm.

Other specimens.—In addition to the holotype I have seen specimens
from the following localities (Fig. 34): Alberta: Soda Lake. Northwest Ter-
ritories: Aklavik; Good Hope; Hay River. Saskatchewan: Parkside; Quill
Lake. Wyoming: Gibbon Falls, Yellowstone National Park.

Discussion.—This species is known from Aklavik, Northwest Territories
to Yellowstone National Park, Wyoming. Collection data has its distribution
restricted to a narrow band just east of the continental divide.

Pherbellia propages Steyskal
Figs. 9, 27-29, 34

The holotype male was examined and agrees well with Figs. 9, 27-29.
Collection data for holotype: Aweme, Manitoba, 9 Oct. 1915, N. Criddle,
National Museum of Natural History Type no. 67877. Wing length 2.7 mm.

Other specimens.—In addition to the holotype I have seen specimens
from the following localities (Fig. 34): Saskatchewan: Hudson Bay; Can-
tyre. Michigan: Midland Co. Minnesota: Anoka Co., Cedar Creek Bog;
Norman Co.; Ramsey Co., University Farm, St. Paul. North Dakota: Bur-
leigh Co., Long Lake; Leonard.

Discussion.—Collection data reveal this species to be more or less re-
stricted to north central United States and south central Canada. Its known
distribution overlaps in part with P. marthae and P. paludum.

VOLUME 84, NUMBER | 3)/)

ACKNOWLEDGMENTS

I express deep appreciation to the following individuals and institutions
for their assistance or loan of material: B. E. Cooper, J. E. H. Martin, and
R. V. Peterson, Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario; P. H. Arnaud, Jr., California Academy of Sciences, San
Francisco; C. O. Berg, Cornell University, Ithaca, New York; B. A. Foote,
Kent State University, Kent, Ohio; R. L. Westcott, Oregon State Depart-
ment of Agriculture, Salem; T. W. Fisher and J. C. Hall, University of
California, Riverside; L. Knutson and G. C. Steyskal, Insect Identification
and Beneficial Insect Introduction Institute, USDA, Beltsville, Maryland
and Systematic Entomology Laboratory, USDA, Washington, D.C., re-
spectively. I am particularly grateful to T. W. Fisher for his counsel and
encouragement over the years in the taxonomy of sciomyzid flies.

LITERATURE CITED

Bratt, A. D., L. V. Knutson, B. A. Foote, and C. O. Berg. 1969. Biology of Pherbellia
(Diptera: Sciomyzidae). Cornell Univ. Exp. Stn. Mem. 404, 247 pp.

El’berg, K. Y. 1965. New Palaeartic genera and species of flies of the family Sciomyzidae
(Diptera, Acalyptrata). Entomol. Obozr. 44(1): 189-198.

Foote, B. A. 1961. The marsh flies of Idaho and adjoining areas (Diptera, Sciomyzidae). Am.
Midl. Nat. 65(1): 144-167.

Melander, A. L. 1920. Review of the Nearctic Tetanoceridae. Ann. Entomol. Soc. Am. 13:
305-332, | plate.

Rozkosny, R. 1966. Ceskgslovenské druhy malakofagni éeledi Sciomyzidae (Diptera). Folia
Prirodovedecke Fak. J. E. Purkyne, Biologia 15: 1-111.

~ Steyskal, G. C. 1961. The North American Sciomyzidae related to Pherbellia fuscipes (Mac-

quart). Pap. Mich. Acad. Sci. Arts Lett. 46: 405-415.
—.. 1965. Sciomyzidae, pp. 685-695. Jn Stone et al., A Catalog of the Diptera of America
North of Mexico. U.S. Dep. Agric., Agric. Handb. 276, 1696 pp.

| ———. 1966. The Nearctic species of Pherbellia Robineau-Desvoidy, subgenus Oxytaenia

Sack (Diptera, Sciomyzidae). Pap. Mich. Acad. Sci. Arts Lett. 51: 31-38.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 38-66

SYNOPSIS OF THE GLYPHIPTERIGIDAE (LEPIDOPTERA:
COPROMORPHOIDEA) OF THE WORLD

JOHN B. HEPPNER

Department of Entomology, Smithsonian Institution, Washington, D.C.
20560.

Abstract.—A brief history and review of the current taxonomic status of
the family Glyphipterigidae is discussed for the world fauna. Included are
a world checklist of genera and species, notes on type-localities (including a
map and table of faunal distributions) for all 326 species, a terminal bibliog-
raphy to original descriptions of taxa, and an index to species. Carmentina
Meyrick is transferred from Yponomeutidae to Glyphipterigidae.

In preparing a revision of the North American Glyphipterigidae, as well
as a world catalog of genera formerly associated with the family (Heppner,
in press b), the following list of included species was completed. The Glyphip-
terigidae sensu Meyrick (1914c) involved one of the most mixed conglom-
erations of unrelated genera ever assembled as one family in the Lepidoptera
in the last 100 years. Representatives of 24 families as divergent and unre-
lated as Incurvariidae and Zygaenidae were included in Glyphipterigidae by
Meyrick, primarily on the basis of various similar characters and on similar
superficial appearance, especially the iridescently colored markings of the
forewings of many of these moths. These common “‘facies’’ undoubtedly
are the result of convergent evolution among unrelated taxa as a response
to behavioral patterns for diurnal flight activity, which is the normal activity
of the majority of species among these unrelated moths.

The Glyphipterigidae are characterized by adult characters of a tineoid
abdominal articulation, naked haustellum, head smooth-scaled, labial palpus
upcurved and usually with ultimate segment longest, maxillary palpus 24
segmented, ocellus present and large, forewings having a pterostigma and
usually with a developed chorda, and male genitalia without a socius, uncus,
or gnathos. The larvae are mostly borers; setal patterns include 2 L-group
prothoracic setae; prolegs are vestigial; and spiracles are often projected on
conelike protrusions. The pupa is unspined and the spiracles are on projec-
tions like in the larva. Some species, notably in Ussara, produce a netlike
pupal case; most other genera pupate in the larval host branch or stem.

VOLUME 84, NUMBER 1 39

As was pointed out in an earlier paper (Heppner, 1977), the main body
of the Glyphipterigidae sensu Meyrick, comprises four major distinct fam-
ilies: Glyphipterigidae sensu stricto (Copromorphoidea), Immidae (Immoi-
dea), Brachodidae (Sesioidea), and Choreutidae (Sesioidea). Another larger
group of genera are now placed in Tortricidae as the tribe Hilarographini
(Chlidanotinae). These five groups comprise in total over 1200 species. The
remaining genera and species not now in these families are distributed
among another 20 families: Incurvariidae, Tineidae, Psychidae, Gracillari-
idae, Oecophoridae, Elachistidae, Agonoxenidae, Gelechiidae, Copro-
morphidae, Epermeniidae, Plutellidae, Yponomeutidae, Douglasiidae, Acro-
lepiidae, Heliodinidae, Pyralidae, Zygaenidae, Limacodidae, Geometridae,
and Noctuidae (Heppner, in press b).

All genera previously associated with Glyphipterigidae are listed and as-
signed to appropriate families in a catalog of these generic names (Heppner,
in press b). Achecklist of Immidae has been completed (Heppner, in press a).
Checklists of Brachodidae and Choreutidae are available in a review of the
Sesioidea (Heppner and Duckworth, 1981). The Hilarographini (Tortricidae)
will be reviewed in a future paper.

The following checklist of the genera and species remainng in Glyphip-
terigidae is presented here to complete the synopses of the majority of the
species formerly assigned to this family sensu lato. The synopsis of genera
below gives an overview of the family and the generic distribution of the
326 species remaining in Glyphipterigidae. Table 1 shows these species as
they are distributed among the various faunal regions as based upon their
type-localities. The map (Fig. 1) illustrates the world distribution of the
type-localities of Glyphipterigidae species (most species are known only
from the type-locality). The surprising aspect seen on the map is the lack
of described species of Glyphipterigidae from such areas as New Guinea,
Sumatra, the Philippines, and temperate South America (Chile and Argen-
tina), areas which should have some species, indeed in most cases large
faunas, for this family. Future collections from such areas undoubtedly will
greatly increase the current size of the family; this can be partially substan-
tiated merely with the North American fauna, where the present total of 15
species rises to 38 in the revision shortly to be in press. Such areas as Japan,
New Zealand, western Europe, and parts of Australia, as well as North
America, probably are now fairly well known for Glyphipterigidae.

The world checklist has the type-species noted for each genus, the page
of original description following the year of publication of each name, with
the general type-locality for each species noted at the right margin. Local-
ities in brackets are of subsequent determination. Letters after Australia
refer to New South Wales (NSW), Queensland (Q), South Australia (SA),
Tasmania (T), Victoria (V), and Western Australia (WA). Other abbrevia-
tions are: ab. = aberration; emend. = emendation; ICZN = International

40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

SCALE ON EQUATOR

000
=

Fig. 1. Distribution map of the family Glyphipterigidae by type-locality of species in the

VOLUME 84, NUMBER 1 4]

world (western and eastern hemispheres).

42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Faunal distribution of Glyphipterigidae species. N = Nearctic; NL = Neotropical:
P = Palearctic; E = Ethiopian; O = Oriental; A = Australian; OC = Oceana.

Genus N NL P E O A Oc Total

Chrysocentris 7 |
Irinympha l
Ernolytis 1
Carmentina | 3
Cotaena 2
Myrsila |
Lepidotarphius l
Tetracmanthes 1
Phalerarcha 2
Cronicombra 6

2

2

tN

Taeniostolella

Machlotica

Abrenthia |
Neomachlotica |
Trapeziophora

Ussara 9
Electrographa 1
Rhabdocrates |

Apistomorpha

Phryganostola

Pantosperma

Circica

Glyphipterix 7 4] 59 20 40 85 1 2S.
Diploschizia 6 3

5 73 60 31 48 97 2 326

— WW

th
ho
_

Noe De
OW NK DK KK WRK PRK NNDAN KR KK NDA |= — CO

Faunal totals I

Commission on Zoological Nomenclature; missp. = misspelling; nom.
nud. = nomen nudum; nom. dub. = nomen dubium; nom. oblit. = nomen
oblitum; preocc. = preoccupied; var. = variety.

Generic Synopsis of Glyphipterigidae

Genus Number of species

(oe)

Chrysocentris Meyrick, 1914
Irinympha Meyrick, 1932
Ernolytis Meyrick, 1922
Carmentina Meyrick, 1930
Cotaena Walker, [1865]
Myrsila Boisduval, [1875]
Lepidotarphius Pryer, 1877
Tetracmanthes Meyrick, 1925
Phalerarcha Meyrick, 1913

Ne SB BNO = =

VOLUME 84, NUMBER 1 43

Cronicombra Meyrick, 1920
Taeniostolella Fletcher, 1940
Machlotica Meyrick, 1909
Abrenthia Busck, 1915
Neomachlotica Heppner, 1981
Trapeziophora Walsingham, 1892
Ussara Walker, 1864
Electrographa Meyrick, 1912

_ Rhabdocrates Meyrick, 1931
Apistomorpha Meyrick, 1881

_ Phryganostola Meyrick, 1881
Pantosperma Meyrick, 1888
Circica Meyrick, 1888
Glyphipterix Hubner, [1825] 2
Diploschizia Heppner, 1981

tn —
WOWN SB De KS KH WK RK NN A

Glyphipterigidae (Copromorphoidea)
Chrysocentris Meyrick, 1914b: 284 (Type-species: C. clavaria Meyrick,

1914).
chalcotypa (Bradley, 1965: 105) (Glyphipterix). New combination. Zaire
chrysozona (Meyrick, 1921b: 112) (Glyphipteryx South Africa
[sic]). New combination.
clavaria Meyrick, 1914b: 284. Malawi
ditiorana (Walker, 1863: 348) (Sciaphila). Malaysia (Sarawak)
eupepla Meyrick, 1930b: 7. Madagascar
infuscata Ghesquiere, 1940: 32. Zaire
phaeometalla Meyrick, 1937: 125. Zaire
urania Meyrick, 1920b: 297. South Africa
Irinympha Meyrick, 1932a: 274 (Type-species: /. aglaograpta Meyrick,
1932).
aglaograpta Meyrick, 1932a: 275. Uganda
Ernolytis Meyrick, 1922: 488 (Type-species: E. chlorospora Meyrick, 1922).
chlorospora Meyrick, 1922: 488. Fiji

Carmentina Meyrick, 1930a: 597 (Type-species: C. iridesma Meyrick, 1930).
Metapodistis Meyrick, 1933: 372 (Type-species: M. chrysosema Meyrick,
1933). New synonymy.

chrysosema (Meyrick, 1933: 372). New combination. Solomon Is.

iridesma Meyrick, 1930a: 598. Solomon Is.

molybdora (Meyrick, 1912: 57) (Glyphipteryx [sic]). Sri Lanka
New combination.

molybdotoma (Diakonoff and Arita, 1979: 93). New combination. Japan

perculta (Diakonoff, 1979: 300). New combination. Taiwan

polychrysa (Meyrick, 1934b: 523). New combination. Indonesia (Java)

44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cotaena Walker, [1865]: 21 (Type-species: C. mediana Walker, [1865]).
mediana Walker, [1865]: 21. Brazil (Para)
plenella (Busck, 1914: 61) (Glyphipteryx [sic]). Panama

Myrsila Boisduval, [1875]: 433 (Type-species: M. auripennis Boisduval,
[1875]).
auripennis Boisduval, [1875]: 433. Brazil (Para)
Lepidotarphius Pryer, 1877: 235 (Type-species: L. splendens Pryer, 1877 [=
Glyphipteryx [sic] perornatella Walker, 1864)).
Desmidoloma Erschoff, 1892: 671 (Type-species: Staintonia fulgens Er-
schoff, 1877 [=Glyphipteryx [sic] perornatella Walker, 1864]).
perornatella (Walker, 1864: 840) (Glyphipteryx [sic]). China
splendens Pryer, 1877: 235.
fulgens (Erschoff, 1877: 347) (Staintonia).

Tetracmanthes Meyrick, 1925: 136 (Type-species: 7. astrocosma Meyrick,

1925).

astrocosma Meyrick, 1925: 136. South Africa

Phalerarcha Meyrick, 1913a: 100 (Type-species: P. chrysorma Meyrick,
1913).

chrysorma Meyrick, 1913a: 101. Guyana

eumitrella (Busck, 1914: 60) (Ussara). New combination. Panama

Cronicombra Meyrick, 1920a: 327 (Type-species: C. granulata Meyrick,
1920).

deltodes (Walsingham, 1914: 301) (Glyphipteryx Mexico (Tabasco)
[sic]). New combination.

essedaria Meyrick, 1926a: 306. Peru

granulata Meyrick, 1920a: 327. Brazil (Para)

lamella (Busck, 1914: 59) (Porpe). Panama

palpella (Walsingham, 1914: 300) (Glyphipteryx Mexico (Tabasco)
[sic]). New combination.

phaeobathra (Meyrick, 1932a: 276). Brazil (Goias)
(Ussara). New combination.

porphyrospila (Meyrick, 1926a: 306) Peru

(Machlotica). New combination.
Taeniostolella Fletcher, 1940: 109 (Type-species: Taeniostola celophora

Meyrick, 1920) [replacement name].

Taeniostola Meyrick, 1920a: 326 (Type-species: 7. celophora Meyrick,
1920) [preocc.].

celophora (Meyrick, 1920a: 327) (Taeniostola). Brazil (Para)

litura (Felder and Rogenhofer, 1875: 14) Brazil (“‘Amazonas’’)
(Oecophora). New combination.

Machlotica Meyrick, 1909b: 36 (Type-species: M. chrysodeta Meyrick,
1909).

VOLUME 84, NUMBER | 45

Maclotica [sic] Busck, 1915: 87, missp.

chrysodeta Meyrick, 1909b: 37. Bolivia
eurymolybda Meyrick, 1926a: 306. Peru
Abrenthia Busck, 1915: 87 (Type-species: A. cuprea Busck, 1915).

cuprea Busck, 1915: 87. USA (Pennsylvania)
Neomachlotica Heppner, 1981b: 479 (Type-species: N. spiraea Heppner,

1981).

actinota (Walsingham, 1914: 301) (Glyphipteryx [sic]). Mexico (Tabasco)
atractias (Meyrick, 1909b: 38) (Machlotica). Bolivia
nebras (Meyrick, 1909b: 37) (Machlotica). Bolivia
spiraea Heppner, 1981b: 481. USA (Florida)

Trapeziophora Walsingham, 1892: 529 (Type-species: 7. gemmula Wal-
singham, 1892).
gemmula Walsingham, 1892: 530. St. Vincent

Ussara Walker, 1864: 800 (Type-species: U. decoratella Walker, 1864).
Setiostoma Felder and Rogenhofer, 1875: 18 (pl. 138, f. 1) (Type-species:
S. flaviceps Felder and Rogenhofer, 1875) [not Setiostoma Zeller, 1875
(Oecophoridae: Stenomatinae)]. New synonymy.
Usara [sic] Busck, [1934]: 182, missp.

ancobathra Meyrick, 1932a: 275. Brazil (Amazonas)
ancyristis Meyrick, 1920a: 334. Brazil (Para)
arquata Meyrick, 1926a: 307. Colombia
chalcodesma Meyrick, 1913a: 67. Guyana
chrysangela Meyrick, 1922: 488. Peru
decoratella Walker, 1864: 801. Brazil (Amazonas)
eurythmiella Busck, 1914: 59. Panama
flaviceps (Felder and Rogenhofer, 1875: 18) Brazil (Amazonas)
(Setiostoma).
hilarodes (Meyrick, 1909c: 430) (Glyphipteryx India (Assam)
[sic]). New combination.
lochrysa Meyrick, 1921a: 184. Indonesia (Java)
olyranta Meyrick, 1931: 184. Brazil (Rio Grande do Sul)
polyastra Meyrick, 1937: 126. South Africa
semicoronis Meyrick, 1932b: 115. Ethiopia
Electrographa Meyrick, 1912: 63 (Type-species: E. thiolychna Meyrick,
1912).
thiolychna Meyrick, 1912: 63. Burma

Rhabdocrates Meyrick, 1931: 183 (Type-species: R. sporomantis Meyrick,
1931).
sporomantis Meyrick, 1931: 184. Peru

46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Apistomorpha Meyrick, 1881: 247 (Type-species: A. argyrosema Meyrick,

1881).
argyrosema Meyrick, 1881: 247. Australia (NSW)
Phryganostola Meyrick, 1881: 248 (Type-species: P. drosophaes Meyrick,
1881).
achlyoessa Meyrick, 1881: 252. New Zealand

achlyoéssa Meyrick, 1881: 252, incorrect spelling.
acyloessa [sic] (Ferro et al., 1977: 9) (Glyphipteryx [sic]), missp.

ataracta Meyrick, 1888: 88. New Zealand
drosophaes Meyrick, 1881: 249. Australia (NSW)
euthybelemna Meyrick, 1881: 250. Australia
macrantha Lower, 1905: 113. Australia (V)
oxymachaera Meyrick, 1881: 251. New Zealand

Pantosperma Meyrick, 1888: 89 (Type-species: P. holochalca Meyrick,
1888).

holochalcum Meyrick, 1888: 89. New Zealand
Circica Meyrick, 1888: 88 (Type-species: C. cionophora Meyrick, 1888).

cionophora Meyrick, 1888: 88. New Zealand
xestobela Meyrick, 1888: 89. New Zealand

Glyphipterix Hubner, [1825]: 421 (Type-species: G. linneella Hubner, [1825]
[=Tinea bergstraesserella Fabricius, 1781]) [ICZN ruling pending: case
2115 (Diakonoff and Heppner, 1977)].

Heribeia Stephens, 1829: 49 (Type-species: Tinea forsterella Fabricius,
1787).

Aechmia Treitschke, 1833: 69 (Type-species: Phalaena (Tortrix) fyeslella
[sic] Fabricius, 1794 [=Phalaena thrasonella Scopoli, 1763)).

Aecimia [sic] Boisduval, 1836: 138, missp.

Glyphipteryx Zeller, 1839: 203, emend. [not Glyphipteryx Curtis, 1827
(Agonoxenidae)].

Glyphiteryx [sic] Fischer von Roslerstamm, 1841: 233, missp.

Anacampsoides Bruand, 1850: 32 (Type-species: Heribeia simpliciella
Stephens, 1834) [nom. oblit.].

Glyphopteryx Herrich-Schaffer, 1854: 92, emend.

Glyphiptoryx [sic] Mann and Rogenhofer, 1878: 500, missp.

Glyphptieryx [sic] Turati, 1879: 203, missp.

Glyphipterys [sic] Christoph, 1882: 38, missp.

Glyphyteryx [sic] Hampson, 1918: 387, missp.

Glyphteryx [sic] Watt, 1920: 439, missp.

acinacella Meyrick, 1883: 193 (Glyphipteryx [sic]). Australia (V)

acronoma Meyrick, 1888: 86 (Glyphipteryx [sic]). New Zealand

VOLUME 84, NUMBER 1 47

acrothecta Meyrick, 1881: 244 (Glyphipteryx [sic]). New Zealand
actinobola Meyrick, 1881: 241 (Glyphipteryx [sic]). Australia (NSW)
aechmiella (Duponchel, 1838: 533) (Elachista). France
oechmiella [sic] (Duponchel, 1838: pl. 308, f. 7) (Elachista), missp.
aerifera Meyrick, 1912: 57 (Glyphipteryx [sic]). New Zealand
albimaculella Heinemann, [1876]: 397 (Glyphipteryx [sic]). Germany
alpha Moriuti and Saito, 1964: 61. Japan
altiorella Bauer, 1923: 167 (Glyphipteryx [sic}). Austria
amblycerella Meyrick, 1883: 189 (Glyphipteryx [sic]). Australia (V)
ametris Diakonoff, 1979: 307. Taiwan
~amphipeda Meyrick, 1920b: 295 (Glyphipteryx [sic]). South Africa
amseli Diakonoff, 1978: 45. China (Chekiang)
anaclastis Meyrick, 1907a: 130 (Glyphipteryx [sic]). Australia
angulosella (O. Costa, 1834a: 4) (Oecophora). Italy
pernicipennella (O. Costa, [1836]: [301]) (Elachista (Ornix)).
~ antidoxa Meyrick, 1909c: 435 (Glyphipteryx [sic]). Sri Lanka
archimedica Meyrick, 1921b: 113 (Glyphipteryx [sic]). Zimbabwe
argophracta Meyrick, 1926b: 338 (Glyphipteryx [sic]). South Africa
argyrelata Turner, 1932: 192 (Glyphipteryx [sic]). Australia (Q)
argyroguttella Ragonot, 1885: lvi (Glyphipteryx [sic]). Italy
ab. paurographella Ragonot, 1885: lvi (Glyphipteryx [sic]).
argyromis Meyrick, 1907c: 985 (Glyphipteryx [sic]). Sri Lanka
argyrotoxa Turner, 1913: 214 (Glyphipteryx [sic]). Australia (Q)
asterias (Meyrick, 1907d: 154) (Anticrates). India (Assam)
asteriella Meyrick, 1881: 235 (Glyphipteryx [sic]). Australia (NSW)
asteronota Meyrick, 1881: 240 (Glyphipteryx [sic]). New Zealand
astrapaea Meyrick, 1881: 245 (Glyphipteryx [sic]). New Zealand
atelura Meyrick, 1920a: 331 (Glyphipteryx [sic]). Guyana
aulogramma Meyrick, 1907a: 121 (Glyphipteryx [sic]). New Zealand
autopetes Meyrick, 1907a: 118 (Glyphipteryx [sic]). Australia (WA)
bactrias Meyrick, 191la: 67 (Glyphipteryx [sic]). New Zealand
basifasciata Issiki, 1930: 427 (Glyphipteryx [sic]). Japan
jezonica Matsumura, 1931: 1079 (Glyphipteryx [sic]).
bergstraesserella (Fabricius, 1781: 302) (Tinea). Germany

bergstraessella [sic] (Fabricius, 1794: 321) (Tinea), missp.
lineella (Hubner, 1796: pl. 65, f. 436) (Tinea).

lineana (Hubner, [1796-99]: pl. 14, f. 84) (Tortrix).

linneana [sic] Hubner, [1825]: 421, missp.

linneella [sic] Htibner, [1825]: 421, missp.

treitschkeana (Frolich, 1828: 76) (Tortrix).

bergstraesella [sic] Bruand, 1850: 48 (Glyphipteryx [sic]), missp.
pietruskii Nowicki, 1864: 22 (Glyphipteryx [sic]).

48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ab. arcuatella Klemensiewicz, 1907: 19 (Glyphipteryx [sic]).
bergstrdsserella [sic] Amsel, 1950: 26 (Glyphipteryx [sic]), missp.

beta Moriuti and Saito, 1964: 61. Japan

Japonicella Matsumura, 1931: 1079 (Glyphipteryx [{sic]), not Zeller, 1877.
bicornis Meyrick, 1909c: 432 (Glyphipteryx [sic]). India (Assam)
bifasciata Walsingham, 1881: 321 (Glyphipteryx {sic]). USA (California)
bifasciella Amsel, 1959: 61 (Glyphipteryx [sic]). Iraq
bohemani (Zeller, 1852: 116) (Aechmia). South Africa
brachyaula Meyrick, 1907a: 127 (Gtyphipteryx [sic]). Australia (Q)
brachydelta Meyrick, 1916: 418 (Glyphipteryx [sic]). New Zealand
californiae Walsingham, 1881: 320 (Glyphipteryx USA (California)

[sic]).

calliactis Meyrick, 1914a: 112 (Glyphipteryx [sic]). New Zealand
callicrossa Meyrick, 1907a: 119 (Glyphipteryx [sic]). Australia (WA)
callidelta Meyrick, 1922: 492 (Glyphipteryx [sic]). Peru
calliscopa Lower, 1905: 112 (Glyphipteryx [sic]). Australia (V)
callithea Meyrick, 1921b: 113 (Glyphipteryx [sic]). South Africa
canachodes Meyrick, 1909c: 434 (Glyphipteryx [sic]). India (Madras)
carenota Meyrick, 1909c: 431 (Glyphipteryx [sic]). India (Assam)
caudatella Walsingham, 1897: 119 (Glyphipteryx [sic]). Grenada
cestrota Meyrick, 1915b: 221 (Glyphipteryx [sic]). Peru
chalcodaedala Turner, 1913: 215 (Glyphipteryx [sic]). Australia (Q)
chalcostrepta Meyrick, 1907a: 128 (Glyphipteryx [sic]). Australia (T)
chionosoma Diakonoff, 1978: 54. China (Kiangsu)
chrysallacta Meyrick, 1922: 491 (Glyphipteryx [sic]). Peru
chrysoplanetis Meyrick, 1881: 238 (Glyphipteryx [sic]). Australia
circumscriptella Chambers, 1881: 291 USA (Massachusetts)

(Glyphipteryx [sic]).
circumscripta [sic] Dyar, 1900: 84 (Glyphipteryx [sic]) missp.

clearcha Meyrick, 192la: 185 (Glyphipteryx [sic]). Indonesia (Java)
climacaspis Meyrick, 1920b: 296 (Glyphipteryx [sic]). South Africa
codonias Meyrick, 1909a: 15 (Glyphipteryx [sic]). New Zealand
colorata Meyrick, 1913a: 103 (Glyphipteryx [sic]). Guyana
columnaris Meyrick, 1913a: 69 (Glyphipteryx [sic]). Guyana
cometophora Meyrick, 1881: 231 (Glyphipteryx [sic]). Australia
compastis Meyrick, 1923b: 618 (Glyphipteryx [sic]). India (Assam)
conjunctella (O. Costa, 1834a: 5) (Oecophora). Italy

colluripennella (O. Costa, 1834b: 77) (Ornix), nom. nud.
colluripennella (O. Costa, [1836]: [301]) (Elachista (Ornix)).

conosema Meyrick, 1913a: 69 (Glyphipteryx [sic]). Guyana
cornigerella Zeller, 1877: 403 (Glyphipteryx [sic]). Colombia
crassilunella Rebel, 1916: 157 (Glyphipteryx [sic]). Greece (Crete)

crinita Meyrick, 1913a: 102 (Glyphipteryx [sic]). Guyana

VOLUME 84, NUMBER I 49

crotalotis Meyrick, 1909c: 432 (Glyphipteryx [sic]). India (Assam)

cultrata Meyrick, 1912: 56 (Glyphipteryx [sic]). India (Assam)

cyanophracta Meyrick, 1883: 186 (Glyphipteryx [sic]). | Australia (NSW)

cyanochalca Meyrick, 1883: 185 (Glyphipteryx [sic]). Australia (NSW)
lyelliana Lower, 1893: 182 (Glyphipteryx [sic]).

danilevskii Diakonoff, 1978: 49. USSR (Kazakh SSR)
decachrysa Meyrick, 1918a: 36 (Glyphipteryx [sic]). South Africa
deliciosa Diakonoff, 1978: 5S. China (Chekiang)
delta Moriuti and Saito, 1964: 62. Japan
deuterastis Meyrick, 1907a: 121 (Glyphipteryx [sic]). Australia (WA)
-diaphora Walsingham, 1910: 258 (Glyphiptery x [sic]). Madeira Is.
dichalina Meyrick, 1911b: 290 (Glyphipteryx [sic]). Seychelles Is.
dichorda Meyrick, 191la: 76 (Glyphipteryx [sic]). New Zealand
| diplotoxa Meyrick, 1920b: 296 (Glyphipteryx [sic]). South Africa
dolichophyes Diakonoff, 1978: 50. China (Kiangsu)
| xyridota Meyrick, 1935: 87 (Glyphipteryx [sic]), not Meyrick, 1918.

-enclitica Meyrick, 1909c: 430 (Glyphipteryx [sic]). India (Mysore)
-epastra Meyrick, 1922: 490 (Glyphipteryx [sic]). Peru
| equitella (Scopoli, 1763: 254) (Phalaena). [Austria]

majorella (Heydenreich, 1851: 87) (Aechmia), nom. nud.
majorella Mann, 1854: 590 (Glyphipteryx [sic]), nom. nud.
var. majorella (Herrich-Schaffer, 1854: 93) (Aechmia).

| maiorella [sic] Prohaska, 1905: 19 (Glyphipteryx [sic]), missp.

)erastis Meyrick, 191la: 76 (Glyphipteryx [sic]). New Zealand
| erebanassa Meyrick, 1934a: 41 (Glyphipteryx [sic]). | China (Kwangtung)
euastera Meyrick, 1881: 236 (Glyphipteryx [sic]). New Zealand
euleucotoma Diakonoff and Arita, 1976: 208. Japan
expurgata Meyrick, 1922: 493 (Glyphipteryx [sic]). Peru
falcigera Meyrick, 1913a: 102 (Glyphipteryx [sic]). Guyana
forsterella (Fabricius, 1781: 509) (Tinea). Germany

humerella (Htibner, [1800-05]: pl. 42, f. 292) (Tinea), not

| Denis and Schiffermuller, 1775 [Gelechiidae].

lucasella (Duponchel, 1838: 440) (Aechmia).

oculatella (Zeller, 1850: 157) (Aechmia).

oculatella Mann, 1854: 591 (Glyphipteryx [sic]), nom. nud.
form nivicaput Diakonoff, 1979: 311, invalid [ICZN, Art. 15].

fortunatella Walsingham, 1908: 989 (Glyphipteryx [sic]). Canary Is.
-funditrix Diakonoff and Arita, 1976: 211. Japan
_fuscoviridella (Haworth, 1828: 569) (Tinea). [England]

albicostella (Duponchel, 1838: 397) (Adela).

aereinitidella (Milliere, 1854: 60) (Cephalispheira).
gamma Moriuti and Saito, 1964: 62. Japan
gaudialis Diakonoff and Arita, 1976: 198. Japan

50 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

gemmatella (Walker, 1864: 1022) (Gelechia).
gemmipunctella (Walker, 1869: 86) (Gelechia).
atristriella Zeller, 1877: 398 (Glyphipteryx [sic]).

chrysolithella Meyrick, 1881: 229 (Glyphipteryx [sic]).

gemmula Diakonoff and Arita, 1976: 194.
gianelliella Ragonot, 1885: lvi (Glyphipteryx [sic]).
gonoteles Meyrick, 1907a: 119 (Glyphipteryx [sic]).

grapholithoides Walsingham, 1891: 116 (Glyphipteryx [sic]).

gypsonota Turner, 1927: 158 (Glyphipteryx [sic]).
halimophila Lower, 1893: 183 (Glyphipteryx [sic]).
haplographa Turner, 1927: 157 (Glyphipteryx [sic]).
harpogramma Turner, 1913: 214 (Glyphipteryx [sic]).
haworthana (Stephens, 1834: 262) (Heribeia).

haworthella (Stephens, 1829: 49) (Heribeia), nom. nud.

zonella (Zetterstedt, [1839]: 1009) (Oecophora).

Sierra Leone
Australia (V)

Japan

Italy

Australia
South Africa
Australia (T)
Australia (SA)
Australia (T)
Australia
[England]

howarthana [sic] Jordan, 1886: 154 (Glyphipteryx [sic]), missp.

hemipempta Meyrick, 1909c: 429 (Glyphipteryx [sic]).

heptaglyphella Le Marchand, 1925: 314 (Glyphipteryx [sic]).

holodesma Meyrick, 1883: 190 (Glyphipteryx [sic]).
hologramma Meyrick, 1920a: 333 (Glyphipteryx [sic]).
hyperlampra Turner, 1913: 216 (Glyphipteryx [sic]).
idiomorpha Meyrick, 1917: 10 (Glyphipteryx [sic]).
imparfasciata Arita, 1979: 77.
indomita Meyrick, 1922: 491 (Glyphipteryx [sic]).
invicta Meyrick, 1920a: 334 (Glyphipteryx [sic]).
iinvicta [sic] Clarke, 1969: 68, missp.
iocheaera Meyrick, 1881: 243 (Glyphipreryx |sic)).
ioclista Meyrick, 1913a: 70 (Glyphipteryx [sic]).
iometalla Meyrick, 1881: 232 (Glyphipteryx [sic]).
isoclista Meyrick, 1925: 136 (Glyphipteryx [sic]).
isozela Meyrick, 1907a: 124 (Glyphipteryx [sic]).
japonicella Zeller, 1877: 400 (Glyphipteryx [sic}).
lamprocoma Meyrick, 1907a: 127 (Glyphipteryx [sic]).
lamprosema Turner, 1926: 144 (Glyphipteryx [sic]).
leptocona Meyrick, 1922: 493 (Glyphipteryx [sic]).
leptosema Meyrick, 1888: 87 (Glyphipteryx [sic}).
leucargyra Turner, 1927: 157 (Glyphipteryx [sic]).
leucocerastes Meyrick, 1881: 239 (Glyphipteryx [sic]).

leucophragma Meyrick, 1923a: 564 (Glyphipteryx [sic)).

leucoplaca Turner, 1913: 217 (Glyphipteryx [sic]).
longistriatella Rebel, 1940: 37 (Glyphipteryx [sic]).
loricatella (Treitschke, 1833: 70) (Aechmia).

Sri Lanka
France
Australia (T)
Brazil (Para)
Australia (Q)
South Africa

Japan (Ryukyu Is.)
Brazil (Amazonas)

Brazil (Para)

New Zealand
Guyana
Australia (Q)
Fiji

Australia

Japan

Australia (SA)
Australia (NSW)
Peru

New Zealand
Australia (T)
Australia (NSW)
Angola
Australia
Azores Is.
Hungary

VOLUME 84, NUMBER I

51

loricutella [sic] Desmarest, [1857]: 282 (Glyphipteryx [sic]), missp.

luteocapitella Caradja, 1926: 41 (Glyphip-
teryx [sic]).
luteomaculata Arita, 1979: 79.
lycnophora Turner, 1913: 213 (Glyphipteryx [sic]).
macraula Meyrick, 1907a: 120 (Glyphipteryx [sic]).
macrodrachma Diakonoff, 1979: 305.
madagascariensis Viette, 1951: 340 (Glyphipteryx [sic]).
magnatella Erschoff, 1877: 346 (Glyphipteryx [sic]).
maritima Diakonoff, 1979: 303.

_marmaropa Turner, 1913: 218 (Glyphipteryx [sic]).
maschalis Meyrick, 1909c: 433 (Glyphipteryx [sic]).
medianella Turati, 1930: 211 (Glyphipteryx [sic]).

_medica Meyrick, 1911b: 291 (Glyphipteryx [sic]).

-melania Diakonoff and Arita, 1976: 213.

_-mesaula Meyrick, 1907a: 121 (Glyphipteryx [sic]).

-metasticta Meyrick, 1907a: 120 (Glyphipteryx [sic]).

-meteora Meyrick, 1881: 237 (Glyphipteryx [sic]).

| chalceres Turner, 1913: 215 (Glyphipteryx [sic]).

-metron Diakonoff, 1948: 206 (Glyphipteryx [sic]).

-metronoma Meyrick, 1907a: 123 (Glyphipteryx [sic]).

-minorella Snellen, 1882: 753 (Glyphipteryx [sic]).

equitella of authors, not Scopoli, 1763.

_molybdastra Meyrick, 1923a: 564 (Glyphipteryx [sic)).

-~monodonta Diakonoff, 1948: 206 (Glyphipteryx [sic]).
montisella Chambers, 1875: 292 (Glyphipteryx [sic]).

USSR (Kemerovo ASSR)

Japan (Ryukyu Is.)

Australia (Q)
Australia (T)
China (Chekiang)
Madagascar
USSR (Irkutsk)
USSR (Maritime)
Australia

Sri Lanka
Greece (Rhodes)
Seychelles Is.
Japan

Australia (SA)
New Zealand
Australia (NSW)

Indonesia (Buru)
Australia (V)
Netherlands

Angola
Indonesia (Buru)
USA (Colorado)

montinella [sic] Chambers, 1877: 14 (Glyphipteryx [sic]), missp.
montella Meyrick, 1913b: 44 (Glyphipteryx [sic]), emend.

teryx [sic]).
nattani Gozmany, 1954: 277 (Glyphipteryx [sic]).
-necopina Philpott, 1927: 88 (Glyphipteryx [sic]).
neochorda Meyrick, 1922: 493 (Glyphipteryx [sic]).
nephoptera Meyrick, 1888: 87 (Glyphipteryx [sic]).
|
:

|
|
morangeila Felder and Rogenhofer, 1875: 10 (Glyphip-

nicaeella Méschler, 1866: 146 (Glyphipteryx [sic]).
| nigromarginata Issiki, 1930: 426 (Glyphipteryx {[sic]}).
suzukii Matsumura, 1931: 1079 (Glyphipteryx [sic]).

nugella Felder and Rogenhofer, 1875: 10 (Glyphipteryx [sic]).

—octatoma Diakonoff, 1978: 48.

octonaria Philpott, 1924: 210 (Glyphipteryx [sic]).
okui Diakonoff and Arita, 1976: 206.

oligastra Meyrick, 1926a: 308 (Glyphipteryx [sic]).
orthodeta Meyrick, 1922: 490 (Glyphipteryx [sic]).

New Zealand

Hungary
New Zealand
Peru

New Zealand
France

Japan

Colombia
China (Chekiang)
New Zealand
Japan

Colombia

Brazil (Amazonas)

52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ortholeuca Meyrick, 1921b: 113 (Glyphipteryx [sic]). South Africa
orthomacha Meyrick, 1920a: 331 (Glyphipteryx [sic]). Australia (Q)
orymagdis Meyrick, 1909c: 435 (Glyphipteryx [sic]). Sri Lanka
oxycopis Meyrick, 1918b: 195 (Glyphipteryx [sic]). India (Assam)
oxydonta Diakonoff, 1978: 57. China (Chekiang)
oxyglypta (Meyrick, 1929: 519) (Acrolepia). New combination. Panama
oxytricha Meyrick, 1928: 424 (Glyphipteryx [sic]). South Africa
palaeomorpha Meyrick, 1881: 242 (Glyphipteryx [sic]). Australia (Q)
paradisea Walsingham, 1897: 119 (Glyphipteryx [sic]). Grenada
parazona Meyrick, 1907a: 125 (Glyphipteryx [sic]). Australia (V)
perfracta Meyrick, 1922: 493 (Glyphipteryx [sic]). Peru
perimetalla Lower, 1905: 112 (Glyphipteryx [sic]). Australia (V)
persica Diakonoff, 1979: 309. Iran
pertenuis Diakonoff, 1979: 306. Tunisia
pharetropis Meyrick, 1907a: 125 (Glyphipteryx [sic]). Australia (V)
phosphora Meyrick, 1907a: 126 (Glyphipteryx [sic]). Australia (SA)
plagiographa Bradley, 1965: 106. Uganda
platydisema Lower, 1893: 183 (Glyphipteryx [sic]). Australia (V)
platyochra Meyrick, 1920a: 333 (Glyphipteryx [sic]). Brazil (Para)
polychroa Lower, 1897: 23 (Glyphipteryx [sic]). Australia (V)
polyzela Meyrick, 1920a: 331 (Glyphipteryx [sic]). Australia (Q)
protomacra Meyrick, 1907a: 117 (Glyphipteryx [sic]). Australia (WA)
pseudostoma Meyrick, 1922: 492 (Glyphipteryx [sic]). Guyana
psychopa Meyrick, 1909c: 433 (Glyphipteryx [sic]). Sri Lanka
pygmaeella Rebel, 1896: 132 (Glyphipteryx [sic]). Canary Is.
pyristacta Turner, 1913: 213 (Glyphipteryx [sic]). Australia (Q)
pyrogastra Meyrick, 1909c: 429 (Glyphipteryx [sic]). India (Assam)
pyrophora Turner, 1913: 217 (Glyphipteryx [sic]). Australia (NSW)
quadragintapunctata Dyar, 1900: 84 (Glyphipteryx [sic]). USA (Kansas)
refractella Zeller, 1877: 402 (Glyphipteryx [sic]). Colombia
regula Diakonoff and Arita, 1976: 202. USSR (Kuril Is.)
repletana (Walker, 1864: 1037) (Ussara). Brazil (Amazonas)
rhanteria Turner, 1913: 216 (Glyphipteryx [sic]). Australia (Q)
rhinoceropa Meyrick, 1935: 86 (Glyphipteryx [sic]). China (Chekiang)
rhodanis Meyrick, 1909c: 434 (Glyphipteryx [sic]). India (Assam)
rugata Meyrick, 1915a: 203 (Glyphipteryx [sic]). New Zealand
sabella Newman, 1856: 299 (Glyphipteryx [sic]). Australia (V)
saurodonta Meyrick, 1913a: 68 (Glyphipteryx [sic]). Canada (Ontario)
schoenicolella Boyd, 1859: 144 (Glyphipteryx [sic]). England
schultzella Amsel, 1949: 88 (Glyphipteryx [sic]). Germany
scintilla C. Clarke, 1926: 420 (Glyphipteryx [sic]). New Zealand

sclerodes Meyrick, 1909c: 432 (Glyphipteryx [sic]). Sri Lanka

| VOLUME 84, NUMBER 1 53

scolias Meyrick, 1910: 73 (Glyphipteryx New Zealand (Kermadec Is.)
[sic]).

- semiflavana Issiki, 1930: 426 (Glyphipteryx [sic]). Japan

_ semifravana [sic] Kodama, 1961: 45 (Glyphipteryx [sic]), missp.

_ semilunaris Wollaston, 1879: 436 (Glyphipteryx [sic]). St. Helena
semisparsa Meyrick, 1918b: 195 (Glyphipteryx [sic]). India (Madras)
seppella (Fabricius, 1776: 296) (Tinea). England
septemstrigella Zeller, 1877: 405 (Glyphipteryx [sic]). Colombia
sexguttella Toll, 1936: 404 (Glyphipteryx [sic]). Poland

_ similis Philpott, 1928: 369 (Glyphipteryx [sic]). New Zealand

| simplicella Christoph, 1882: 38 (Glyphipteryx [sic]). USSR (Maritime)
simpliciella (Stephens, 1834: 263) (Heribeia). [England]

cramerella of authors, not Fabricius, 1777 [Gracillariidae].

cognatella (Stephens, 1834: 261) (Heribeia).

variella Zeller, 1839: 203 (Glyphipteryx [sic]), not Fabricius, 1794
[Incurvariidae].

fischeriella (Zeller, 1839: 204) (Aechmia).

roeslerstammella (Fischer von Roéslerstamm, 1841: 242) (Aechmia).

desiderella (Fischer von Roslerstamm, 1841: 242) (Aechmia).

desideratella [sic] (Duponchel, 1842: 322) (Aechmia), missp.

_speculans Meyrick, 1922: 490 (Glyphipteryx [sic]). Brazil (Amazonas)
_ speculiferella Christoph, 1882: 37 (Glyphipteryx [sic]). USSR (Maritime)
-stasichlora Meyrick, 1931: 184 (Glyphipteryx [sic}). Peru
stelucha Meyrick, 1909d: 375 (Glyphipteryx [sic}). South Africa
stilata Meyrick, 1912: 57 (Glyphipteryx [sic]). Sri Lanka
struvei Amsel, 1938: 113 (Glyphipteryx [sic]). Germany
sulcosa Diakonoff, 1978: 58. Italy (Sardinia)
synarma Meyrick, 1909c: 431 (Glyphipteryx [sic]). India (Assam)
syndecta Meyrick, 1915b: 221 (Glyphipteryx [sic}). Peru
synorista Meyrick, 1922: 492 (Glyphipteryx [sic]). Brazil (Amazonas)
talhouki Diakonoff, 1978: 61. Lebanon
tetrachrysa Meyrick, 1907b: 749 (Glyphipteryx [sic]). Sri Lanka
tetrasema Meyrick, 1883: 191 (Glyphipteryx [sic]). Australia (T)
thrasonella (Scopoli, 1763: 253) (Phalaena). [Austria]

fueslella (Fabricius, 1787: 250) (Tinea).

fyeslella (Fabricius, 1794: 318) (Tinea).

ahornerella (Hiibner, 1796: pl. 26, f. 178) (Tinea).

seppella (Hiibner, 1796: pl. 32, f. 223) (Tinea), not Fabricius, 1776.
aillyella (Hubner, 1796: pl. 64, f. 431) (Tinea).

triguttella (Donovan, 1804: pl. 382, f. 1) (Phalaena).

fueslii (Haworth, 1828: 531) (Gracillaria), emend.

equitella (Treitschke, 1833: 71) (Aechmia), not Scopoli, 1763.

54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

poeyella (Duponchel, 1838: 438) (Aechmia).

fuselella [sic] (Zeller, 1839: 203), (Tinea) missp. [in synonymy].
fiesiella [sic] (Westwood, 1840: 112) (Aechmia), missp.
cladiella Stainton, 1859: 364 (Glyphipteryx [sic]).

var. albardella Snellen, 1882: 752 (Glyphipteryx [sic]).

var. eyndhoveniella Snellen, 1882: 752 (Glyphipteryx [sic]).

ab. nitens Bankes, 1907: 204 (Glyphipteryx [sic]).

thravonella [sic] Meyrick, 1914c: 28 (Glyphipteryx [sic]), missp.

transversella (Walker, 1864: 849) (Argyresthia). New Zealand
trigonaspis Meyrick, 1907a: 131 (Glyphipteryx [sic]). Australia (WA)
trigonodes Arita, 1979: 79. Japan (Ryukyu Is.)
tripedila Meyrick, 1912: 58 (Glyphipteryx [sic]). India (Assam)
‘‘triplaca’’ Turner; Meyrick, 1913b: 44 (Glyphipteryx Australia (Q)
[sic]), nom. nud., nom. dub.
tripselia Meyrick, 1907a: 129 (Glyphipteryx [sic]). Australia (V)
triselena Meyrick, 1881: 234 (Glyphipteryx [sic]). New Zealand
tungella Felder and Rogenhofer, 1875: 10 (Glyphipteryx New Zealand
[sic]).
umbilici Hering, 1927: 431 (Glyphipteryx [sic]). Canary Is.
uncta Meyrick, 1913a: 101 (Glyphipteryx [sic]). Guyana
unguifera Meyrick, 1922: 489 (Glyphipteryx [sic]). Peru
unifasciata Walsingham, 1881: 322 (Glyphipteryx [sic]).. USA (California)
variata Meyrick, 1913a: 104 (Glyphipteryx [sic]). Guyana
versicolor Meyrick, 1913a: 103 (Glyphipteryx [sic]). Guyana

voluptella Felder and Rogenhofer, 1875: 10 (Glyphip- _ Brazil (Amazonas)
teryx [sic]}).

xanthoplecta Meyrick, 1922: 491 (Glyphipteryx [sic]). Peru
xyridota Meyrick, 1918b: 194 (Glyphipteryx [sic]). India (Assam)
zalodisca Meyrick, 1920a: 332 (Glyphipteryx [sic]). Brazil (Para)
zelota Meyrick, 1888: 86 (Glyphipteryx [sic]). New Zealand
zermattensis Amsel, 1932: 20 (Glyphipteryx [sic]). Germany

Diploschizia Heppner, 198la: 311 (Type-species: Glyphipteryx [sic] im-
pigritella Clemens, 1863).
glaucophanes (Meyrick, 1922: 489) (Glyphipteryx [sic]) Brazil (Amazonas)
habecki Heppner, 1981la: 317 USA (Florida)
impigritella (Clemens, 1863: 9) (Glyphipteryx [sic]). USA (Pennsylvania)
exoptatella (Chambers, 1875: 234) (Glyphipteryx [sic]).

kimballi Heppner, 198la: 325. USA (Florida)
lanista (Meyrick, 1918b: 195) (Glyphipteryx [sic]). USA (N. Carolina)
minimella Heppner, 198la: 315. USA (Florida)

regia Heppner, 198la: 320. USA (Florida)

|
|

VOLUME 84, NUMBER 1 55
tetratoma (Meyrick, 1913a: 68) (Glyphipteryx [sic]). Guyana
urophora (Walsingham, 1914: 300) (Glyphipteryx [sic]). Guatemala

ACKNOWLEDGMENTS

Preliminary research resulting in this summary was partially made pos-
sible by a grant from the National Science Foundation (76-12550 DEB) and
the University of Florida, Department of Entomology and Nematology
(IFAS), Gainesville, Florida (while at this institution), for study of collec-
tions at the British Museum (Natural History), London, England, in 1976—
77. | also acknowledge the support of the Smithsonian Institution in this
visit to London, as well as subsequent research on this group of moths.
Special thanks are also due to my colleagues, J. F. G. Clarke (Smithsonian
Institution) and A. Diakonoff (Rijksmuseum van Natuurlijke Historie, Lei-
den, Netherlands), who have long studied Glyphipterigidae and have al-
lowed many periods of discussion regarding this family.

LITERATURE CITED

Amsel, H. G. 1932. Neue mitteleuropaische Kleinschmetterlinge und Bemerkungen Uber
| Melasina lugubris Hb. und M. ciliaris O. (Lep.). Dtsch. Entomol. Z. 46: 18-24, 1 pl.
. 1938. Neue oder seltene Kleinschmetterlinge aus dem nordwestdeutschen Faunen-
gebiet, nebst Bemerkungen tiber die Rassenbildung bei Gelechia ericetella Hb. in Mittel-
und Siideuropa (Lepidoptera: Gelechiidae, Glyphipterygidae). Abh. Naturwiss. Ver.
Bremen 30: 108-114.
; ———. 1949. Eine neue deutsche Glyphipteryx-Art (Lep. Glyphipterygidae). Entomon 1: 88-
| 89.
| ———. 1950. Kleinschmetterlinge aus dem badischen Schwarzwald. Beitr. Naturkd. Forsch.
| Siidwestdtschl. 9: 26-28.
. 1959. Irakische Kleinschmetterlinge, II. Bull. Soc. Entomol. Egypte 43: 41-83.
Arita, Y. 1979. Three new Glyphipterix species from Ishigaki Island, the Ryukyu Islands
(Lepidoptera: Glyphipterigidae). Tyo to Ga 30: 77-82.
Bankes, E. R. 1907. Glyphipteryx thrasonella, Scop., ab. nitens, n. ab. Entomol. Mon. Mag.
43: 204.
Bauer, E. 1923. Eine neue Glyphipteryx (Microlep.) aus den Alpen. Entomol. Mitt. 12: 167-
168.
Boisduval, J. B. A. D. de. 1836. Histoire naturelle des insectes. Species général des Lepi-
dopteres. Vol. 1. Roret, Paris. 690 pp.
| ———.. [1875]. Sphingides, Sesiides, Castniides. Jn Boisduval, J. B. A. D. de and A. Gueneée,
Histoire naturelle de insectes. Spécies général des Lépidopteres Heteroceres. Vol. 1.
| Roret, Paris. 568 pp.
Boyd, T. 1859. Notes on an entomological tour in Cornwall. Entomol. Weekly Intelligencer
| 1858: 142-144.
Bradley, J. D. 1965. Microlepidoptera. Jn Ruwenzori Expedition 1952. Vol. II, no. 12. Br.
Mus. (Nat. Hist.), London. Pp. 81-148.

Bruand, C. T. 1850. Catalogue systematique et synonymique des Lepidopteres de Deéparte-
ment du Doubs. Tineides. Mém. Soc. Emul. Doubs 3(3): 23-68.

Busck, A. 1914. New genera and species of microlepidoptera from Panama. Proc. U.S. Natl.
Mus. 47: 1-67.

56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

———. 1915. Descriptions of new North American microlepidoptera. Proc. Entomol. Soc.

Wash. 17: 79-94.

. [1934]. Microlepidoptera of Cuba. Entomol. Am. 13: 151-202, pls. 30-36.

Caradja, A. von. 1926. Ueber einige bei Sutschansk gesammelte Pyraliden und sonstige Klein-
falter. Dtsch. Entomol. Z. Iris 40: 36-43.

Chambers, V. T. 1875. Teneina [sic] of Colorado. Cincinnati Q. J. Sci. 2: 289-305.

. 1877. Tineina. Can. Entomol. 9: 13-15.

. 1881. New species of Tineina. J. Cincinnati Soc. Nat. Hist. 3: 289-296.

Christoph, H. 1882. Neue Lepidopteren des Amur-Gebietes. Bull. Soc. Imp. Natr. Moscou
57: 5—47.

Clarke, C. E. 1926. New species of Lepidoptera. Trans. N. Z. Inst. 56: 417-421.

Clarke, J. F. G. 1969. Catalogue of the type specimens of Microlepidoptera in the British
Museum (Natural History) described by Edward Meyrick. Vol. VI: Glyphipterigidae,
Gelechiidae (A-C). Br. Mus. (Nat. Hist.), London. 537 pp., 267 pls.

Clemens, B. 1863. American micro-lepidoptera. Proc. Entomol. Soc. Phil. 2: 4-14.

Costa, O. G. 1834a. Specie nuove di Lepidotteri del Regno di Napoli. I. Naples. 8 pp., | pl.

1834b. Cenni zoologici ossia descrizione sommaria delle specie nuove di animali

discoperti in diverse contrade del Regno nell anno 1834. Annuario Zool. (Naples) 1834:

1-90.

—.. [1836]. Asopia. In Fauna del Regno di Napoli ossia enumerazione di tutti gli animali
che abitano le diverse regioni di questo regno e le acque che le bagnano contenente la
descrizione de nuovi 0 poco esattamente conosciuti. Lepidotteri. Torchi, Naples. Pp.
[196-197; and 301].

Desmarest, E. [1857]. Papillons nocturnes. Jn Chenu, J. C., Encyclopédie d’histoire naturelle
ou traité complet de cette science d’apres les travaux des naturalists les plus éminents
de tous les pays et de toutes les €poques. Vol. 10. Havard, Paris. 321 pp., 39 pls.

Diakonoff, A. 1948. Fauna Buruana. Microlepidoptera I. Treubia 19: 197-219.

. 1978. Descriptions of new genera and species of the so-called Glyphipterygidae sensu

Meyrick, 1913 (Lepidoptera). Zool. Verh. (Leiden) 160: 1-63.

———. 1979. Descriptions of new taxa of the so-called Glyphipterigidae auctorum (Lepidop-
tera). Zool. Meded. (Leiden) 54; 291-312.

Diakonoff, A. and Y. Arita. 1976. New and old species of Glyphipterigidae and a new species
of Acrolepiidae from Japan and the Far East (Lepidoptera). Zool. Meded. (Leiden) 50:
179-219.

———. 1979. Three new species of the so-called Glyphipterigidae auctorum from Japan (Lep-
idoptera). Zool. Meded. (Leiden) 54: 87-100.

Diakonoff, A. and J. B. Heppner. 1977. Proposed use of the plenary powers to designate a
type-species for the nominal genus Glyphipterix Hiibner, [1825] (Lepidoptera, Glyphip-
terygidae). Z. N. (S.) 2115. Bull. Zool. Nomencl. 34: 81-84.

Donovan, E. 1792-1813. The natural history of British insects; explaining them in their several
states, with the periods of their transformations, their food, oeconomy and c. Together
with the history of such minute insects as require investigation by the microscope.
London. 16 vols.

Duponchel, P. A. J. 1838. Nocturnes. Jn Godart, J. B., ed., Histoire naturelle des Lépidop-

teres ou Papillons de France. Vol. 11. Méquignon-Marvis, Paris. 720 pp., pls. 287-314.

. 1842. Nocturnes. Suppl. Jn Godart, J. B., ed., Histoire naturelle des Lépidoptéres ou

Papillons de France. Suppl. Méquignon-Marvis, Paris. 555 pp., pls. 51-90.

Dyar, H. G. 1900. Notes on some North American Yponomeutidae. Can. Entomol. 32: 37-
41, 84-86.

Erschoff, N. 1877. Diagnosen neuer Lepidopteren aus den verschiedenen Provinzen des rus-
sischen Reiches. Hor. Entomol. Soc. Ross. 12: 336-348.

VOLUME 84, NUMBER | 57

—. 1892. Verzeichniss von Schmetterlinge aus Central-Siberien. Mem. Lepid. Romonoff
6: 670-672, pl. 16.

Fabricius, J. C. 1776. Genera insectorum eorumque characteres naturales secundum numer-
um, figuram, situm et proportionem omnium partium oris asiecta mantissa specierum
nuper detectrum. Kiel. 310 pp.

———. 1781. Species insectorum exhibentes eorum differentias specificas, synonyma auc-
torum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. Bohn,
Hamburg and Cologne. 2 vols., 510 pp.

_| ———. 1787. Mantissa insectorum sistens species nuper detectas adiectis synonymis, obser-

| vationibus, descriptionibus, emendationibus. Proft, Hafniae. 382 pp.

——. 1794. Entomologia systematica emendata et aucta. Secundum classes, ordines, gen-
era, species adjectis synonymis, locis, observationibus, descriptionibus. Proft, Hafniae.
Vol. 3, part 2, 349 pp.

Felder, R. and A. F. Rogenhofer. 1875. Zweite Abtheilung: Lepidoptera. Atlas. Jn Felder,
C., R. Felder, and A. F. Rogenhofer, Reise der Osterreichischen Fregatte Novara um
die Erde in den Jahren 1857, 1858, 1859 unter den Behilfen des Commodore B. von
Wiillerstorf-Urbair. Zoologischer Theil. [part]. Vienna. 20 pp., pls. 108-140.

_ Ferro, D. N., A. D. Lowe, R. G. Ordish, K. G. Somerfield, and J. C. Watt. 1977. Standard

names for common insects of New Zealand. Bull. Entomol. Soc. N. Z. 4: 1-42.

_ Fischer von Roslerstamm, J. A. 1834-1843. Abbildungen zur Berichtigung und Erganzung der

Schmetterlingkunde, besonders der Microlepidopterologie als Supplement zu Treitschke’s

und Hubner’s europaeischen Schmetterlingen, mit erlauterndem Text. Leipzig. 308 pp.,

100 pls.

_ Fletcher, T. B. 1940. New generic names for microlepidoptera. Entomol. Rec. J. Var. 52: 17-
19, 109.

Frolich, G. F. von. 1828. Enumeratio tortricum, L., regno Wirttembergico indigenarum.
Tubingen. 102 pp.

Ghesquiere, J. 1940-42. Catalogue raisonnés de la faune entomologique du Congo Belge.
Lépidopteres, microlépidopteres. Ann. Mus. Congo Belge (Turvuren), Zool. (3) 7: 1-

240, 6 pls.
Gozmany, L. 1954. Studies on microlepidoptera. Ann. Hist.-Nat. Mus. Natl. Hung. 5: 273-
285.

Hampson, G. F. 1918. Some small families of the Lepidoptera which are not included in the
key to the families in the catalogue of Lepidoptera Phalaenae, a list of families and
subfamilies of the Lepidoptera with their types, and a key to the families. Novit. Zool.
25: 366-394.

Haworth, A. H. 1803-1828. Lepidoptera Britannica. J. Murray, London. 609 pp. [4 parts].

Heinemann, H. von and M. F. Wocke. [1876]. Die Schmetterlinge Deutschlands und der
Schweiz. Zweite Abtheilung. Kleinschmetterlinge. Band II: Die Motten und Federmot-
ten. Heft II. Schwetschke, Braunschweig. Pp. 389-825.

Heppner, J. B. 1977. The status of the Glyphipterigidae and a reassessment of relationships
in yponomeutoid families and ditrysian superfamilies. J. Lepid. Soc. 31: 124-134.

|) —. 198la. Revision of the new genus Diploschizia (Lepidoptera: Glyphipterigidae) for

) North America. Fla. Entomol. 64: 309-336.

| —. 1981lb. Neomachlotica, a new genus of Glyphipterigidae (Lepidoptera). Proc. Ento-

mol. Soc. Wash. 83: 479-488.

. In press a. Review of the family Immidae (Immoidea) with a world checklist. J. Lepid.

Soc.

. In press b. A world catalog of genera associated with the Glyphipterigidae auctorum

(Lepidoptera: Copromorphoidea). J. N. Y. Entomol. Soc.

58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Heppner, J. B. and W. D. Duckworth. 1981. Classification of the superfamily Sesioidea (Lep-
idoptera: Ditrysia). Smithson. Contrib. Zool. 314: 1-144.

Hering, M. 1927. Die Minenfauna der Canarischen Inseln. Zool. Jahrb. Abt. Syst. Okol.
Geogr. Tiere 53: 405-486.

Herrich-Schaffer, G. A. W. 1847-1855. Die Schaben und Federmotten. Jn Systematische
Bearbeitung der Schmetterlinge von Europa, zugleich als Text, Revision und Supple-
ment zu Jakob Hiibner’s Sammlung europaischer Schmetterlinge. Vol. 5. G. J. Manz,
Regensburg. 394 + 52 pp., 132 pls.

Heydenreich, G. H. 1851. Lepidopterorum europaeorum catalogus methodicus. Systematisch-
es Verzeichniss der europaeischen Schmetterlinge. J. Klinkhardt, Leipzig. 130 pp.

Hiibner, J. 1796-[1838]. Sammlung europaischer Schmetterlinge. Augsburg. 9 pts.

. 1816-[1825]. Verzeichniss bekannter Schmettlinge [sic]. Augsburg. 431 pp.

Issiki, S. T. 1930. New Japanese and Formosan microlepidoptera. Ann. Mag. Nat. Hist. (10)
6: 422-431.

Jordan, K. 1886. Die Schmetterlingsfauna nordwest-Deutschlands, insbesondere die lepi-
dopterologischen Verhaltnisse der Umgebung von Gottingen. Zool. Jahrb. Zeitsch. Syst.
Geogr. Biol. Thiere, Suppl. 1: 1-164.

Klemensiewicz, S. 1907. O nowychi malo znanych gatunkach motyli fauny galicyjskiej. Spra-
wozd. Komis. Fizyograf. PAU (Warsaw) 40: 3-31.

Kodama, T. 1961. The larvae of Glyphipterygidae (Lepidoptera) in Japan (I). Osaka Fac.
Agric., Entomol. Lab. 6: 35-45.

Le Marchand, S. 1925. Glyphipteryx heptaglyphella, n. sp. Amat. Papill. 2: 314-316.

Lower, O. B. 1893. New Australian Lepidoptera. Trans. R. Soc. S. Aust. 17: 146-185.

. 1897. Descriptions of new species of Australian Lepidoptera, with notes on synonymy.

Proc. Linn. Soc. N.S.W. 22: 10-32.

. 1905. New Australian Lepidoptera No. 22. Trans. R. Soc. S. Aust. 29: 103-115.

Mann, J. 1854. Aufzahlung der Schmetterlinge, gesammelt auf einer Reise im Auftrag des k.
k. zoologischen Museums nach Oberkrain und dem Kiistenlande, in den Monaten Mai
und Juni 1854, als Beitrag zur Fauna des Osterreichischen Kaiserstaates. Verh. Zool.-
Bot. Ver. Wien 4: 545-596.

Mann, J. and A. F. Rogenhofer. 1878. Zur Lepidopteren-Fauna des Dolomiten-Gebietes.
Verh. Zool.-Bot. Ges. Wien 27: 491-500.

Matsumura, S. 1931. 6000 illustrated insects of Japan-Empire. Tokyo. 1496 + 191 pp.

Meyrick, E. 1881. Descriptions of Australian micro-lepidoptera. IV: Tineina. Proc. Linn. Soc.
N.S.W. 5: 204-271.

—. 1883. Descriptions of Australian micro-lepidoptera. VII: Revisional. Proc. Linn. Soc.
N.S.W. 7: 148-202.

—. 1888. Descriptions of New Zealand Tineina. Trans. N. Z. Inst. 20: 77-106.

—. 1907a. Descriptions of Australasian micro-lepidoptera. XIX. Plutellidae. Proc. Linn.
Soc. N.S.W. 32: 47-150.

—. 1907b. Descriptions of Indian micro-lepidoptera. III. J. Bombay Nat. Hist. Soc. 17:

730-754.

——. 1907c. Descriptions of Indian micro-lepidoptera. IV. J. Bombay Nat. Hist. Soc. 17:
976-994.

—. 1907d. Descriptions of Indian micro-lepidoptera. V. J. Bombay Nat. Hist. Soc. 18:
137-160.

—. 1909a. Notes and descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 41:
5-16.

—. 1909b. Descriptions of micro-lepidoptera from Bolivia and Peru. Trans. Entomol.
Soc. Lond. 1909: 13-43.

VOLUME 84, NUMBER | 59

_ 1909c. Descriptions of Indian micro-lepidoptera. IX. J. Bombay Nat. Hist. Soc. 19:
410-437.

_ 1909d. New South African microlepidoptera. Ann. South Afr. Mus. 5: 349-379.

. 1910. Lepidoptera from the Kermadec Islands. Trans. N. Z. Inst. 42: 67-73.

_ 191la. Notes and descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 43:
58-78.

_ 1911b. Tortricina and Tineina. Jn Reports of the Percy Sladen Trust Expedition to
the Indian Ocean in 1905, under the leadership of My. J. Stanley Gardiner, M. A. Vol.
) 3. Trans. Linn. Soc. Lond. Zool. (2) 14: 263-307.

. 1912. Glyphipterygidae. Exotic Microlepid. 1: 35-63.

. 1913a. Glyphipterygidae. Exotic Microlepid. 1: 67—70, 98-104.

. 1913b. Carposinidae, Heliodinidae, Glyphipterygidae. Jn Wagner, H.. ed., Lepidop-
i! terorum catalogus. Part 13. W. Junk, Berlin. 53 pp.

_ 1914a. Descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 46: 101-1 18.

. 1914b. Glyphipterygidae. Exotic Microlepid. 1: 283-284.

. 1914c. Lepidoptera Heterocera. Fam. Glyphipterygidae. Jn Wytsman, P., ed., Genera
insectorum. Fasc. 164. Brussels. 39 pp., 2 pls.

_ 1915a. Descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 47: 201-204.

|, ———. 1915b. Descriptions of South American micro-lepidoptera. Trans. Entomol. Soc.
li Lond. 1915: 201-256.

_ 1916. Descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 48: 414-419.
1917. Descriptions of South African micro-lepidoptera. Ann. South Afr. Mus. 17: l-

_ 1918a. Descriptions of South African micro-lepidoptera. Ann. Transvaal Mus. 6: 7—

. 1918b. Glyphipterygidae. Exotic Microlepid. 2: 191-196.
. 1920a. Glyphipterygidae. Exotic Microlepid. 2: 325-338.
———. 1920b. Descriptions of South African micro-lepidoptera. Ann. South Afr. Mus. 17:
273-318.
_ 1921a. New microlepidoptera. Zool. Meded. (Leiden) 6: 145-202.
_ 1921b. Descriptions of South African micro-lepidoptera. Ann. Transvaal Mus. 8: 49-
148.
. 1922. Glyphipterygidae. Exotic Microlepid. 2: 481-493.

1923a. Voyage du Comte Jacques de Rohan-Charot dans |’Angola. Description
d’éspeces nouvelles de microlépidopteres. Bull. Mus. Natl. Hist. Nat. (Paris) 29: 563-
566.
———. 1923b. Glyphipterygidae. Exotic Microlepid. 2: 616-619.
————. 1925. Glyphipterygidae. Exotic Microlepid. 3: 132-136.
—_. 1926a. Glyphipterygidae. Exotic Microlepid. 3: 301-308.
__ 1926b. New South African micro-lepidoptera. Ann. South Afr. Mus. 23: 325-351.
———. 1928. Glyphipterygidae. Exotic Microlepid. 3: 420-424.
——. 1929. The micro-lepidoptera of the ‘‘St. George’” Expedition. Trans. Entomol. Soc.

Lond. 76: 489-521.
. 1930a. Hyponomeutidae [sic]. Exotic Microlepid. 3: 591-598.
. 1930b. Glyphipterygidae. Exotic Microlepid. 4: 1-7.
. 1931. Glyphipterygidae. Exotic Microlepid. 4: 180-185.
. 1932a. Glyphipterygidae. Exotic Microlepid. 4: 273-276.
1932b. Entomological expedition to Abyssinia, 1926-7. Microlepidoptera. Trans.
Entomol. Soc. Lond. 80: 107-120.
———. 1933. Glyphipterygidae. Exotic Microlepid. 4: 370-372.
1934a. Pterophoridae-Tortricidae-Tineidae. Jn Caradja, A. von and E. Meyrick, Ma-

60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

terialien zu einer Microlepidopteren-Fauna Kwangtungs. Dtsch. Entomol. Z. Iris 48:
28-43.

——. 1934b. Glyphipterygidae. Exotic Microlepid. 4: 523-524.

—_——. 1935. List of microlepidoptera of Chekiang, Kiangsu and Hunan. Jn Caradja, A. von

and E. Meyrick, Materialien zu einer Microlepidopteren Fauna der chinesischen Prov-

incen Kiangsu, Chekiang und Hunan. Friedlander, Berlin. Pp. 44-96.

. 1937. Glyphipterygidae. Exotic Microlepid. 5: 123-126.

Milliere, P. 1854. Description de nouvelles éspeces de microlépidopteres. Ann. Soc. Entomol.
Fr. (3) 2: 59-68.

Moriuti, S. and T. Saito. 1964. Glyphipterix semiflavana Issiki and the allied new species
from Japan (Lepidoptera: Glyphipterigidae). Entomol. Rev. Japan 16: 60-63, pl. 9-10.

Moschler, H. B. 1866. Aufzahlung der in Andalusien 1865 von Herrn Graf v. Hoffmannsegg
gesammelten Schmetterlinge. Berlin. Entomol. Z. 10: 136-146.

Newman, E. 1856. Characters of a few Australian Lepidoptera, collected by Mr. Thomas R.
Oxley. Trans. Entomol. Soc. Lond. 1855: 281-300.

Nowicki, M. S. 1864. Microlepidopterorum species novae. Jagellon Univ., Krakow. 31 pp.,
1 pl.

Philpott, A. 1924. Notes and descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst.
55: 207-214.

—. 1927. Notes and descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 58: 80—
90.

—. 1928. Notes and descriptions of New Zealand Lepidoptera. Trans. N. Z. Inst. 58:
359-370.

Prohaska, K. 1905. Beitrag zur Microlepidopteren-Fauna von Steirmark und Karnten. Jah-
resber. K. K. Ersten Staats-Gymnas. (Graz) 1905: 1-29.

Pryer, W. B. 1877. Descriptions of new species of Lepidoptera from North China. Cist.
Entomol. 2: 231-235, pl. 4.

Ragonot, E. L. 1885. [Contribution]. Bull. Soc. Entomol. Fr. 1885: lvi.

Rebel, H. 1896. Dritter Beitrag zur Lepidopteren-fauna der Canaren. Ann. Naturhist. Hofmus.
(Vienna) 11: 102-148, pl. 3.

—. 1916. Die Lepidopterenfauna Kretas. Ann. Naturhist. Hofmus. (Vienna) 30: 66-172,
pl. 4.

———. 1940. Die Lepidopterenfauna des Azorischen Archipels. Mit 1. Anhang: eine Lepi-
dopteren-Ausbeute von Madeira. Comm. Biol. 8: 1-59, pls. 1-2.

Scopoli, J. A. 1763. Entomologia Carniolica exhibens insecta Carnioliae indigena et distributa
in ordines, genera, species, varietates. Methodo Linnaeana. Vienna. 419 pp., 43 pls.
{1972 reprint: Akademische Verlag, Graz].

Snellen, P. C. T. 1882. De Vlinders van Nederland. Microlepidoptera. Brill, Leiden. 2 vols.,
1196 pp., 14 pls.

Stainton, H. T. 1859. A manual of British butterflies and moths. Vol. II]. (Comprising the
slender-bodied and small moths). Van Voorst, London. 480 pp.

Stephens, J. F. 1829. The nomenclature of British insects; being a compendious list of such

species as are contained in the systematic catalogue of British insects, and forming a

guide to their classification, and c. and c. Baldwin and Cradock, London. 68 pp. [June

1, 1829].

1831-1835. Illustrations of British entomology; or, a synopsis of indigenous insects:
containing their generic and specific distinctions; with an account of their metamor-
phoses, times of appearance, localities, food, and economy, as far as practicable. Haus-
tellata. Vol. IV. London. 436 pp., pls. 33-40.

Toll, S. 1936. Untersuchung der Genitalien bei Pyrausta purpuralis L. und P. ostrinalis Hb.,
nebst Beschreibung 11 neuer Microlepidopteren-Arten. Ann. Mus. Zool. Polonici 11:
403-413, pls. 47-49.

VOLUME 84, NUMBER 1 61

Treitschke, F. 1833. Schaben. Geitschen. Jn Ochsenheimer, F., Die Schmetterlinge von Eu-
ropa. Vol. 9, pt. 2. Fleischer, Leipzig. 294 pp.

Turati, E. 1879. Contribuzione alla fauna Lepidotterologica Lombarda. Boll. Soc. Entomol.

Ital. 11: 153-208, pls. 7-8.

. 1930. Lepidotteri di Rodi. Mem. Soc. Entomol. Ital. 9: 196-214.

Turner, A. J. 1913. Studies in Australian micro-lepidoptera. Proc. Linn. Soc. N.S.W. 38: 174-

228.

. 1926. Studies in Australian Lepidoptera. Trans. R. Soc. S. Aust. 50: 120-155.

—. 1927. New and little-known Tasmanian Lepidoptera. Pap. R. Soc. Tasmania 1926:

119-164.

. 1932. New Australian Lepidoptera. Trans. R. Soc. S. Aust. 56: 175-196.

Viette, P. 1951. Quelques microlépidopteres de Madagascar. Mém. Inst. Sci. Madagascar
(Ser. A) 5: 337-346.

Walker, F. 1863. Tortricites & Tineites. Jn List of the specimens of lepidopterous insects in
the collection of the British Museum. Part 28. Br. Mus., London. Pp. 288-561.

— . 1864. Tineites. Jn ibid. Pt. 29-30. Br. Mus., London. Pp. 563-1096.

—. [1865]. Supplement. /n ibid. Pt. 31. Br. Mus., London. 321 pp.

—. 1869. Characters of undescribed Lepidoptera Heterocera. E. W. Janson, London. 112

Pp.

Walsingham, T. de G. 1881. On some North American Tineidae. Proc. Zool. Soc. Lond. 1881:

301-325, pls. 35-36.

. 1891. African micro-lepidoptera. Trans. Entomol. Soc. Lond. 1891: 63-132, pls. 3-7.

—. 1892. On the micro-lepidoptera of the West Indies. Proc. Zool. Soc. Lond. 1891: 492—
549.

— . 1897. Revision of the West-Indian micro-lepidoptera, with descriptions of new species.
Proc. Zool. Soc. Lond. 1897: 54-183.

—. 1908. Microlepidoptera of Tenerife. Proc. Zool. Soc. Lond. 1907: 911-1034, pls. 51-
DS:

—. 1910. Madeiran Tineina (Lepidoptera). Entomol. Mon. Mag. 46: 257-259.

—. 1914. Hemerophilidae. Jn Tineina, Pterophorina, Orneodina and Pyralidina and He-
pialina (part). Jn Godman, F. D. and O. Salvin, Biologia Centrali-Americana. Insecta.
Lepidoptera-Heterocera. Vol. 4. London. Pp. 300-319, pl. 9.

Watt, M. N. 1920. The leaf-mining insects of New Zealand. Trans. N. Z. Inst. 52: 439-466,
pl. 30.

Westwood, J. O. [1838]-1840. Synopsis of the genera of British insects. London. 158 pp.

Wollaston, T. V. 1879. Notes on the Lepidoptera of St. Helena, with descriptions of new
species. Ann. Mag. Nat. Hist. (5) 3: 219-233, 329-343, 415-441.

Zeller, P. C. 1839. Versuch einer naturgemassen Eintheilung der Schaben. Isis v. Oken 32:

167-220.

. 1850. Verzeichniss der von Herrn Jos. Mann beobachteten toscanischen Microlepi-

doptera. Entomol. Z. (Stettin) 11: 59-64, 134-136, 139-162, 195-212.

—. 1852. Lepidoptera Microptera, quae J. A. Wahlberg in Caffrorum terra collegit. Glo-

gau. 120 pp. [reissued in [1854]: Kongl. Svenska Vetensk. Akad. Handl. 1852: 1-120].

. 1877. Exotische Microlepidoptera. Hor. Soc. Entomol. Ross. 13: 3-493, 6 pls.

Zetterstedt, J. W. [1839]-1840. Insecta Lapponica. Voss., Lipsiae. 1140 pp.

Alphabetical List of Species

achlyoessa Meyrick, Phryganostola acronoma Meyrick, Glyphipterix
achlyoéssa Meyrick, Phryganostola acrothecta Meyrick, Glyphipterix
acinacella Meyrick, Glyphipterix actinobola Meyrick, Glyphipterix

62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

actinota (Walsingham), Neomachlotica

acyloessa [sic] (Ferro et al.), Phryganostola
achlyoessa

aechmiella (Duponchel), Glyphipterix

aereinitidella (Milliere), Glyphipterix

aerifera Meyrick, Glyphipterix

aglaograpta Meyrick, Irinympha

ahornerella (Hiibner), Glyphipteryx thra-
sonella

aillyella (Hiibner), Glyphipterix thrasonella

albardella Snellen, Glyphipterix thrasonella

albicostella (Duponchel), Glyphipterix fus-
coviridella

albimaculella Heinemann, Glyphipterix

alpha Moriuti and Saito, Glyphipterix

altiorella Bauer, Glyphipterix

amblycerella Meyrick, Glyphipterix

ametris Diakonoff, Glyphipterix

amphipeda Meyrick, Glyphipterix

amseli Diakonoff, Glyphipterix

anaclastis Meyrick, Glyphipterix

ancobathra Meyrick, Ussara

ancyristis Meyrick, Ussara

angulosella (Costa), Glyphipterix

antidoxa Meyrick, Glyphipterix

archimedica Meyrick, Glyphipterix

arcuatella Klemensiewicz, Glyphipterix
bergstraesserella

argophracta Meyrick, Glyphipterix

argyrelata Turner, Glyphipterix

argyroguttella Ragonot, Glyphipterix

argyromis Meyrick, Glyphipterix

argyrosema Meyrick, Apistomorpha

argyrotoxa Turner, Glyphipterix

arquata Meyrick, Ussara

asterias (Meyrick), Glyphipterix

asteriella Meyrick, Glyphipterix

asteronota Meyrick, Glyphipterix

astrapaea Meyrick, Glyphipterix

astrocosma Meyrick, Tetracmanthes

ataracta Meyrick, Phryganostola

atelura Meyrick, Glyphipterix

atractias (Meyrick), Neomachlotica

atristriella Zeller, Glyphipterix gemmipunc-
tella

aulogramma Meyrick, Glyphipterix

auripennis Boisduval, Myrsila

autopetes Meyrick, Glyphipterix

bactrias Meyrick, Glyphipterix

basifasciata Issiki, Glyphipterix

bergstraesella [sic] Bruand, Glyphipterix
bergstraesserella

bergstraessella [sic] (Fabricius), Glyphip-
terix bergstraesserella

bergstraesserella (Fabricius), Glyphipterix

bergstrdsserella [sic] Amsel, Glyphipterix
bergstraesserella

beta Moriuti and Saito, Glyphipterix

bicornis Meyrick, Glyphipterix

bifasciata Walsingham, Glyphipterix

bifasciella Amsel, Glyphipterix

bohemani (Zeller), Glyphipterix

brachyaula Meyrick, Glyphipterix

brachydelta Meyrick, Glyphipterix

californiae Walsingham, Glyphipterix

calliactis Meyrick, Glyphipterix

callicrossa Meyrick, Glyphipterix

callidelta Meyrick, Glyphipterix

calliscopa Lower, Glyphipterix

callithea Meyrick, Glyphipterix

canachodes Meyrick, Glyphipterix

carenota Meyrick, Glyphipterix

caudatella Walsingham, Glyphipterix

celophora (Meyrick), Taeniostolella

cestrota Meyrick, Glyphipterix

chalcodaedala Turner, Glyphipterix

chalceres Turner, Glyphipterix meteora

chalcodesma Meyrick, Ussara

chalcostrepta Meyrick, Glyphipterix

chalcotypa (Bradley), Chrysocentris

chionosoma Diakonoff, Glyphipterix

chlorospora Meyrick, Ernolytis

chrysallacta Meyrick, Glyphipterix

chrysangela Meyrick, Ussara

chrysodeta Meyrick, Machlotica

chrysolithella Meyrick, Glyphipterix gem-
mipunctella

chrysoplanetis Meyrick, Glyphipterix

chrysorma Meyrick, Phalerarcha

chrysosema (Meyrick), Carmentina

chrysozona (Meyrick), Chrysocentris

cionophora Meyrick, Circica

circumscripta [sic] Dyar, Glyphipterix cir-
cumscriptella

circumscriptella Chambers, Glyphipterix

cladiella Stainton, Glyphipterix thrasonella

clavaria Meyrick, Chrysocentris

clearcha Meyrick, Glyphipterix

climacaspis Meyrick, Glyphipterix

codonias Meyrick, Glyphipterix

cognatella (Stephens), Glyphipterix simpli-
ciella

colluripennella (Costa), Glyphipterix sim-
pliciella

I ee

VOLUME 84, NUMBER |

colorata Meyrick, Glyphipterix

columnaris Meyrick, Glyphipterix

cometophora Meyrick, Glyphipterix

compastis Meyrick, Glyphipterix

conjunctella (Costa), Glyphipterix

conosema Meyrick, Glyphipterix

cornigerella Zeller, Glyphipterix

cramerella (of authors), Glyphipterix sim-
pliciella

crassilunella Rebel, Glyphipterix

crinita Meyrick, Glyphipterix

crotalotis Meyrick, Glyphipterix

cultrata Meyrick, Glyphipterix

cuprea Busck, Abrenthia

cyanophracta Meyrick, Glyphipterix

cyanochalca Meyrick, Glyphipterix

danilevskii Diakonoff, Glyphipterix

decachrysa Meyrick, Glyphipterix

decoratella Walker, Ussara

deliciosa Diakonoff, Glyphipterix

delta Moriuti and Saito, Glyphipterix

deltodes (Walsingham), Cronicombra

desideratella [sic] (Duponchel), Glyphipte-
rix simpliciella

desiderella (ROslerstamm), Glyphipterix
simpliciella

deuterastis Meyrick, Glyphipterix

diaphora Walsingham, Glyphipterix

dichalina Meyrick, Glyphipterix

dichorda Meyrick, Glyphipterix

diplotoxa Meyrick, Glyphipterix

ditiorana (Walker), Chrysocentris

dolichophyes Diakonoff, Glyphipterix

drosophaes Meyrick, Phryganostola

enclitica Meyrick, Glyphipterix

epastra Meyrick, Glyphipterix

equitella (of authors), Glyphipterix mino-
rella

equitella (Scopoli), Glyphipterix

equitella (Treitschke), Glyphipterix thraso-
nella

erastis Meyrick, Glyphipterix

erebanassa Meyrick, Glyphipterix

essedaria Meyrick, Cronicombra

euastera Meyrick, Glyphipterix

euleucotoma Diakonoff and Arita, Glyphip-
terix

eumitrella (Busck), Phalerarcha

eupepla Meyrick, Chrysocentris

eurymolybda Meyrick, Machlotica

eurythmiella Busck, Ussara

euthybelemna Meyrick, Phryganostola

63

exoptatella (Chambers), Diploschizia impi-
gritella

expurgata Meyrick, Glyphipterix

eyndhoveniella Snellen, Glyphipterix thra-
sonella

falcigera Meyrick, Glyphipterix

fiesiella (Westwood), Glyphipterix thraso-
nella

fischeriella (Zeller), Glyphipterix simplici-
ella

flaviceps (Felder and Rogenhofer), Ussara

forsterella (Fabricius), Glyphipterix

fortunatella Walsingham, Glyphipterix

fueslella (Fabricius), Glyphipterix thraso-
nella

fueslii (Haworth), Glyphipterix thrasonella

fulgens (Erschoff), Lepidotarphius peror-
natella

funditrix Diakonoff and Arita, Glyphipterix

fuscoviridella (Haworth), Glyphipterix

fuselella [sic] (Zeller), Glyphipterix thraso-
nella

fyeslella (Fabricius), Glyphipterix (ibid.)

gamma Moriuti and Saito, Glyphipterix

gaudialis Diakonoff and Arita, Glyphipterix

gemmatella (Walker), Glyphipterix

gemmipunctella (Walker), Glyphipterix

gemmula Diakonoff and Arita, Glyphipterix

gemmula Walsingham, Trapeziophora

gianelliella Ragonot, Glyphipterix

glaucophanes (Meyrick), Diploschizia

gonoteles Meyrick, Glyphipterix

granulata Meyrick, Cronicombra

grapholithoides Walsingham, Glyphipterix

gypsonota Turner, Glyphipterix

habecki Heppner, Diploschizia

halimophila Lower, Glyphipterix

haplographa Turner, Glyphipterix

harpogramma Turner, Glyphipterix

haworthana (Stephens), Glyphipterix

haworthella (Stephens), Glyphipterix ha-
worthana

hemipempta Meyrick, Glyphipterix

heptaglyphella Le Marchand, Glyphipterix

hilarodes (Meyrick), Ussara

holochalca Meyrick, Pantosperma

holodesma Meyrick, Glyphipterix

hologramma Meyrick, Glyphipterix

howarthana [sic] Jordan, Glyphipterix ha-
worthana

humerella (Hubner), Glyphipterix forster-
ella

64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

hyperlampra Turner, Glyphipterix

idiomorpha Meyrick, Glyphipterix

iinvicta [sic] Clarke, Glyphipterix invicta

imparfasciata Arita, Glyphipterix

impigritella (Clemens), Diploschizia

indomita Meyrick, Glyphipterix

infuscata Ghesquiere, Chrysocentris

invicta Meyrick, Glyphipterix

iocheaera Meyrick, Glyphipterix

iochrysa Meyrick, Ussara

ioclista Meyrick, Glyphipterix

iometalla Meyrick, Glyphipterix

iridesma Meyrick, Carmentina

isoclista Meyrick, Glyphipterix

isozela Meyrick, Glyphipterix

japonicella Matsumura, Glyphipterix beta

japonicella Zeller, Glyphipterix

jezonica Matsumura, Glyphipterix basifas-
ciata

kimballi Heppner, Diploschizia

lamella (Busck), Cronicombra

lamprocoma Meyrick, Glyphipterix

lamprosema Turner, Glyphipterix

lanista (Meyrick), Diploschizia

leptocona Meyrick, Glyphipterix

leptosema Meyrick, Glyphipterix

leucargyra Turner, Glyphipterix

leucocerastes Meyrick, Glyphipterix

leucophragma Meyrick, Glyphipterix

leucoplaca Turner, Glyphipterix

lineana (Hubner), Glyphipterix bergstraes-
serella

lineella Hubner, Glyphipterix (ibid.)

linneana [sic] Hiibner, Glyphipterix (ibid.)

linneella Hubner, Glyphipterix (ibid.)

litura (Felder and Rogenhofer), Taeniosto-
lella

longistriatella Rebel, Glyphipterix

loricatella (Treitschke), Glyphipterix

loricutella [sic] Desmarest, Glyphipterix
loricatella

lucasella (Duponchel), Glyphipterix forster-
ella

luteocapitella Caradja, Glyphipterix

luteomaculata Arita, Glyphipterix

lycnophora Turner, Glyphipterix

lyelliana Lower, Glyphipterix cyanochalca

macrantha Lower, Phryganostola

macraula Meyrick, Glyphipterix

macrodrachma Diakonoff, Glyphipterix

madagascariensis Viette, Glyphipterix

magnatella Erschoff, Glyphipterix

maiorella [sic] Prohaska, Glyphipterix equi-
tella

majorella (Herrich-Schaffer), Glyphipterix
equitella

majorelia (Heydenreich), Glyphipterix equi-
tella

majorella Mann, Glyphipterix equitella

maritima Diakonoff, Glyphipterix

marmaropa Turner, Glyphipterix

maschalis Meyrick, Glyphipterix

mediana Walker, Cotaena

medianella Turati, Glyphipterix

medica Meyrick, Glyphipterix

melania Diakonoff and Arita, Glyphipterix

mesaula Meyrick, Glyphipterix

metasticta Meyrick, Glyphipterix

meteora Meyrick, Glyphipterix

metron Diakonoff, Glyphipterix

metronoma Meyrick, Glyphipterix

minimella Heppner, Diploschizia

minorella Snellen, Glyphipterix

molybdastra Meyrick, Glyphipterix

molybdora (Meyrick), Carmentina

molybdotoma (Diakonoff and Arita), Carmen-
tina

monodonta Diakonoff, Glyphipterix

montella Meyrick, Glyphipterix montisella

montinella [sic] Chambers, Glyphipterix
montisella

montisella Chambers, Glyphipterix

morangella Felder and Rogenhofer, Gly-
phipterix

nattani Gozmany, Glyphipterix

nebras (Meyrick), Neomachlotica

necopina Philpott, Glyphipterix

neochorda Meyrick, Glyphipterix

nephoptera Meyrick, Glyphipterix

nicaeella Moschler, Glyphipterix

nigromarginata Issiki, Glyphipterix

nitens Bankes, Glyphipterix thrasonella

nivicaput Diakonoff, Glyphipterix forster-
ella

nugella Felder and Rogenhofer, Glyphipte-
rix

octatoma Diakonoff, Glyphipterix

octonaria Philpott, Glyphipterix

oculatella Mann, Glyphipterix forsterella

oculatella (Zeller), Glyphipterix forsterella

okui Diakonoff and Arita, Glyphipterix

oligastra Meyrick, Glyphipterix

olyranta Meyrick, Ussara

orthodeta Meyrick, Glyphipterix

VOLUME 84, NUMBER 1

ortholeuca Meyrick, Glyphipterix
orthomacha Meyrick, Glyphipterix
orymagdis Meyrick, Glyphipterix
oxycopis Meyrick, Glyphipterix
oxydonta Diakonoff, Glyphipterix
oxyglypta (Meyrick), Glyphipterix
oxymachaera Meyrick, Phryganostola
oxytricha Meyrick, Glyphipterix
palaeomorpha Meyrick, Glyphipterix
palpella (Walsingham), Cronicombra
paradisea Walsingham, Glyphipterix
parazona Meyrick, Glyphipterix
_ paurographella Ragonot, Glyphipterix ar-
gyroguttella
perculta (Diakonoff), Carmentina
perfracta Meyrick, Glyphipterix
perimetalla Lower, Glyphipterix
pernicipennella (Costa), Glyphipterix angu-
losella
perornatella (Walker), Lepidotarphius
_ persica Diakonoff, Glyphipterix
pertenuis Diakonoff, Glyphipterix
phaeobathra (Meyrick), Cronicombra
phaeometalla Meyrick, Chrysocentris
| pharetropis Meyrick, Glyphipterix
phosphora Meyrick, Glyphipterix
| pietruskii Nowicki, Glyphipterix bergstraes-
serella
| plagiographa Bradley, Glyphipterix
| platydisema Lower, Glyphipterix
platyochra Meyrick, Glyphipterix
plenella (Busck), Cotaena
poeyella (Duponchel), Glyphipterix thraso-
nella
polyastra Meyrick, Ussara
polychroa Lower, Glyphipterix
| polychrysa (Meyrick), Carmentina
| polyzela Meyrick, Glyphipterix
porphyrospila (Meyrick), Cronicombra
‘protomacra Meyrick, Glyphipterix
pseudostoma Meyrick, Glyphipterix
psychopa Meyrick, Glyphipterix
pygmaeella Rebel, Glyphipterix
pyristacta Turner, Glyphipterix
pyrogastra Meyrick, Glyphipterix
pyrophora Turner, Glyphipterix
quadragintapunctata Dyar, Glyphipterix
trefractella Zeller, Glyphipterix
regia Heppner, Diploschizia
regula Diakonoff and Arita, Glyphipterix

|

65

repletana (Walker), Glyphipterix

rhanteria Turner, Glyphipterix

rhinoceropa Meyrick, Glyphipterix

rhodanis Meyrick, Glyphipterix

roeslerstammella (Fischer v. Roslerstamm),
Glyphipterix simpliciella

rugata Meyrick, Glyphipterix

sabella Newman, Glyphipterix

saurodonta Meyrick, Glyphipterix

schoenicolella Boyd, Glyphipterix

schultzella Amsel, Glyphipterix

scintilla Clarke, Glyphipterix

sclerodes Meyrick, Glyphipterix

scolias Meyrick, Glyphipterix

semicoronis Meyrick, Ussara

semiflavana Issiki, Glyphipterix

semifravana [sic] Kodama, Glyphipterix
semiflavana

semilunaris Wollaston, Glyphipterix

semisparsa Meyrick, Glyphipterix

seppella (Fabricius), Glyphipterix

seppella (Hiibner), Glyphipterix thrasonella

septemstrigella Zeller, Glyphipterix

sexguttella Toll, Glyphipterix

similis Philpott, Glyphipterix

simplicella Christoph, Glyphipterix

simpliciella (Stephens), Glyphipterix

speculans Meyrick, Glyphipterix

speculiferella Christoph, Glyphipterix

spiraea Heppner, Neomachlotica

splendens Pryer, Lepidotarphius perorna-
tella

sporomantis Meyrick, Rhabdocrates

stasichlora Meyrick, Glyphipterix

stelucha Meyrick, Glyphipterix

stilata Meyrick, Glyphipterix

struvei Amsel, Glyphipterix

sulcosa Diakonoff, Glyphipterix

suzukii Matsumura, Glyphipterix nigromar-
ginata

synarma Meyrick, Glyphipterix

syndecta Meyrick, Glyphipterix

synorista Meyrick, Glyphipterix

talhouki Diakonoff, Glyphipterix

tetrachrysa Meyrick, Glyphipterix

tetrasema Meyrick, Glyphipterix

tetratoma (Meyrick), Diploschizia

thiolychna Meyrick, Electrographa

thrasonella (Scopoli), Glyphipterix

thravonella [sic] Meyrick, Glyphipterix thra-
sonella

transversella (Walker), Glyphipterix

66 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

treitschkeana (Froélich), Glyphipterix bergs-
traesserella

trigonaspis Meyrick, Glyphipterix

trigonodes Arita, Glyphipterix

triguttella (Donovan), Glyphipterix thraso-
nella

tripedila Meyrick, Glyphipterix

triplaca Meyrick, Glyphipterix “‘triplaca”’

tripselia Meyrick, Glyphipterix

triselena Meyrick, Glyphipterix

tungella Felder and Rogenhofer, Glyphip-
terix

umbilici Hering, Glyphipterix

uncta Meyrick, Glyphipterix

unguifera Meyrick, Glyphipterix

unifasciata Walsingham, Glyphipterix

urania Meyrick, Chrysocentris

urophora (Walsingham), Diploschizia

variata Meyrick, Glyphipterix

variella Zeller, Glyphipterix simpliciella

versicolor Meyrick, Glyphipterix

voluptella Felder and Rogenhofer, Glyphip-
terix

xanthoplecta Meyrick, Glyphipterix

xestobela Meyrick, Circica

xyridota Meyrick, Glyphipterix

xyridota Meyrick, Glyphipterix dolicho-
phyes

zalodisca Meyrick, Glyphipterix

zelota Meyrick, Glyphipterix

zermattensis Amsel, Glyphipterix

zonella (Zetterstedt), Glyphipterix hawor-
thana

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 67-80

| AN ADDITIONAL STUDY ON THE NESTING BEHAVIORS OF
SPECIES OF MISCOPHUS (HYMENOPTERA: SPHECIDAE)

FRANK E. KURCZEWSKI

| Department of Environmental and Forest Biology, S.U.N.Y. College of
| Environmental Science and Forestry, Syracuse, New York 13210.

Abstract.—Additional observations made on nesting females of Misco-
_phus (Miscophus) americanus Fox, M. (Nitelopterus) slossonae barberi
| (Krombein), and M. (Nitelopterus) s. slossonae (Ashmead) in New York
_and Florida extend the range in variation in their ecologies and nesting
' behaviors. Miscophus americanus females nested in sand cliffs, left en-
trances open and oriented and brought prey to the nests in flight. Their nests
/ were short and simple and were stored with 3-14 tiny, mostly Theridion
\ spp. (Theridiidae). Miscophus slossonae barberi nested in flat sand, usually
left entrances open and oriented by running around the entrances, and trans-
| ported prey on the ground or in short, low flights. Their longer, simple nests
| were stocked with 5-14 tiny lycosids and salticids. Miscophus s. slossonae
| constructed the longest and deepest nests of the three taxa, and stored 7-
8 tiny spiders of a variety of families (Theridiidae, Micryphantidae, Oxy-
} opidae, Lycosidae, Gnaphosidae, Salticidae). This subspecies nested in flat
| sand, closed nest entrances or left them open during provisioning, oriented
| in flight or on the ground, transported prey in low, short flights or on the
| ground, and often took longer than a day to complete a nest.

I
|

In 1969 I summarized much of the known biological and behavioral in-
formation for the Nearctic and exotic Miscophus, including species in the
subgenera Miscophus and Nitelopterus. Bohart and Menke (1976) encap-
|} suled this information in three paragraphs, and Krombein (1979) summa-
rized the known ecological data, prey records, and biological references for

species of Miscophus. Neither compilation added much new information to
the existing pool of data on the nesting behaviors of species of Miscophus

|

simply because no studies of species in this genus were published during
the 1970's except for a note on Nitelopterus (=Miscophus) evansi Krom-
bein by Evans (1970). The intent of the present paper is to provide new
information on species in both Nearctic subgenera in order to extend the
range of biological knowledge for this little-studied group of wasps.

68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The species of Miscophus which are treated in this paper, their localities,
dates of observation and collection, and associated ethological note numbers
are as follows: M. (Miscophus) americanus Fox—Selkirk Shores St. Pk.,
Oswego Co., N.Y., 14 June 1971-22 August 1972 (MI 3-6, 8-13); Arcadia,
DeSoto Co.. Fla.. 27 March 1972 (MI-7); M. (Nitelopterus) slossonae bar-
beri (Krombein)—Archbold Biol. Sta., L. Placid, Highlands Co., Fla., 15
March-6 April 1973 (NI-21-24); M. (Nitelopterus) s. slossonae (Ashmead)—
Arcadia, DeSoto Co., Fla., 8 April 1971-28 March 1973 (NI-17-20, 25). The
wasps and prey have been deposited in the insect and invertebrate collec-
tions of the S.U.N.Y. College of Environmental Science and Forestry, Syr-
acuse.

Miscophus (Miscophus) americanus Fox

Eleven nests were constructed into vertical sand cliffs. The nest at Ar-
cadia, Fla., was situated halfway up a 2.4 m steep cliff, whereas the 10 nests
at Selkirk Shores St. Pk., N.Y., were located from 18 to 76 (x, 53) cm from
the top of a 1.8 m high cliff. Although I was unable to locate their nests,
provisioning females of M. americanus were seen flying to and from a 1 m
high sand cliff near the entrance to Presque Isle St. Pk., Erie Co., Pa.,
during 4-9 July 1971.

Several females began nests in shallow pre-existing depressions in the
cliff face or beneath overhanging rootlets in concealed situations. , Two such
nest entrances were only | cm apart, although the cells of these respective
nests were separated by 3 cm.

Burrow construction components conformed to those described by Kurc-
zewski (1969), although false starts were made less frequently when nesting
in sand cliffs than when nesting in flat sand. The duration of burrow con-
struction for four females averaged 35 (21-44) minutes.

All prey (32) of females of M. americanus brought to the nest were carried
in flight. Nesting in sand cliffs may restrict females to practice this method
of transport. Size of prey influenced the speed of flight but not type of
transport. The mechanisms of prey carriage were similar to those described
by Kurezewski (1969).

The period of time between 13 consecutive provisioning trips of two fe-
males averaged 10 (2-29) minutes. Neither size of prey nor position within
the sequence influenced the period of time between such trips. Seven larger
prey were taken into the open entrances, released 1-2 mm inside the bur-
rows, and pulled in from inside after the wasps had turned around, whereas
11 smaller prey were transported directly into open burrows without females
releasing the spiders. One female exited in flight from 3 to 30 (z, 8.4; N =
10) seconds after entering with prey. The time between consecutive exits
decreased steadily from 1130 hours (30 seconds) to 1326-1334 hours (3 sec-

VOLUME 84, NUMBER | 69

onds), perhaps a reflection of increased familiarity with the burrow confines
and/or increased sand surface temperature.

Upon taking the last prey into the nest females of M. americanus briefly
appeared headfirst inside the entrance, turned around, and reentered the
burrow. From 5 to 9 minutes later they were observed in their entrances
making nest closures. Details of the closure were as described by Kurczew-
ski (1969), except for extensive use of the mandibles to break down sand
from the walls of the burrows. Five closures were completed by females
between 1315 and 1605 hours. They averaged 9.9 (8-15) minutes in duration.
This closure activity culminated in burrows filled with loose sand for much
of their lengths from the fully provisioned cells to within 1-S mm of the
entrances.

Ten nests of M. americanus were short, simple (spurless) and unicellular.
Four nests exhibited some lateral curvature upon entering the cliff face,
whereas six went in rather straight. The burrows coursed inward almost
horizontally with most declining from 5 to 15° with the horizon. Nest en-
trances were 2.5—3.0 mm in diameter and the burrows, 2.0—2.5 mm. Burrow
lengths ranged from 2.0 to 3.8 (x, 2.8) cm, including cell length. The ellip-
soid-shaped cells were 4-6 mm wide, 4—6 mm high and 6-12 mm long.

Females of M. americanus preyed almost totally on spiders of the family
Theridiidae as follows: Theridion differens Emerton, 18 ad. 2, 1 pen. ¢; 19
imm.; 7. flavonotatum Becker, 2 ad. 2; T. murarium Emerton, 22 imm.;
T. redimitum (Linn.), 1 imm.; and, T. sp., 5 imm. In Florida one wasp
provisioned partly with four immature Araneus sp. (Araneidae), an excep-
tional record. Many of the prey were moving the legs and extruding silk
from the spinnerets.

Nine fully provisioned cells of M. americanus contained from 3 to 14 (x,
7.7) prey per cell. The individual spiders ranged in weight from 0.5 to 5.9
(x, 1.9; N = 67) mg. Four captor wasps averaged 4.4 (3.0-5.7) mg in weight.
The contents of eight fully provisioned cells ranged in weight from 8.0 to
20.0 (%, 14.7) mg.

The prey of M. americanus were placed variously in the cells as follows:
ventral side upward, head inward, 15; dorsal side upward, head inward, 26;
dorsal side upward, head outward, 13; sideways, head upward or down-
ward, 8. No prey were placed ventral side upward and head outward. There
was no particular pattern as to the exact positioning of these spiders within
the cell, i.e., individuals positioned dorsal side upward and head inward
were found at the back or front ends of the cell or toward the center.

The egg of M. americanus was attached to the spider as described and
figured by Kurczewski (1969). Eggs measured 1.25-1.50 x 0.40-0.45 mm.
In eight cells four eggs were attached to the base of the left side of the
spider’s abdomen and four to the base of the right side. The egg-bearing

70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

spiders were placed in the cells either dorsal side upward and head inward
(6) or dorsal side upward and head outward (2). They were positioned in
the cells near the back (3), middle (2), or front end (3). Spiders with eggs
weighed 1.8—5.9 (%, 3.2) mg and were the largest (3), second largest (3), third
largest (1), or second smallest (1) individuals in the cells. In one cell there
was an egg of the same size and shape as that of M. americanus on the floor
in addition to the egg attached to the spider!

Miscophus (Nitelopterus) slossonae barberi Krombein

Three nests were excavated and examined in flat, bare, white sand at the
Archbold Biological Station. Several other females were observed nesting
in flat or sloping, bare, white sand near Lakes Annie and Placid, Highlands
Co., Fla., during March and April 1973, but data were not gathered on these
individuals.

Females searching for nesting sites made numerous false starts, especially
in the morning between 1100 and 1200 hours (E.S.T.). Components of bur-
row construction were similar to those described by Krombein and Kurc-
zewski (1963) and Kurczewski (1969). One wasp started a burrow in a
depression in bare sand and completed it | hour and 58 minutes later. Fe-
males completing burrows exited headfirst, turned 180°, walked back and
forth on or around the tumulus, entered and exited from the burrow from
one to four times, and then began walking increasingly farther away from
the entrance in circular or zig-zag routes, an apparent form of orientation.
These movements increased in rapidity until the females flew off to hunt.

Prey transport involved a female grasping the spider with the mandibles
by its forelegs or pedipalps, head forward and ventral side upward, and
walking rapidly forward on the ground or making short, ‘“‘hopping’’ flights
using the wings. Several prey were released and abandoned when the pro-
visioning wasps were attacked by foraging ants. Such females returned to
their nests and immediately closed the nest entrances with sand. Some
wasps periodically returned to their nests without prey, examined the open
entrance and flew off or entered the burrow, exited several seconds later
and flew away.

The size of prey influenced the manner in which the provisioning wasp
entered her open nest. Females with 16 ‘‘smaller’’ prey went directly into
their open nests without releasing the spiders, whereas wasps with 9
“‘larger’’ prey released the spiders just inside the entrances, went into the
burrows, turned around and pulled the prey inside. Such wasps spent an
average of 19 (5-42; N = 15) minutes between consecutive prey. These
females reappeared in their entrances from 10 to 180 (¢, 49; N = 14) seconds
after placing their prey in the nests. The longest periods of time were re-
corded near the end of the provisioning sequence.

Components of final closing the nest were essentially identical to those

VOLUME 84, NUMBER 1 7

described for the nominate subspecies by Kurczewski (1969). Three final
closures averaged 38 (27-46) minutes in duration. Closed nests contained
burrows completely filled with sand. One female attempted to bring prey
into a burrow that had been filled with sand and contained prey with one of
the spiders bearing a wasp’s egg!

The three nests of M. slossonae barberi contained straight or slightly
curved burrows with diameters of 2 mm and ended in single cells. These
burrows ranged from 4.9 to 6.1 (¥, 5.3) cm in length and from 2.9 to 3.9 (x,
3.5) cm in depth, including cell length and height, respectively. The cells
averaged 3.5 (3-4) mm high, 3.5 (3-4) mm wide and 5.5 (S—6) mm long. The
diameter of the burrow entrance in all three nests was 2.5 mm.

The females preyed upon tiny wolf and jumping spiders as follows: LY-
COSIDAE: Pirata sp., 1 imm.; Schizocosa sp., 1 imm. SALTICIDAE:
Habrocestum sp. nr. pulex (Hentz), 2 ad. 2,6 ad. d, | imm. d, 15 imm.
The spiders were incompletely paralyzed, moved their legs, and extruded
silk from the spinnerets.

The three fully provisioned cells contained from 5 to 14 (x, 8.3) prey per
cell. The spider individuals ranged in weight from 0.2 to 2.8 (¥, 1.2; N =
26) mg. One wasp weighed 2.2 mg. The cell contents ranged in weight from
4.3 to 15.4 @, 9.2) mg.

The prey of M. slossonae barberi were placed in various positions within
the cells as follows: ventral side upward, head inward, 9; dorsal side up-
ward, head inward, 11; dorsal side upward, head outward, 3; and, sideways,
head upward or downward, 2. None of the prey were positioned ventral
side upward and head outward.

The egg of this subspecies was affixed to the spider’s abdomen as indi-
cated and figured for M. americanus by Kurczewski (1969). Eggs measured
from 1.2 to 1.8 mm in length. All three eggs were affixed to the left sides of
the spiders’ abdomens and the three spiders with eggs were positioned dor-
sal side upward and head inward in the cells. The three spiders were located
at the front or back end or in the middle of the cell. Egg-bearing spiders
weighed from 1.4 to 2.2 (¥, 1.9) mg. They were the largest prey in each of
the three cells.

Miscophus (Nitelopterus) s. slossonae (Ashmead)

Four nests were constructed into the white sand flats bordering the Peace
River, west of Arcadia, Fla. One nest was dug into the slope of an ant-lion
pit, near the top, another was concealed beneath two overhanging grass
blades, a third into the edge of a cow’s hoofprint, and a fourth beneath an
overhanging Mimosa plant. A fifth wasp attempted to dig in a depression
caused by a marking stake. Only one female attempted to nest in bare sand,
devoid of vegetation or depressions.

Burrow construction components were identical to those described by

72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Kurczewski (1969). One wasp searched for only 10 minutes before remaining
in one place and completing her burrow. She took 50 minutes to completion,
beginning at 1159 and ending at 1249 hours (E.S.T.). She removed sand
from the burrow to a distance of 20 mm from the entrance. The resulting
tumulus was elongate-ovoidal in shape.

One wasp made a 2-second-long temporary closure of the nest entrance
after completing the burrow, after which the entrance remained one-third
open at the top. She then ran around the area of the entrance for 30 seconds
and flew away. This female returned from her hunting excursions several
times but never entered the burrow. Two other females made no closures
of their entrances while obtaining a total of six prey. Another wasp, how-
ever, after leaving the nest open during her first two provisioning trips made
a complete closure of the nest entrance as she obtained her third prey. After
placing the spider inside, she exited and closed the entrance with sand. The
duration of this closure was 25 seconds; she periodically interrupted five
times to turn toward the entrance. This was followed by an orientation
flight.

Another wasp opened her nest and entered at 1012 hours (E.S.T.), after
spending the night in a short burrow 50 cm away. She exited, turned and
entered her nest twice more, then made an 18-second-long temporary clo-
sure, including two turns toward the entrance. She, too, made an orientation
flight following the closure. The female returned both without and with prey
between 1042 and 1052 hours, each time closing the entrance with sand
upon leaving. She brought two prey to the nest at 1101 and 1114 hours but
did not close the entrance with sand upon leaving. The female made no
other closures of this nest until she finally closed it after ovipositing at 1201
hours.

Other females observed later in the day (1400-1530 hours) made brief
temporary closures of their nest entrances, which included turning toward
the entrance one or more times, and culminated in low orientation flights,
or, they left the nest entrance open between consecutive provisions and
oriented by running around the entrance on the sand. In two cases females
hunted spiders on the sand near their nest entrances. One wasp stung a
small spider only 1.5 m from her entrance. Subsequent prey were captured
even closer to the entrance in all directions of the compass.

Details of prey transport for this subspecies were described by Krombein
(1964) and Kurczewski (1969). Smaller wasps with black and red abdomens
brought prey on the ground, whereas one noticeably larger provisioning
female with a mostly all-red abdomen made 30-cm-long flights. Females
spent from 5 to 44 (¥, 16; N = 9) minutes between consecutive prey. Neither
size of prey nor position within the provisioning sequence influenced the
period of time between consecutive trips. Two larger prey were released
inside the entrances by females and pulled in from inside the burrow, after

VOLUME 84, NUMBER 1 if)

the wasps had turned around. Eight smaller spiders were taken into the
nests without being released or were released and taken in as described
above. In the latter case most such releases were determined by a temporary
closure of the nest entrance in which case the wasp had to relinquish the
spider in order to remove the sand fill. But one release occurred with an
open entrance prior to making the final nest closure.

Females of M. s. slossonae exited from 15 seconds to 3.5 minutes (N =
7) later. In the case of the wasp spending a long period of time inside the
nest prior to exiting, such entries had been deterred by a sand-filled en-
trance, il.e., temporary closure. One female twice removed her nest closure
without relinquishing the spider, i.e., she held the spider with the middle
legs as she removed the sand fill with the forelegs.

Another wasp began her final closure after entering her open nest with
prey at 1159 hours. The components were identical to those described for
this subspecies by Kurczewski (1969). She completed filling the burrow and
distributing sand on the area of the entrance at 1216, taking a total of 17
minutes for the closure. Females that did not begin and complete nests in
a single day closed their burrow entrances completely with sand in late
afternoon, returning often after several minute intervals to throw additional
sand on the fill. They then moved 0.5—1.0 m away and dug short burrows
in which they spent the night, facing head outward. Females emerged head
first from these resting burrows usually between 0930 and 1030 hours
(E.S.T.) and completed provisioning their nests after opening the entrances.

Nests of M. s. slossonae were moderately short, shallow and unicellular.
Nests were either straight or curved considerably. Nest entrances and bur-
rows were |.5—2.0 mm in diameter. Four burrows averaged 7.8 (7.4—8.4) cm
in length and 5.6 (4.9-6.0) cm in depth, including cell length and height. The
elongate-ovoidal cells were 3.5—4.0 cm wide and 6—7 mm long.

Females of this subspecies preyed upon a variety of small spiders as
follows: THERIDIIDAE: Steatoda fulva (Keys.), |imm. 6. MICRYPHAN-
TIDAE: Grammonata sp. nr. texana Banks, | ad. 2; Grammonata sp., 5
imm. OXYOPIDAE: Oxyopes salticus Hentz, | imm. LYCOSIDAE: Pirata
sp., | imm.; Arctosa littoralis (Hentz), 1 imm. GNAPHOSIDAE: Micaria
sp., limm. 6. SALTICIDAE: Pellenes sp., 7 imm. Prey were incompletely
paralyzed and moved their legs and emitted silk from the spinnerets.

Two completely provisioned nests contained 7 and 8 prey. Two unfinished
nests held 5 and 2 spiders. Individual spiders weighed from 0.5 to 3.5 (x,
1.3; N = 22) mg. Two wasps weighed 1.5 and 2.0 mg. The contents of two
fully provisioned cells weighed 9.0 and 11.5 mg.

Prey of M. s. slossonae were placed variously in the cells as follows:
ventral side upward, head inward, 8; dorsal side upward, head inward, 11;
and, dorsal side upward, head outward, 3. No prey were placed ventral side
upward and head outward or sideways. The contents of one cell with 7 prey

74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

exemplifies the heterogeneity of the prey positioning: 2, dorsal side upward,
head outward at the back end; 3, dorsal side upward, head inward in the
middle: and, 2, ventral side upward, head inward at the front end.

Eggs of M. s. slossonae were attached to the spiders as described and
figured for M. americanus by Kurczewski (1969). Eggs ranged from 1.0 to
1.5 mm in length. Two eggs were affixed to the bases of the left sides of the
spiders’ abdomens. These prey were placed dorsal side upward and head
inward in the middle of the cells. Spiders bearing eggs weighed 1.0 and 1.5
mg; these prey were the second largest individuals in the cells, in each case
animm. Pellenes sp.

DISCUSSION

Miscophus (Miscophus) americanus, M. (Nitelopterus) slossonae bar-
beri, and M. (Nitelopterus) s. slossonae are multivoltine throughout much
of their range. Even in upstate New York, M. americanus nests from mid-
June through September. All species inhabit areas of sandy soil with sparse
vegetation, preferring to build burrows in pits, depressions and hoof- and
footprints. The nests observed in the present study were constructed from
the sand surface. Whereas both subspecies of M. (Nitelopterus) slossonae
preferred to nest in flat sand, M. (Miscophus) americanus dug nests into
vertical sand cliffs. This is in sharp contrast with my earlier (1969) paper in
which I reported M. americanus nesting in flat or slightly sloping sandy
soils in New York and Florida.

The behavioral components involved in digging the burrows are similar
for all species. Soil removal is accompanied by little or no leveling of
the tumulus in front of the entrance. Females of M. (Miscophus) ameri-
canus spent less time during burrow construction than either subspecies of
M. (Nitelopterus) slossonae, probably a reflection of the shorter burrows
of the former species.

Upon completion of the burrow females of M. (Miscophus) americanus
and M. (Nitelopterus) slossonae barberi left the nest entrance open as they
went in search of prey. Some females of M. (Nitelopterus) s. slossonae left
entrances open whereas other females, especially those nesting in late morn-
ing, made brief sand closures of their entrances prior to hunting. Both sub-
species of M. (Nitelopterus) slossonae were observed making nest closures
during the provisioning sequence, typical slossonae more frequently and
under more varied conditions than slossonae barberi. Closures made by the
latter subspecies were observed during periods of ant interference. The
extent of temporary closure and the situations surrounding its presence or
absence have been discussed in some detail by Krombein and Kurczewski
et and Kurcezewski (1969), suffice it to say it is an extremely variable

ehavior.

Females of M. (Miscophus) americanus and both subspecies of M. (Ni-

VOLUME 84, NUMBER 1 75

telopterus) slossonae were observed making orientation flights or orienta-
tion ‘‘runs’’ or ‘“‘walks”’ following burrow construction and exit from the
entrance. Orientation flights predominated in the case of M. americanus
females nesting in sand cliffs, whereas those females nesting in flat or slight-
ly sloping sand in an earlier study (Kurczewski, 1969) oriented predomi-
nantly by running or walking around the entrance. In M. slossonae, espe-
cially s. slossonae, females which made temporary nest closures usually
oriented by flying, whereas those which left nest entrances open oriented
by running around the entrances. In neither M. s. slossonae nor M. slos-
sonae barberi was there a correlation between the form of orientation and
the manner in which the prey was brought to the nest. Miscophus ameri-
canus females, which often oriented in flight in front of sand cliffs, brought
prey to their nests in flight.

Females of M. (Nitelopterus) s. slossonae hunted and captured prey near
their nest entrances, whereas those of M. (Miscophus) americanus often
flew rather long distances in order to obtain prey. The former species preyed
upon either wandering or web-spinning spiders on or near the sand surface
which were transported to the nest mostly on the ground, whereas M.
americanus females took mostly combfooted spiders from webs above
ground and brought these prey to their nests in flight. In at least M. (Ni-
telopterus) s. slossonae the prey is stung and paralyzed incompletely. In-
complete paralysis of the prey involving spiders moving their legs and emit-
ting silk from the spinnerets was the rule also in M. slossonae barberi and
M. (Miscophus) americanus. In contrast to earlier studies of species in this
genus (Cazier and Mortenson, 1965; Krombein and Evans, 1954; Krombein
and Kurczewski, 1963), I found no evidence of amputation of the legs of
the prey in the present study.

The mechanisms of prey transport among species in this genus have been
summarized by Kurczewski (1969). The exact manner of transport and the
extent of the flight varies with the species of wasp and the relative sizes of
the females and their prey. Miscophus (Nitelopterus) s. slossonae, for ex-
ample, practiced ground transport as well as carrying prey in short flights,
whereas M. (Miscophus) americanus invariably transported prey in long,
extensive flights. Both Kurczewski (1969) and Bohart and Menke (1976)
have speculated on the relationship between the method of prey transport
and the length of wings and degree of wing venation in some of the species.

In the species of Miscophus studied herein the manner in which prey was
taken into the nest was dependent upon the absence or presence of a tem-
porary closure and size of spider. Smaller spiders were usually taken di-
rectly into the open burrow, whereas larger prey were often released tem-
porarily just outside or inside the entrance and pulled in from inside after
the wasp had turned around.

The behavioral components involved in final closing the nests are similar

76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Species of prey of some species of Miscophus.

i
Species of Prey No. Specimens, Stage Source

Miscophus (Miscophus) americanus

THERIDIIDAE
Theridion australe Banks Simm. Kurczewski, 1969
Theridion differens Emerton 1Sjades22 lepenaicis Kurczewski, 1969; MI-3-6,
37 imm. 8-13
Theridion flavonotatum Becker 2 ad. 2 MI-7
Theridion murarium Emerton 24 imm. Kurczewski, 1969; MI-3, 4,
(te)5 2 Hil) ik}
Theridion redimitum (Linn.) | imm. MI-3
Theridion sp. 5 imm. MI-7
ARANEIDAE
Araneus sp. 4 imm. MI-7
Miscophus (Nitelopterus) slossonae barberi
LINYPHIIDAE
Meioneta formica (Emerton) lad. 9 Krombein and Kurcezewski,
1963
LYCOSIDAE
Pirata sp. | imm. NI-24
Geolycosa sp. | imm. Krombein, 1964
Schizocosa sp. | imm. NI-22
Lycosa sp. | imm. Krombein, 1964
SALTICIDAE
Habrocestum sp. nr. pulex Dradee 2 Gradeao: NI-21-24
(Hentz) limm. 6; 15 imm.
Pellenes sp. | imm. Krombein and Kurezewski,
1963
Miscophus (Nitelopterus) s. slossonae
THERIDIIDAE
Steatoda erigoniformis 10 imm. Kurezewski, 1969
(Cambridge)
Steatoda fulva (Keys.) limm. 6 NI-17
MICRYPHANTIDAE
Grammonata sp. nr. texana laden? NI-18
Banks
Grammonata sp. Simm. NI-17-19
LIN YPHIIDAE
Meioneta formica (Emerton) lad. 9 Krombein and Kurczewski,
1963

eee

VOLUME 84, NUMBER 1 77

Table 1. Continued.

Species of Prey No. Specimens, Stage Source
TETRAGNATHIDAE
Tetragnatha laboriosa Hentz 4 imm. Kurezewski, 1969
OXYOPIDAE
Oxyopes salticus Hentz 1 imm. NI-18
LYCOSIDAE
Pirata sp. 1 imm. NI-17
Pardosa sp. | imm Kurezewski, 1969
Arctosa littoralis (Hentz) | imm. NI-20
Arctosa sp. 1 imm. Krombein and Kurezewski,
1963
Lycosa sp. 4 imm. Krombein and Kurezewski,
1963; Krombein, 1964
GNAPHOSIDAE
Micaria sp. limm. 3 NI-20
SALTICIDAE
Habrocestum pulex (Hentz) | imm. Krombein and Evans, 1955
Metaphidippus galathea (Walck.) | imm. Krombein and Evans, 1954
Metaphidippus sp. 1 imm. Krombein, 1964
Pellenes sp. 7 imm. NI-18, 19
DICTYNIDAE
Dictyna altamira Gertsch and ieial, & Krombein and Kurezewski,
Davis 1963

in not only the species studied herein but also in many other species in both
subgenera. The typical closing components in their exact sequence have
been detailed by Kurczewski (1969).

The nests of most species of Miscophus are relatively short, shallow and
unicellular, the exception being M. evansi (Evans, 1963, 1970) and some
exotic species. Although data on nest dimensions, as summarized by Kurc-
zewski (1969), indicated that the nests of M. (Miscophus) americanus and
the two subspecies of M. (Nitelopterus) slossonae are similar, an analysis
of the burrow lengths (and cell depths in M. slossonae) in the present study
revealed that nests of M. (Nitelopterus) s. slossonae are longer and deeper
than those of M. (Nitelopterus) slossonae barberi and about twice as long
as those of M. (Miscophus) americanus. The differences between the nest
dimensions of these species clearly reflect differences in the habitats in
which they nested. Cells of M. americanus were longer, wider and higher

78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

than those of either subspecies of M. slossonae and prey biomass was
usually greater.

Kurezewski (1969) summarized the prey specificity or lack thereof of
species in the subgenera Miscophus and Nitelopterus. Most of the species
appear to restrict their hunting activities to specific ecological situations
where particular kinds of prey occur. Thus, M. (Miscophus) americanus
preyed almost entirely upon combfooted web-spinning spiders belonging to
the family Theridiidae (Table 1). The record of this species using immature
araneids in Florida is exceptional, although females of M. americanus ac-
cepted anesthetized immatures of Eustala (Araneidae) when they were sub-
stituted at the nest entrance for prey theridiids (Kurczewski, 1969). Both
subspecies of M. (Nitelopterus) slossonae were more general in their se-
lection of prey. Miscophus slossonae barberi preyed upon small, immature
Lycosidae and Salticidae (Table 1). One record of Meioneta formica (Lin-
yphiidae) for this subspecies was exceptional (Krombein and Kurczewski,
1963). Thus, this subspecies of M. slossonae captures mostly wandering
spiders on or near the sand surface. Miscophus s. slossonae is heteroge-
neous in its selection of prey, taking no less than nine families of spiders
from a variety of ecological situations (Table 1). Underscoring this lack of
specificity is the fact that members of the Theridiidae, Micryphantidae, Lin-
yphiidae, Tetragnathidae and Dictynidae are web-spinners, whereas those
of the Oxyopidae, Lycosidae, Gnaphosidae and Salticidae are wanderers.

The number of prey stored in the fully provisioned cells of the species of
Miscophus does not vary inversely with the sizes of the prey, as indicated
by Kurezewski (1969). This is substantiated by the diversity in the biomass
as evidenced by the total weights of the prey contents, e.g., M. americanus
(8-20 mg). Some of the fully provisioned cells of this species and of M.
slossonae barberi contained much empty space whereas others were fully
packed with prey. Variable weather conditions and/or prey abundance may
influence the number of prey stored in the cells.

The prey of M. (Miscophus) americanus and both subspecies of M.
(Nitelopterus) slossonae were placed in the cell variously with a prepon-
derance of spiders positioned dorsal side upward and head inward (48), then
ventral side upward and head inward (32) and dorsal side upward and head
outward (19). No prey were positioned ventral side upward and head out-
ward. Egg-bearing prey of these species were positioned either dorsal side
upward and head inward (11) or dorsal side upward and head outward (2).
The dorsal side upward positioning of the egg-bearing spider is related to
the affixment of the egg, i.e., laterally on one side or the other to the base
of the spider’s abdomen (see Fig. 5, Kurczewski, 1969). The egg may extend
well beyond the spider’s dorsal abdominal terga and would become dis-
lodged if the prey were placed ventral side upward on the floor of the cell.
Egg-bearing spiders were placed in the cells either at the front (4) or back
(4) ends or toward the middle (4), with no set pattern for either M. ameri-

i
]
|

VOLUME 84, NUMBER | 79

canus or either subspecies of M. slossonae. This find is in agreement with
previous studies on these species and M. kansensis (Kurczewski, 1969).
Spiders with eggs were frequently larger than other prey in the cells or of
the same size. Whereas the mean weights of prey for M. americanus, M.
slossonae barberi, and M. s. slossonae were 1.9, 1.2, and 1.3 mg, those for
egg-bearing prey of these species were 3.2, 1.9, and 1.3 mg, respectively.

Some unusual ovipositional behaviors were noted in the present study.
One cell of M. americanus contained two eggs of the same size and shape,
one affixed to the prey in the position typical of the genus and the other
lying on the floor of the cell. It is possible that one egg was dislodged from
a spider during oviposition, inducing the female to oviposit again on the
same or another prey; or, a second wasp may have inadvertently entered
the wrong nest and oviposited and this egg was dislodged by the resident
female as she laid her egg. Despite the fact that Kurczewski (1969) stated
that the egg of species of Miscophus is not deposited until the full comple-
ment of prey has been placed in the cell, the possibility of rare instances of
progressive provisioning should not be overlooked. One cell of M. ameri-
canus was Closed with only three spiders, an egg, and considerable empty
space. Although I observed final closure of this nest the female or another
wasp was collected with prey near the nest entrance minutes later. The final
closure I observed may have been an extensive, several-minute-long tem-
porary closure. A filled nest of M. slossonae barberi was excavated and
contained five prey and an egg; shortly thereafter a female with prey arrived
and attempted to enter this nest. Whether these observations are, in fact,
variations of progressive provisioning behavior or misinterpretations of ob-
servations are unknown but they are certainly worth noting.

The behavioral observations presented herein substantiate Bohart and
Menke’s (1976) inclusion of the former genus Nifelopterus as a subgenus of
Miscophus. My data reveal striking similarities among species in the sub-
genera Miscophus and Nitelopterus in nest-site selection, burrow construc-
tion, orientation, mechanisms of prey transport, final closure, nest archi-
tecture, prey size and their placement in the cell and position of egg
attachment. Noteworthy behavioral differences between species in these
subgenera include nesting microhabitat and related manner of prey trans-
port, temporary closure activity and kinds (families) of prey. The latter are
rather minor distinctions which are ecologically related. Similar differences
can be found among species belonging to many genera, e.g., Tachysphex,
Ammophila (Evans, 1959, 1965) and Bembix (Evans, 1957, 1966; Evans and
Matthews, 1973, 1975).

ACKNOWLEDGMENTS

I am indebted to the late Richard Archbold and his staff, particularly
James Lane, for providing the excellent facilities of the Archbold Biological
Station, Lake Placid, Fla., where some of my studies were undertaken. I

80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

thank Richard C. Miller for assisting me with some of the field studies on
Miscophus americanus and Roy A. Norton for identifying the prey spiders.

LITERATURE CITED

Bohart, R. M. and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. Univ.
Calif. Press, Berkeley. 695 pp.

Cazier, M. A. and M. A. Mortenson. 1965. Studies on the bionomics of sphecoid wasps. IV.
Nitelopterus laticeps and Nitelopterus texanus (Hymenoptera: Sphecidae). Pan-Pac.
Entomol. 41: 21-29.

Evans, H. E. 1957. Studies on the comparative ethology of digger wasps of the genus Bembix.
Comstock Publ. Assoc., Ithaca, N.Y. 248 pp.

—. 1959. Observations on the nesting behavior of digger wasps of the genus Ammophila.
Am. Midl. Nat. 62: 449-473.

—. 1963. Notes on the prey and nesting behavior of some solitary wasps of Jackson Hole,
Wyoming. Entomol. News 74: 233-239.

—. 1965. Simultaneous care of more than one nest by Ammophila azteca Cameron (Hy-
menoptera, Sphecidae). Psyche (Camb. Mass.) 72: 8-23.

—. 1966. The comparative ethology and evolution of the sand wasps. Harvard Univ.
Press, Cambridge, Mass. 526 pp.

———. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming. Bull.
Mus. Comp. Zool. 140: 451-511.

Evans, H. E. and R. W. Matthews. 1973. Systematics and nesting behavior of Australian

Bembix sand wasps (Hymenoptera, Sphecidae). Mem. Am. Entomol. Soc. 20: 1-387.

. 1975. The sand wasps of Australia. Sci. Am. 233: 108-115.

Krombein, K. V. 1964. Results of the Archbold Expeditions. No. 87. Biological notes on
some Floridian wasps. Am. Mus. Novit. 2201: 1-27.

——.. 1979. Larridae, pp. 1617-1650. In Krombein, K. V. et al., eds., Catalog of Hyme-
noptera in America North of Mexico, Vol. 2. Smithsonian Institution Press, Washington,
Die

Krombein, K. V. and H. E. Evans. 1954. A list of the wasps collected in Florida, March 29
to April 5, 1953, with biological annotations (Hymenoptera, Aculeata). Proc. Entomol.
Soc. Wash. 56: 225-236.

——. 1955. Annotated list of the wasps collected in Florida, March 20 to April 3, 1954
(Hymenoptera, Aculeata). Proc. Entomol. Soc. Wash. 57: 223-235.

Krombein, K. V. and F. E. Kurczewski. 1963. Biological notes on three Floridian wasps
(Hymenoptera: Sphecidae). Proc. Biol. Soc. Wash. 76: 139-152.

Kurezewski, F. E. 1969. Comparative ethology of female digger wasps in the genera Mis-

cophus and Nitelopterus (Hymenoptera: Sphecidae, Larrinae). J. Kans. Entomol. Soc.
42: 470-509.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 81-84

FOODS OF CAULOPHILUS SPP., PARTICULARLY THE
BROADNOSED GRAIN WEEVIL, C. ORYZAE
(GYLLENHAL), BASED ON INTERCEPTION RECORDS
(COLEOPTERA: CURCULIONIDAE: COSSONINAE)

DONALD R. WHITEHEAD

Systematic Entomology Laboratory, I[BII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Throughout the native range of the broadnosed grain weevil,
Caulophilus oryzae (Gyllenhal), its most common agriculturally important
food is avocado seed. Although C. oryzae has been introduced elsewhere,
it is not an efficient stored product pest; it needs moist food. Infestation
records confirm various other foods but not banana, whereas other Central
American Caulophilus are intercepted primarily from banana detritus.

The main purpose of this note is to confirm that the broadnosed grain
weevil, C. oryzae (Gyllenhal), uses numerous food resources in addition to
avocado seeds. A second purpose is to comment on other members of the
genus occasionally found with plant materials shipped to the United States
from tropical America. This information is from specimen data in the Na-
tional Museum of Natural History (USNM) and augments previously pub-
lished records.

Caulophilus oryzae (Gyllenhal)

Kuschel (1962) reviewed the known species of Caulophilus Wollaston and
recognized C. oryzae as the correct name for the species previously known
as C. latinasus (Say). Junior synonyms are Rhyncolus lauri Gyllenhal, Cau-
lophilus sculpturatus Wollaston, and Cossonus pinguis Horn.

Caulophilus oryzae has been reported as a pest of various plant materials
(fruits—acorns, avocado, chickpeas, corn, millet; roots—ginger, sweet po-
tato, taro) and, specifically, of stored products (Chittenden, 1911; Cotton,
1956; McFarlane, 1963; Richards and Herford, 1930). It is most frequently
found associated with avocado seeds, and this led me to question that the
avocado seed pest and the stored product pest really are the same. I sorted
and examined materials in the USNM and prepared appropriate genital dis-

82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sections, and I conclude that the various samples of C. oryzae are conspe-
cific.

The following associations with infested plant materials are noted: avo-
cado seeds—numerous records (Florida, California, Puerto Rico, Mexico,
Guatemala). Corn—several records and series (Florida, South Carolina,
Puerto Rico, Jamaica, Mexico). Pigeon peas—one series (Bahamas). Span-
ish pear—one series (Mexico). Seeds of mammy-apple—one series (Puerto
Rico). Cola nuts—one series (Jamaica). Yam tubers—several series (Ja-
maica). Dry ginger roots—several series (Barbados, Jamaica). Philodendron
seed pod—one specimen (Mexico). Other quarantine interceptions have
been made from various orchids, poinsettia flowers, pineapples, and cycads,
apparently as hitchhikers.

Caulophilus rufotestaceus (Champion)

The USNM has numerous individuals and series intercepted from banana
leaves and detritus (throughout Mexico and Central America), one specimen
from sweet potato (Panama), and one from orchids (Mexico).

Other Caulophilus Species

Caulophilus costatus Champion.—Four interception records from banan-
as (Guatemala, Honduras).

Caulophilus lineaticollis (Champion).—One interception record from ba-
nanas (Panama) and one from sweet potato (Panama).

Caulophilus quichensis (Champion).—One interception record, among
series of C. rufotestaceus, from bananas (Nicaragua).

DISCUSSION

Kuschel (1962) included 16 described species in Caulophilus, all except
the Nearctic C. dubius (Horn) from the American tropics. Most of these
species were transferred from other genera (Stenomimus Wollaston, Tyttho-
mimus Champion).

The records of the four Caulophilus species from banana detritus all date
from a period in the mid-1930’s, when special attention was paid to potential
pests introduced with shipments of bananas. Since then, changes in shipping
procedures such as refrigeration and cleaner cargoes have cut immigration
considerably. There are similar associations with banana detritus for various
members of the cossonine genera Pseudopentarthrum Wollaston, Rhinan-
isus Broun, and Rhyncolus Germar, although not in the numbers recorded
for Caulophilus rufotestaceus. In contrast, despite the wide diversity of
interception records for C. oryzae, none is an association with banana.

. Bananas and banana-like plants (Musaceae, Araceae) with fleshy stem
lussues are probable ancestral hosts for members of Caulophilus in general.
Thus, the single record of C. oryzae from a philodendron seed pod (Araceae)

VOLUME 84, NUMBER | 83

may have significance, as may a similar record of an unidentified Caulo-
philus species. If stem tissues are the ancestral food source for members of
the genus, adaptation to fruits of the same or related plants may be an
intermediate step toward adaptation to such foods as avocado seeds. Inter-
ception records show that avocado seeds are the most frequent cultivated
food source for C. oryzae, but more information is needed about nonculti-
vated foods.

Despite its name, C. oryzae should not be considered an important stored
product pest, at least of cereals, because it does not thrive on the commodity
if well dried and undamaged. It does not feed on whole grain or seed that
is dry and hard but will attack cracked or damaged seeds such as those
already attacked by other stored product pests, and it is common on cereals
only on the growing plant or just after harvest when the seed is still soft
(Cotton, 1956; McFarlane, 1963; Quintana et al., 1960). In Florida avocado
plantations, I found it to be abundant in seeds of fallen fruits which had
opened husks and little remaining pulp; it is not a pest of the fruits, only of
the seeds.

The native range of C. oryzae includes Guatemala, Mexico, and perhaps
Florida and the West Indies (see Chittenden, 1911), regions inhabited by
avocado and other species of Persea Gaertn. (Lauraceae). The beetle does
not have the cosmopolitan distribution characteristic of the other stored
product weevils (Sitophilus spp., Rhynchophorinae). To the north, C. ory-
zae ranges at least to South Carolina, probably a secondary extension. Rec-
ords from California (Barrett, 1931), Hawaii (Beardsley, 1960), Madeira
(type-locality of sculpturatus), Sweden (type-locality of oryzae), and else-
where are the result of introductions. Caulophilus oryzae does not appear
to be notably adept at colonization, and I suspect that permanent coloni-
zations are likely only in avocado growing regions.

ACKNOWLEDGMENTS

I thank J. R. Gorham, Food and Drug Administration, USDA, and R. D.
Gordon, R. W. Hodges, and A. S. Menke, Systematic Entomology Labo-
ratory, USDA, for critical review of this manuscript.

LITERATURE CITED

Barrett, R. E. 1931. A new pest of avocado in California. Pan-Pac. Entomol. 7: 191.

Beardsley, J. W. 1960. [Note]. Proc. Hawaii. Entomol. Soc. 17: 176.

Chittenden, F. H. 1911. The broad-nosed grain weevil. (Caulophilus latinasus Say.). Papers
on insects affecting stored products. U.S. Dep. Agric. Bur. Entomol. Bull. 96: 19-24.

Cotton, R. T. 1956. Pests of stored grain and stored grain products. Burgess Publ. Co.,
Minneapolis. 306 pp.

Kuschel, G. 1962. Some notes on the cossonine genus Caulophilus Wollaston with a key to
the species (Coleoptera: Curculionidae). Coleopt. Bull. 16: 1-4.

84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

McFarlane, J. A. 1963. An annotated record of Coleoptera, Lepidoptera, Hemiptera and
Hymenoptera associated with stored produce in Jamaica. Tropical Agric. 40: 211-216.

Quintana, R., D. A. Wilbur, and W. R. Young. 1960. Insectos del grano almacenado que
infestan al maiz en el campo. Agric. Tec. Mex. 10: 40-43.

Richards, O. W. and G. V. B. Herford. 1930. Insects found associated with cacao, spices and
dried fruits in London warehouses. Ann. Appl. Biol. 17: 367-395.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 85-96

ARBORICHTHONIUS N. GEN., AN UNUSUAL ENARTHRONOTE
SOIL MITE (ACARINA: ORIBATEI) FROM ONTARIO

Roy A. NORTON

S.U.N.Y. College of Environmental Science and Forestry, Syracuse,
New York 13210.

Abstract.—Arborichthonius styosetosus, n. gen., n. sp., is proposed based
on the holotype female collected from soil, St. Lawrence Islands National
Park, Ontario. Unusual plesiomorphies include independent tibial and gen-
ual solenidia, astegasimy, and a partially divided femur I. Apomorphies
include unusual erectile setae and bidactyly.

The suprafamilial group of macropyline oribatid mites forming the Enar-
thronota (unspecified rank) of Grandjean (1946c), or the synonymous cohort
Euarthronota of Balogh and Mahunka (1979), constitutes an assemblage of
families of which all but three are usually considered to represent isolated
superfamilies. Except for the Brachychthoniidae which is rich in species,
most are apparently relicts, represented by one or a few genera each with
one or several nominal species. They often exhibit curious mixtures of rath-
er primitive and specialized structures. With regard to certain characters
which are quite fixed in the Brachypylina, for example secondary body
divisions and leg chaetotaxy, striking diversity is manifest in the Enarthro-
nota.

The purpose of this paper is to describe a new enarthronote mite exhib-
iting some of the most primitive character states found in the group, and at
the same time, specializations not observed in any other oribatid mite taxon.
A new genus is proposed, based on this species, but pending the conclusion
of ongoing research on enarthronote phylogeny, no familial assignment will
be suggested here.

The holotype (deposited in the Canadian National Collection, Ottawa) is
an adult female collected from moist soil beneath moss, on Thwartway
Island, St. Lawrence Islands National Park, Ontario, Canada, September
1976, by Evert E. Lindquist and Ian M. Smith, whom I thank for allowing
me to study it. No other specimens are known, and due to flattening in the
original slide mount general body shape and natural dimensions cannot be
definitively described. Despite this, its morphological uniqueness justifies

86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the proposals which follow. Morphological terminology used in the descrip-
tion is mostly that developed by F. Grandjean (see Travé and Vachon, 1975
for references).

Arborichthonius Norton, New Genus

Diagnosis.—With character states of the Enarthronota (Grandjean, 1969)
qualified as follows. Distinctly astegasime. Subcapitulum anarthric. Body
form dichoid, with light sclerotization. 32 pairs of notogastral setae. One
secondary ‘‘type S’’ scissure (Grandjean, 1946c) present, bearing erectile
setae e, and ey. Setae f;, f erectile, inserted on adjacent intercalary scler-
ites in pair of circular unsclerotized depressions. No suprapleural band;
sclerotization continuous between dorsal and lateral regions. Peranal seg-
ment absent. Epimera I with one pair of setae. Palp five segmented. Famulus
with bract. Solenidial formula unique (see below). Solenidial coupling var-
iously expressed: absent (seta d present) on genua I-III; present (imperfect)
on genu IV: absent (d present) on tibia III; hyperperfect (d absent) on tibiae
I and II. Femur I incompletely divided into basi- and telofemur. Ambula-
crum with symmetrical bidactyly. Adoral seta or, enlarged, flattened; or,
minute. No genital tracheae or eyes.

Type species.—Arborichthonius styosetosus, n. sp.

Arborichthonius styosetosus Norton, New Species

Etymology.—The Latin generic prefix arbori (meaning tree) and Greek
trivial prefix styo (meaning to erect or stiffen) both refer to the dendriform
erectile setae, f, and fy.

General.—Approximate length of body, not including gnathosoma 245 wm
(an estimate due to slightly crushed specimen). Integument, except as noted,
lightly but distinctly sclerotized, light yellowish tan in color, finely and ir-
regularly punctate. Unsclerotized arthrodial integument of subcapitulum,
legs, and palp distinctly granulate (probable due to localized cerotegument);
that of ventral and lateral regions weakly striate.

Prodorsum (Fig. 1).—Lateral regions of prodorsum unsclerotized. Aspis
broader than long; posterior margin bends sharply ventrad before joining
hysterosomal integument. Rostral tectum very narrow, hardly developed;
anterior margin rounded, entire. Bothridial opening flush with surface of
aspis; internal ridges circular in distal %, longitudinal in sharply narrowed
proximal 2; with distinctly porose walls. Sensillus (ss) of uniform diameter,
Subpectinate, bowed, with longest branches on convex side; branches non-
birefringent in polarized light. Interlamellar seta (in) inserted close to both-
ridium; slightly longer than sensillus, finely attenuate and sparsely barbed,
probably directed vertically in life. Lamellar seta (/e) and rostral seta (ro)
progressively shorter than in and inserted closer to meson. Exobothridial
setae differing in form; xs unusually large, attenuate, length intermediate

VOLUME 84, NUMBER | 87

between in and /e, curving dorsad from insertion anterolaterad of bothri-
dium; xi short, straight. Supracoxal seta e/J hollow, incorporated with in-
tegument (Fig. 2).

Notogaster (Fig. 1).—Suprapleural band absent!. Sclerotization weak or
absent in anterolateral region. Anterior extent of sclerotization unclear, but
greater medially than laterally, where it ends at level of setae c, and cs.
Setal pair c, inserted at ends of narrow dark, internal thickening or rib (ti)
traversing the notogaster. Erectile setae e,, e. large, extending beyond pos-
terior margin of notogaster when in relaxed position; narrowly rounded
distally; with non-birefringent barbs. Setae of pair e, both inserted on un-
paired elongated intercalary sclerite; each seta e, on a laterally tapering
sclerite clearly separated from the central one. Erectile setae f, and f, un-
usually large, dendriform, with thick birefringent branches; tips of branches
acute or narrowly rounded. Seta f, inserted on a simple, trapezoidal inter-
calary sclerite; f, sclerite with a distinct straight or slightly curved medial
arm. Setae ps,, ps2 broad, foliate; bladelike non-birefringent portion later-
ally serrated and occupying *% the setal length (ps,, Fig. 3) or entire and
occupying 4 the setal length (ps,); birefringent portion basally thick and
bifurcated, with paraxial branch short, antiaxial branch long and attenuate.
Remaining setae of several types: c. and cs relatively short, acute, with |
or 2 barbs; cp, c,, d,, dz, hy attenuate, with sparse, indistinct barbs; hs
similar, but with barbs distinct; 4, intermediate in size and general appear-
ance between other / setae and setae of row e; pss small, simple, inserted
its length away from ventral margin of notogaster at level of posterior margin
of genital plate. All but smallest of notogastral setae appear hollow, with
very narrow central canal; canal extends into branches of f,, f2. Lyrifissures
in form of round or elliptical cupule leading internally to dark, sclerotized
tubular canal; im located posterolaterad of seta e, insertion; ip posterolat-
erad of f, insertion; ih located ventrally, anterolaterad of ps, insertion; ips
on extreme ventromedial edge of notogaster at folded junction with ano-
genital region, slightly posteromediad of ps; insertion; lyrifissue ia not ob-
served’.

Ventral region—Epimera of pairs I and II well defined, broadly convex;
each half-epimere separated from its counterpart by narrow, unsclerotized
medial groove. Apodemes ap, and ap, distinct. Epimera III and IV both
flat; pair III indistinctly separated, fused medially with each other and pos-
teriorly with epimera IV in proximal *%; pair IV medially separated by un-
sclerotized, striate integument. Apodemes aps, ap4, and apsj absent. Epi-

1 The specimen could not be observed laterally, but there seems to be no lateral break in the
sclerotization when viewed dorsoventrally.

* It may in fact be present in its usual position posteroventrad of seta c3, but interference
of legs III and IV precluded close examination of this area.

ti
QD Tet ATSF.

Cc.’

tf,

Figs. 1-3. Arborichthonius styosetosus, holotype female. 1, Dorsal aspect, slightly flattened

and broken. 2, Supracoxal seta e/, dorsal aspect. 3, Seta PS», dorsal aspect (hatched area is
birefringent).

VOLUME 84, NUMBER 1 89

meral setation (I to IV): 3-1-3-4 on left, 3-1-3-5 on right. Genital plates with
3 simple setae in lateral row and 5 (left) or 4 (right) in a medial row. Anal
plates immediately adjacent to genital plates and of approximately equal
length; each with 2 setae, in anterior 2. Adanal plates fused behind anal
plates, posterior separation from notogaster indistinct; 3 pairs of adanal
setae. Aggenital region very lightly sclerotized posteriad of single aggenital
seta and confluent with adanal plates; area approximately between aggenital
seta and epimere IV unsclerotized. Ovipositor short, with distinctly plicate
walls; with 9 pairs of setae, 3 pairs on dorsal lobe, 3 on each ventral lobe,
and 3 pairs of coronal setae; Ww, and ¢, large and finely attenuate.

Gnathosoma.—Subcapitulum (Fig. 4) with no evidence of secondary ar-
ticulations. With 4 pairs of subcapitular setae (a, m, n, h) of which a is
clearly thickest and longest. Infrabuccal cleft long; ventral commissure (Ji)
located slightly posteriad of palpal insertion. Dorsal integument of lateral
lips with 4 dark ridges or ‘“‘chevrons’’ which appear to radiate from the base
of seta a in Fig. 4. A pair of sclerotized plates (Fig. 5) present near base of
labrum, mediad of seta m as seen by transparency, which possess a row of
denticles anteriorly and about a dozen longitudinal rows of small rectangular
rasplike teeth; laterad, denticles become smaller and longitudinal rows
shorter. Rutellum narrow, tridentate. Adoral setae heteromorphic; or, re-
duced to small, simple indistinct seta; or, long, acute, undulating, curving
dorsodistally; or, distally broad, flattened, with a small, but distinct medial
notch in the proximal 4. Sclerotized pharyngeal cupola (Co) present, dorsad
of pharynx (Ph); without sagittal carina. Chelicera narrow, of the Hypo-
chthonius type; each digit with 3 teeth, those of the movable digit con-
centrated distally. Cheliceral setae subequal; cha inserted dorsally, chb
laterodorsally. Palp (Fig. 6) with setal formula (trochanter to tarsus) of 0-2-
1-3-9. Tarsal solenidion w medium sized, baculiform, inserted on proximal
4 of tarsus, not reaching distally to tip of segment. Palpal setae smooth or
with several small barbs; homologies, indicated in Figs. 6 and 7, with pattern
similar to Hypochthonius (Grandjean, 1946a). Anteriad of w setal alveoli
with tear-drop shape, pointed anteriorly. Only large, distal compound seta
(fused wl’, ul’, sul) eupathidic; small size precluded determining whether
or not individual branches were hollow.

Legs.—Relatively short, simple in form (Figs. 7, 10, 11, 12); lengths (base
of trochanter to base of claws) of legs I to IV in proportions 1.3:1.0:1.0:1.2.
Basi-telofemur division of leg I indicated by region of non-granulate, un-
sclerotized integument, broadest antiaxially (Fig. 8), narrowest paraven-
trally (Fig. 7), with 2 parts of femur joined by region of sclerotized integu-
ment extending from dorsal midline midway down paraxial face; setae d,

3 If the branches of the compound eupathid are counted separately, the formula is 0-2-1-
3-11.

90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

wv V
~ VY yuuuuuu
uu udU Wy UUUU
iu gu eduUuuU
Sgu yudduu

U4 yyuuu
Uy ybuuu
Wouu

Figs. 4-9. Arborichthonius styosetosus, holotype female. 4, Subcapitulum, ventral aspect.
5, Internal plate of subcapitulum. 6, Palp (same scale as Fig. 4), antiaxial aspect, with enlarged

distal eupathid. 7, Leg | (minus trochanter), paraxial aspect. 8, Trochanter and femur I, an-
tiaxial aspect. 9, Famulus.

VOLUME 84, NUMBER 1 9]

Figs. 10-12. Arborichthonius styosetosus, holotype female. 10, Leg II (minus trochanter),
dorsal aspect. 11, Leg III, dorsal aspect. 12, Leg IV (minus trochanter and base of femur),
paraxial aspect. Seta /’ of tarsus absent on other leg IV.

l’, 1", v” on telofemur, bv” on basifemur. Ambulacra symmetrically bidactyl;
no trace of medial claw or birefringent vestige. Setal formulae (trochanter
to tarsus, famulus included) as follows: leg I (1-5-5-5-18); leg II
(1-6-5-4-16); leg III (2-3-4-4-13); leg IV (2-3-4-4-13 or 14). Probable setal
homologies indicated in illustrations. Many leg setae appear to have thin,
inconspicuous central canal, at least near base. Famulus (Fig. 7, e, Fig. 9)

92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

bifurcate, with ventral branch curved, somewhat spatulate, dorsal branch
in form of overhanging bract. Proral setae (p) smooth, with large central
canal, the only well defined eupathids on tarsus I; subunguinal (s, eupathidic
in most oribatid mites) similar to other tarsal setae. Solenidial formulae
(genu to tarsus) as follows: leg I (2-1-3); leg H (1-1-2); leg HI (1-1-0); leg IV
(1-0-0). Three are long, tactile: tibial solenidia @ I and ¢ II and genual so-
lenidion o IV. Remaining solenidia baculiform (@,I, w.II, w,II, @ HI, o III,
o IV) or piliform (@.I, wl, oI, oI). Seta d coupled to o on leg IV, in
adjacent but separate alveolus. Setae d absent on tibiae I and II.

DISCUSSION
Unusual Pleisomorphies

1. Independence of leg setae d.—Arborichthonius styosetosus is one of
the few Enarthronota which do not completely exhibit the derived close
association, or coupling, of genual and tibial solenidia (on legs where they
exist) with normal setae (usually d) of the respective segments. Some ter-
minology helpful in discussing such couplings can be defined as follows: 1)
Imperfect coupling occurs when the seta and solenidion are inserted in sep-
arate, but adjacent alveoli; 2) perfect, or normal coupling occurs when both
are inserted in a common alveolus, although the seta may be small and
regressive, or even fused to the solenidion; 3) hyperperfect coupling occurs
when the normal seta has regressed to the point of dissappearing, leaving
only the solendion, which is typically very large and tactile in form. Hy-
perperfect coupling may be the culmination of an ontogenetic trend, as seen
in certain brachypyline oribatid mites (such as Epidamaeus) where d be-
comes gradually smaller in the immature instars and is absent in the adult
instar. It also appears to culminate a similar phylogenetic trend, with the
result that seta d may be absent in all instars, as is common in poronotic
Brachypylina. Grandjean (1946b) has called these two apomorphies the Scu-
tovertex stage and Galumna stage, respectively. It is clear, however, that
we are dealing with grades of evolutionary development, which like many
regressive apomorphic states, have been reached independently in multiple
lineages.

Arborichthonius styosetosus exhibits several grades of coupling. There is
none on genua I-III and tibia III, where seta d and the respective baculiform
solenidion are independent and distant. Imperfect coupling is found on genu
IV, where solenidion o is tactile. Hyperperfect coupling is found on tibiae
I and Il; whether or not this develops during ontogeny is not known. The
absence of coupling on one or more genua or tibiae (where solenidia are
present) is known in only three other enarthronote species; these comprise
the superfamily Pterochthonioidea (Grandjean, 1948, 1950; Travé, 1967).
Pterochthonius exhibits perfect and hyperperfect coupling, as well as in-

VOLUME 84, NUMBER 1 93

dependence of seta d, whereas all setae d are independent in Atopochthon-
ius and Phyllochthonius. Among non-enarthronote oribatid mites indepen-
dent setae d (on genua and tibiae with solenidia) are found only in the
Palaeosomata and the genera Eulohmannia and Elliptochthonius, where
coupling is totally absent, and Nehypochthonius, Parhypochthonius, Ge-
hypochthonius, and Perlohmannia, where coupling is only found on certain
legs.

Although the ultrastructural proof is not yet available, it is reasonable to
suggest that the eventual result of the development of coupling and onto-
genetic or phylogenetic loss of the normal seta is a gradual takeover of
dorsal mechanoreceptive function by the large, flagellate, tactile, chemo-
receptive solenidion. Ancestrally, chemoreceptive and dorsal mechanore-
ceptive functions were presumably performed by the separate structures.
From a mechanoreception standpoint, the advantage of coupling, particu-
larly the hyperperfect type, may be that solenidia, which are hollow (and
presumably less dense than normal setae) and elastic, may be less prone to
breakage than similarly sized flagellate normal setae, which are solid and
more brittle.

2. Solenidial formulae.—Also rather primitive is the rich complement of
solenidia; next to Atopochthonius and Phyllochthonius, is the least regres-
sive known in the Enarthronota. Among all known Oribatei (Grandjean,
1964) the specific formula of Arborichthonius is unique.

3. Rostral tectum.—Arborichthonius is one of the few enarthronote genera
to retain the ancestral astegasime condition, whereby the rostral tectum is
poorly developed, not fully covering the retracted mouthparts from above.
Some Brachychthoniidae, such as Liochthonius are astegasime (Grandjean,
1963), as is the haplochthoniid genus Amnemochthonius (Grandjean, 1948).
Both genera contain very small and incompletely sclerotized species which
are regressive in many characters; whether or not the astegasime condition
is also a result of regressive loss of rostral sclerotization is not clear in these
groups. Arborichthonius maintains astegasimy despite light but relatively
uniform general sclerotization.

4. Divided femur I.—This is one of the most interesting character states
exhibited by Arborichthonius. Within the Oribatei, divided femora were
known only in the Palaeosomata and are considered ancestral to the typical
single femur formed by the integration of these two segments (Grandjean,
1954b). Arborichthonius exhibits a partially divided femur (Figs. 7, 8); the
separation is incomplete in the paradorsal region and appears, in effect, to
form a poorly defined hinge. Is this subdivision ancestral or secondarily
derived by localized desclerotization? There is little evidence to support
either conclusion and what does exist is somewhat contradictory. The cen-
tral placement of the division and femoral chaetotaxy support the former
conclusion; seta bv” is a basifemoral seta in the Palaeosomata, d, /’, /” and

94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

v" are telofemoral setae (Grandjean, 1954b). Secondary desclerotization
might have produced a deviant pattern of setal distribution between the two
parts. Also, the fact that the division is on the first leg and not the others
is consistent with Grandjean’s view on retardation of character appearance;
in the Palaeosomata leg I develops the division earlier than the other legs
and would be the last to lose this character if retardation was occurring in
phylogenetic time. On the other hand, the presence of smooth articulating
integument, in contrast to the granulate arthrodial integument between other
leg segments, is consistent with a hypothesis of secondary desclerotization.

Unusual Apomorphies

1. Notogastral setae.—Like certain other Enarthronota, Arborichthonius
has hypertrophied notogastral setae which the mite can raise and lower. The
evidence for this is in the nature of their insertions. In sclerotized Enar-
thronota, erectile setae always insert on an intercalary sclerite which is
bordered enteriorly and posteriorly by unsclerotized integument. In previ-
ously described species these intercalary sclerites are associated with one
of the transverse notogastral scissures characteristic of the Enarthronota,
such that when the notogastral plates immediately in front of and behind the
scissure are pulled apart, presumably by hysterosomal distension but pos-
sibly by bending the hysterosoma ventrally, the unsclerotized connective
integument is stretched and the associated setae erected (Grandjean, 1931).
Grandjean (1946c) has called this kind of scissure ‘‘type S.’’ Atopochthon-
ius, Phyllochthonius and certain Palaeosomata have erectile setae which
apparently function in a similar way, except that there are no articulating
notogastral plates and erection appears to be simply a result of general
stretching of gastronotic integument by hysterosomal distension (Grandjean,
1948). Arborichthonius exhibits a type S scissure bearing erectile setae e,
and e,. Although the three dimensional body shape of the Arborichthmius

styosetosus holotype is difficult to determine, due to flattening on the mi-
croscope slide, the fact that the intercalary sclerite bearing the setae of pair
e, is not divided is evidence that the mite is rather flat in life. Grandjean
(1948) has noted that separation of intercalary sclerites, such as is found
between all other such sclerites of Arborichthonius, is necessary in convex
mites in order to avoid cuticular distortion when setae are erected.

Unlike other known sclerotized enarthronote mites, Arborichthonius has
erectile setae which are not associated with a type S scissure. Setae f, and
f, are inserted in paired circular regions of unsclerotized integument which
are concave when setae are in their resting (horizontal) position. The erectile
nature is certain because of the adjacent, but independent intercalary scler-
ites on which they insert. Hysterosomal distension presumably causes the
unsclerotized integument to bulge outward, erecting the four setae.

Foliose setae ps, and ps,, while not erectile, are unlike any known in

VOLUME 84, NUMBER 1 95

other oribatid mites. Combined with the large dendriform setae of row f,
they give the posterior region a general appearance more characteristic of
certain Scutacaridae (Prostigmata) than an oribatid mite.

2. Pharyngeal cupola.—The most complex, specialized pharyngeal struc-
ture known in oribatid mites is found in the enarthronote superfamily Cos-
mochthonioidea and the derivative Protoplophoridae (Grandjean, 1946c,
1954a, 1962). Included in this aspiratory apparatus is a thin, ventrally con-
cave sclerotized internal plate on which the dilator muscles of the pharynx
originate. Because of the association with pharyngeal musculature, this
plate, or cupola, appears to be a derivative of the epistome of other arachnid
groups. Arborichthonius, Atopochthonius and Phyllochthonius, while lack-
ing most of the pharyngeal specializations of cosmochthonioid mites, do
exhibit a thin cupola dorsad of the pharynx, although it is not as well de-
veloped as in the latter group.

3. Adoral setae.—Arborichthonius belongs to a small group of Enarthro-
nota with unusual adoral setae; this group also includes Hypochthonius,
Hypochthoniella, Atopochthonius, Phyllochthonius, Mesoplophora, and
Archoplophora. Typically, seta or, is highly reduced or vestigial; or, is
large, flattened, spatulate, or otherwise specialized, but always exhibits an
unusual medial notch somewhere near its midpoint; or; is also large, but
attenuate except in the later two genera. This combination of adoral modi-
fications appears to be a synapomorphy.

4. Bidactyly.—Few oribatid mites are symmetrically bidactyl. Except for
Gehypochthonius rhadamanthus Jacot and the enarthronote genera Arbor-
ichthonius, Atopochthonius, and Phyllochthonius, the only known bidactyl
oribatid mites are those in which one of the lateral claws has been lost
(Grandjean, 1939). These four taxa have well developed lateral claws as
adults but not even a vestige of a central claw. One can suggest that this is
an evolutionary grade a step beyond that of Acaronychus, Eulohmannia,
Nehypochthonius, Elliptochthonius, Parhypochthonius, and Gehypo-
chthonius xarifae Strenzke in which the central claw is highly reduced in
size. The presence of symmetrical bidactyly in the Enarthronota as well as
both bidactyly and centrally regressive tridactyly in the genus Gehypo-
chthonius, suggests that symmetrical bidactyly has appeared in more than
one lineage.

LITERATURE CITED

Balogh, J. and S. Mahunka. 1979. New taxa in the system of the Oribatida (Acari). Ann.
Hist.-Nat. Mus. Natl. Hung. 71: 279-290.

Grandjean, F. 1931. Observations sur les oribates (2° série). Bull. Mus. Natl. Hist. Nat. 3:

651-665.

. 1939. L’évolution des ongles chez les oribates (acariens). Bull. Mus. Natl. Hist. Nat.

11: 539-546.

——.. 1946a. Au sujet de l’organe de Claparede, des eupathidies multiples et des taenidies

96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

manidublaires chez les acariens actinochitineux. Arch. Sci. Phys. Nat. Geneve 28: 63—
87.

——. 1946b. La signification évolutive de quelques caracteres des acariens (1" série). Bull.
Biol. Fr. Belg. 79: 297-325.

1946c. Les Enarthronota (acariens). Premiere série. Ann. Sci. Nat. Zool. 8: 213-248.

1948. Les Enarthronota (acariens) (2° série). Ann. Sci. Nat. Zool. 10: 29-58.

1950. Les Enarthronota (acariens) (3° série). Ann. Sci. Nat. Zool. 12: 85-107.

1954a. Les Enarthronota (acariens) (4° série). Ann. Sci. Nat. Zool. 16: 311-335.

. 1954b. Sur les nombres d’articles aux appendices des acariens actinochitineux. Arch.

Sci. (Geneve) 7: 335-362.

1962. Nouvelles observations sur les oribates (2° série). Acarologia 4: 396-422.

1963. Sur deux especes de Brachychthoniidae et leur développement (oribates). Acar-

ologia (Paris) 5: 122-151.

1964. La solénidiotaxy des oribates. Acarologia (Paris) 6: 529-556.

1969. Considérations sur le classement des oribates. Leur division en 6 goupes ma-
jeurs. Acarologia (Paris) 11: 127-153.

Trave, J. 1967. Phyllochthonius aoutii nov. gen., nov. spec., un Enarthronota (acarien oribate)
nouveau de Cote d'Ivoire, avec la création d’une superfamille nouvelle, Phylochtho-
noidea. Zool. Meded. (Leiden) 42: 83-105.

Trave, J. and M. Vachon. 1975. Frangois Grandjean, 1882-1975 (Notice biographique et
bibliographique). Acarologia (Paris) 17: 1-19.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 97-101

NEW SPECIES OF UROSIGALPHUS (HYMENOPTERA:
BRACONIDAE) FROM MEXICO

LESTER P. GIBSON

U.S. Department of Agriculture, Forest Service, Northeastern Forest
Experiment Station, Delaware, Ohio 43015.

Abstract.—Two new species of Urosigalphus subgenus Neourosigalphus
Gibson are described, yucatanensis and meridianus. A key to Mexican and
Central American species is presented.

The subgenus Neourosigalphus was established by Gibson (1972a) and
now includes 23 species of which four are known from Mexico and Central
America (Gibson, 1972b). Two new species of this subgenus, from Mexico,
are described in this paper. Both belong to a group of Neourosigalphus that
bear carapace tubercles and have 14 antennal segments. They do not resem-
ble previously described species from Mexico but do resemble some South
American species.

KEY TO SPECIES OF THE SUBGENUS NEUROSIGALPHUS FROM
MEXICO AND CENTRAL AMERICA

1. Carapace with apical tubercles; 14 antennal segments .............. 4
— Carapace without apical tubercles; 15 or 16 antennal segments ...... 2

2. Antenna 15 segmented; antenna dark brown; hindfemur testaceous
FE eee os ae epee 3

— Antenna 16 segmented; antenna medium brown, hindfemur red
DEO Wie eens ed orate, deve brstars: 2 68 ore ewonne See pepe avocadoae Gibson

3. Palpi yellow; tegula dark testaceous, mesopleuron impunctate cen-
CUA We TTER ES cheyd ef eps z yeu ee a Stay oteps phn oem rags punctifrons Crawford

— Palpi brown to dark brown, tegula dark red black, mesopleuron
Spaisely, punctate. centrally xx. ssi mayo 4 © 426 be neopunctifrons Gibson

4. Carapace tubercles long and well developed, 0.2 mm long (Fig. 3),
2 with ovipositor as long as carapace length ......... flavens Gibson

— Carapace tubercles short, less than 0.12 mm long (Figs. 1, 2), 2 with
OVIPOSiton,2s aswlong.as- carapace length 4/5: isssjassibidu Fyre bye Sebel 5

5. Palpi yellow; wings tinted brown; hindcoxa orange ..............-.
aren OR Niro dual ays Ay uae ee yucatanensis, new species

98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

— Palpi light to medium brown; wings clear; hindcoxa brown .........
i Sarit Io pe tit niin cat cae meridianus, new species

Urosigalphus (Neourosigalphus) yucatanensis Gibson, New Species
Fig. |

General features.—Length 2.5 mm. Head and thorax deep red brown;
carapace dark red brown; abdomen brown; clypeus red apically; labrum
rufotestaceous; mandible rufotestaceous, with red base and apex; palpi pale
yellow; antenna medium brown, lighter ventrally and on scape and pedicel.
Fore- and midlegs yellow tan; hindcoxa and femur orange and hindtibia and
tarsus medium brown. Tegula light red brown. Wings tinted brown, with
brown hairs, costa light brown basally and medium brown apically; stigma
dark brown; the remaining veins medium brown distally but paler basally.

Female.—Head: General anterior outline roundly quadrate. Vertex finely
rugoso-punctate, temples and genal regions finely punctured. Lower face
rounded, evenly finely punctate; fronto-clypeal groove with an oval anterior
tentorial pit near each end. Clypeus elongate quadrate; lower central margin
gently concave. Mandible teeth long and narrow, outer surface rugose.
Ocelli circular, interocellar area slightly raised. Antennal scapes well sep-
arated with area between fossae finely rugoso-punctate and with a shallow,
broad, finely punctate antennal sulcus extending to rear ocelli. Antenna 14-
segmented; scape long and relatively slender, inner margin slightly rounded;
pedicel amost 2 as long as scape. Occipital carina complete.

Thorax: Unevenly punctate. Mesonotum fairly regularly punctate, general
surface uneven; notaulices broad, deep, with large foveae, area of conver-
gence broad, rather quadrate, depressed, with large carinate punctures;
median lobe densely finely punctate with a small, short central carina an-
teriorly; lateral lobes gently flattened, finely punctured with a lateral border
of carinate punctures; posterolateral carina weakly flangelike. Scutellar
groove with 5 dissecting carinae. Scutellum roundly trigonal in dorsal pro-
file, rugoso-carinate; posterior surface abruptly sloped. Propodeum flat pos-
teriorly with a strong transverse carina; central dorsal carina absent, trans-
verse carina proceeds down both sides in an arc to the point of abdominal
articulation; dorsal surface narrow with irregular rugae delimiting irregular
depressions; posterior surface roughly and densely rugoso-punctate; lateral
areas Coarsely rugoso-punctate. Mesopleuron punctate; middle central area
impunctate, lower anterior area with large carinate punctures in a meso-
pleural groove; mesosternum punctate with very small longitudinal carinae.
Lateral regions of pronotum coarsely punctate, upper posterior corner with
a slight bend and with triangular area behind with 2 large punctures sepa-
rated by a carina. Wings tinted brownish; submedian cell longer than the
median basally.

Abdomen: Carapace oval in dorsal and lateral views, longitudinally ru-

VOLUME 84, NUMBER 1 99

Figs. 1-2. 1, Urosigalphus (Neurosigalphus) yucatanensis, type. 2. U. (N.) meridianus,
type.

goso-punctate anteriorly, densely punctate and shiny posteriorly, apex with
2 small apical spines, 0.12 mm long (Fig. 1). Ovipositor 1.0 mm long, nearly
% as long as carapace, ovipositor sheath reddish brown.
Male.—Unknown.
Holotype.—, Progreso, Yucatan, Mexico; 23-VII-62; H. E. Evans Col-
lector. In MCZC.

100 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Urosigalphus (Neurosigalphus) flavens.

Distribution.—Mexico: Yucatan.
Biology and host.—Unknown.

Urosigalphus (Neourosigalphus) meridianus Gibson, New Species
Fig. 2

General features —Length 3.0-3.5 mm. Head and thorax black; carapace
dark red black to black; abdomen medium to dark red brown; clypeus black,
labrum maroon to red black; mandible tan with base and apex dark red;
palpi light to medium brown, antenna medium to dark brown. Fore- and
midlegs yellow tan; hindfemur yellow tan to orange, hindtibia medium
brown and tarsus dark brown; all coxae usually brown to dark brown, darker
than femora. Tegula dark red brown or red black. Wings clear, hyaline with
brown hairs, costa, stigma, and veins medium to dark brown distally but
veins becoming paler basally.

Female.—Differs from yucatanensis as follows: Head: Temples, and ge-
nal regions finely to moderately rugoso-punctate. Lower face evenly finely
rugoso-punctate; anterior tentorial pit round. Mandible teeth long but broad.
Area between antennal fossae finely rugoso-punctate and bicarinate with a
small central carina immediately anterior to median ocellus, and a rugoso-
punctate antennal sulcus.

Thorax: Notaulices broad but shallow with moderate punctures, area of
convergence depressed with moderate to large punctures; lateral lobes
densely finely punctured with a foveolate lateral carina. Scutellar groove
with 3 dissecting carinae. Scutellum coarsely rugoso-punctate throughout.
Propodeum flattened posteriorly with a moderately strong transverse carina;
central dorsal carina short and moderate, lateral areas rather irregularly
carinate and rugoso-punctate. Lower anterior area of mesopleuron with
moderate punctures in a mesopleural groove; mesosternum punctate with
a line of large sized punctures along center line and transverse carinae dis-

VOLUME 84, NUMBER 1 101

tally. Lateral regions of pronotum irregularly punctured, medially sparsely
punctate. Wings clear.

Abdomen: Carapace elongate oval in dorsal and lateral views, rugoso-
punctate posteriorly, apex with small apical spines, 0.04 mm long (Fig. 2).
Ovipositor 1.2 mm long, *% as long as carapace, ovipositor sheath reddish
black.

Male.—Unknown.

Holotype.—¢, Mexico. Yuriria, Guanajuato, 6500’, 7-VIII-62, H. E. Ev-
ans Collector. In MCZC.

Paratypes.—1 2 Patzcuaro, Michoacan, 15-VII-1965, H. E. Evans: 1 2
Chilpancingo, Guerrero, 3800’, 30-VII-1962, H. E. Evans (in MCZC). 1 2
Guadalajara, Jal., 23-28-VII-1965, H. E. Evans (in Rijksmuseum van Na-
tuurlijke Historie).

Distribution.—Mexico: Guanajuato, Michoacan, Guerrero, Jalisco.

Biology and host.—Unknown.

ACKNOWLEDGMENTS

I thank Alfred F. Newton, Jr. of the Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts (MCZC) and Kees van
Achterberg of the Rijksmuseum van Naturrulijke Historie, Leiden, Neth-
erlands for the loan of specimens.

LITERATURE CITED

Gibson, L. P. 1972a. Revision of the Genus Urosigalphus of the United States and Canada
(Hymenoptera: Braconidae). Misc. Publ. Entomol. Soc. Am. &(3 & 4): 83-134.
—.. 1972b. Urosigalphus of Mexico and Central America (Hymenoptera: Braconidae).

Misc. Publ. Entomol. Soc. Am. 8(3 & 4): 135-157.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 102-110

THE PHENOLOGY OF THE PLANT BUGS (HEMIPTERA: MIRIDAE)
ASSOCIATED WITH CEANOTHUS CRASSIFOLIUS IN A
CHAPARRAL COMMUNITY OF SOUTHERN CALIFORNIA

JOHN D. PINTO

Department of Entomology, University of California, Riverside, Califor-
nia 92521.

Abstract.—Seven species of Miridae were found to develop on Ceanothus
crassifolius at a locality in southern California. Mirid activity was confined
to the first five months of the year, the period of lowest temperatures and
greatest precipitation, and also the time of blooming and fruiting for C.
crassifolius. Although levels of synchrony are high, the periods of greatest
abundance of mirid species are spread out through the growing season.

There are little data on the host relationships of western U.S. Miridae. A
relatively recent list of associations (Knight, 1968) does not suggest a large
fauna on Ceanothus. However, I have found that species of Ceanothus and
other Rhamnaceae serve as hosts for a variety of plant bugs in California.
This paper deals with the species on C. crassifolius Torrey, or Hoaryleaf
Ceanothus, an evergreen shrub common between 450-1100 m in the dry
chaparral covered hills of southern California.

The study is based on collections of all instars of Miridae on C. crassi-
folius at a single site throughout most of 1979. Primary objectives were to
determine the number of species developing on this plant, their occurrence
relative to host phenology, and the degree of interspecific seasonal overlap.
Preliminary data of feeding behavior are also presented.

Seven species were found to develop on C. crassifolius. Six belong to the
Phylini and one to the Mirini. Identification to species is possible for only
five of the seven at this time. The phylines collected were Psallus ancorifer
(Fieber), Psallus breviceps Reuter, Psallus sp., Microphylellus bicinctus
(Van Duzee), Lepidopsallus californicus Van Duzee, and Phymatopsallus
sp. near croceguttatus Knight. The single mirine present was Pycnocoris
ursinus Van Duzee. In addition, five adults of Deraeocoris fulgidus (Van

Duzee) were collected. Since this species was not represented by immatures
it is not dealt with further.

VOLUME 84, NUMBER 1 103

SITE DESCRIPTION AND METHODS

The sampling site is located in the hills to the immediate west of Menifee
Valley, an inland southern California valley ca. 5 km east of Lake Elsinore
in SW Riverside Co. Since the phenology of C. crassifolius varies with
microhabitat, collections were restricted to an area of .44 hectare atop a
north-south running ridge (33°39'N, 117°13’W; 550 m elev.). This relatively
small area limited the number of plants, but minimized variation in expo-
sure, drainage and elevation. All samples were from plants judged to be in
approximately the same phenological state.

Considering the high level of flight activity in plant bugs (Southwood,
1960), movement of adults to the study area from plants in more advanced
phenological states is a confounding variable. However, the effect of adult
dispersal was probably minimized by choice of the study site where the host
plants were all in a similar stage of development. The site was also situated
ca. 0.50 km from locales with plants that were obviously disjunct pheno-
logically. Furthermore, the lack of seasonally disjunct adults during sam-
pling suggests that intersite movement was minimal.

The study area is dominated by chaparral but is distinctly influenced by
Coastal Sage Scrub vegetation (see Munz, 1959, for a characterization of
these communities). As is typical of the low hills bordering the hot, dry
interior valleys of southern California, perennial vegetation on the site is
relatively sparse. Only ca. 50% of the surface is covered by perennial can-
opy. The remainder is bare ground, much of it strewn with large granitic
boulders. Annual grasses and herbs cover much of the surface in winter and
spring. Dominant perennials are Salvia mellifera Greene (Labiatae), Eri-
ogonum fasciculatum Bentham (Polygonaceae), and Adenostema fascicu-
latum Hooker and Arnott (Rosaceae). Ceanothus crassifolius itself has a
patchy distribution and contributes only ca. 5% to the total perennial canopy
(E. M. Fisher, personal communication).

Seven relatively large C. crassifolius of 2-3 m height were sampled from
21 January to | December 1979. Immatures of the season’s first mirid species
were already present when collections began. Mirid activity continued from
that time until 4 July. The sampling dates during this time are given in Figs.
1 and 2. A mean of 6.3 days (range, 4-13) intervened between consecutive
collections. On only one occasion (20 June—4 July) did the number of inter-
vening days exceed 8. Samples after 10 July were made twice monthly. This
was the first date after 21 January that sampling failed to collect mirids.

Plant bugs were collected by beating vegetation and allowing specimens
to fall into a modified sweep net, 12 cm deep and 28 cm in diameter. Three
positions at 1-1.5 m height were sampled on each plant. Plant bugs were
quickly aspirated at each position. After all plants were sampled, the entire
catch was killed in cyanide. Immatures were then transferred immediately

104 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

POST BLOOM
PRE BLOOM aes ; >
100 -
|
te

@
Oo
|

T

0)
(@)

PERCENT OF TOTAL Miridae

SAMPLING DATE (1979)

Fig. 1. Phenology of Miridae and host (Ceanothus crassifolius) during 1979. Adults and
immatures combined for each plant bug species. @ = Psallus ancorifer; & = Psallus breviceps;
@ = Psallus sp.; O = Microphylellus bicinctus; x = Lepidopsallus californicus; A = Phy-
matopsallus nr. croceguttatus. Pycnocoris ursinus omitted (see Fig. 2). Summary of host
phenology indicated at top. Open arrow indicates day of maximum bloom; closed arrow in-
dicates time of fruit dehiscence.

to 70% alcohol for storage. Most of the adults were point mounted, the
remainder were kept in alcohol. All collections were made during daylight,
from 0900-1200 hours.

Mirid immatures and adults were readily associated by structural and
color traits common to both. Rearings of six of the species verified these
associations. Psallus ancorifer was the only species not reared. Three to
ten nymphs of each species were reared by placing them with fresh vege-
tation in a covered Petri dish (100 x 10 mm). Plant material was replaced
every other day. Specimens in rearing were also utilized for feeding obser-
vations. Bugs were provided leaves, fruit and stems. Flowers were also
made available to species active early in the season.

Rainfall and mean monthly temperatures were recorded from August
1978—July 1979 (Fig. 3). Precipitation was monitored at the study site. Tem-
peratures are from published U.S. Weather Bureau records from Sun City,
Calif., ca. 3.5 km from the site (433 m elev.). The periodicity of rainfall
during and immediately before this study was normal for southern Califor-
nia, 1.e., winter concentrated. The total amount of precipitation (435 mm),
however, was ca. 65% above normal.

Voucher specimens from this study are deposited in the collections of the
Department of Entomology, University of California, Riverside, and the
National Museum of Natural History, Washington, D.C.

VOLUME 84, NUMBER 1 105

63 3 2) 2

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2129 4 11 16 25 4 11 18 23.1 10 15 2229 6 13 2229 5 13 204 10

SAMPLING DATES (1979)
Fig. 2. Phenology of the seven species of Miridae on Ceanothus crassifolius during 1979.
Dotted lines indicate immatures, solid lines adults.

106 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

h. croceguttatus

L. californicus

M. bicinctus

Ps. sp.
|
Ps. breviceps
Py. ursinus

Ps. ancorifer

25
200 20 Ys
= a
£ 150 15 =
> =
O WJ
EE 100 Z 10 2
- Z Z S
© 50 P Z Z 3.
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AUG SEP OCT NOV DEC JAN FEB MAR’ APR MAY JUN JUL
1978 1979

Fig. 3. Occurrence of the Miridae on Ceanothus crassifolius relative to monthly precipi-
tation (bars) and average monthly temperature (connected dots) during 1978-79. December
occurrence of P. ancorifer assumed from collections in 1980.

RESULTS AND DISCUSSION

The results of sampling are given in Figs. | and 2. Figure | indicates the
percentage of all stages of each species relative to the total number of Mi-
ridae collected per sampling day. Figure 2 considers the immatures and
adults of each species separately as a percentage of total immature, and
adult catch, respectively, per sampling day. The latter also includes the total
number of individuals collected daily.

All seven species are packed into the first six months of the year. The
only mirids collected from 4 July-December were a few adults of various
species assumed to be strays from plants other than C. crassifolius. The
activity period for each species varied from two to three months. There is
no evidence for more than a single generation per year for any of the species.
Since first-instars of all species were collected prior to successive instars,
it is assumed that the summer and autumn are spent in the egg stage.

Although there was substantial synchrony, none of the species overlap
completely (Fig. 2). Instead, they follow one another closely throughout the
season. Sampling revealed the following order of appearance for phyline
species: Psallus ancorifer, Psallus breviceps, Psallus sp., Microphylellus

|
)
|

VOLUME 84, NUMBER 1 107

bicinctus, Lepidopsallus californicus, and Phymatopsallus near crocegut-
tatus. The mirine, Pycnocoris ursinus, was too uncommon to determine
confidently its order of appearance relative to P. ancorifer and P. breviceps.
The pattern of species occurrence shown here is similar to that reported by
Waloff (1968) for the mirids on Scotch broom in England.

Figure 1, containing all instars, shows that each of the six phylines rep-
resented the dominant species at some time during the sampling period.
Only the two earliest (P. ancorifer and P. breviceps) and latest species (L.
californicus and Ph. nr. croeceguttatus) were dominant in the absolute
sense in representing over 50% of the total mirid catch on any given sam-
pling day. However, the immatures and adults of all six species did so
dominate at some time when compared to heterospecific immatures and
adults, respectively (Fig. 2).

The appearance of P. ancorifer, the season’s first species, is not resolved
(Fig. 2). It was not yet present when plants were checked in early December
1978. Additional sampling was not possible until 21 January 1979 at which
time P. ancorifer immatures were in abundance. To better determine the
appearance of this species, host plants were checked weekly during the last
two months of 1980. In that year, first-instars were first collected on 7
December.

The occurrence of species relative to host phenology is summarized in
Fig. |. The flowering period for C. crassifolius is from January to April
(Munz, 1959). At the collecting site sampled plants flowered from 4 February
to 23 March. Psallus ancorifer was the only species active prior to
bloom. The appearance of P. breviceps was roughly concurrent with the
onset of flowering. Psallus sp., M. bicinctus and L. californicus first ap-
peared during flowering but most of their activity was postbloom. The last
species, Ph. nr. croceguttatus, did not appear until well after flowering was
complete, and, in fact, was most abundant after fruit dehiscence. Although
the largest number of species were active during fruiting, species present
during flowering were more abundant. Thus, 936 mirids (* = 133.7/day)
were collected on the seven sampling dates during flowering whereas only
647 (x = 92.4) were taken on the seven dates immediately after bloom.

Although sampling frequency was not designed to detect seasonal varia-
tion in sex ratios, such variation was apparent in P. breviceps, Psallus sp.,
M. bicinctus, and L. californicus. Populations of all four were male-biased
at the beginning of adult activity and female-biased toward the end. In P.
breviceps, for example, collections on 11 and 18 March with 45 males and
26 females deviated significantly from a 1:1 ratio (P < .05) as did samples
on 23 March and | April with 18 males and 58 females (P < .005).

Five of the seven mirids on Ceanothus crassifolius were observed feeding
in the laboratory. Ultimate instar nymphs of Psallus sp., Psallus breviceps,
Microphylellus bicinctus, and Phymatopsallus nr. croceguttatus were com-

108 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

monly observed to insert their proboscis in the underside of leaves. Since
this surface is covered with a woolly pubescence the actual points of inser-
tion relative to leaf venation were not observed. Fruit, stems and the upper
surface of leaves were ignored by these species. Flowers were available
only to Microphylellus bicinctus and Psallus sp. The latter fed on floral
parts but appeared to prefer leaves. The former was observed feeding on
leaves only. Feeding in a single adult of Pyncnocoris ursinus was observed
intermittently over a period of several days. It fed on fruit and ignored all
other vegetative parts provided. Flowers, however, were not available to
this species when observations were made.

The occurrence of several related species on the same host plant is of
evolutionary and ecological interest. This is especially so if the species occur
at the same time. Estimates of interspecific seasonal overlap are given in
Table |. Data are presented for the entire activity period of each species as
well as for the period of adult activity. The degree of seasonal overlap for
species pairs is reported as a percentage of the number of sampling dates
synchrony was confirmed, relative to the total number of sampling days
each species occurred. Indices of synchrony, summarizing degree of sea-
sonal overlap for each species, were also calculated (Table 1).

Seasonal overlap of the seven species is considerable (Table 1). As ex-
pected from Fig. 1, it is least for the first (P. ancorifer) and last (Ph. nr.
croceguttatus) species and greatest for intermediates (P. breviceps, P. sp.,
and M. bicinctus).

The degree of synchrony among congeners is of particular interest. Al-
though three of the species are currently placed in Psallus, it is clear that
P. breviceps is closest to M. bicinctus. It keys to Microphylellus and, ac-
cording to T. J. Henry (personal communication), it is closer to that genus
than to Psallus. The only other congeners are Psallus ancorifer and Psallus
sp. Both species pairs show considerable synchrony (Figs. 1, 2; Table 1).
However, in each case the overlap is primarily between immatures of one
and adults of its relative. This suggests that the potential for interspecific
sexual interaction, at least, is minimal. Possible modes of additional sepa-
ration, such as oviposition sites have not been adequately studied. Waloff
(1968) and Dempster (1964) reported some differences in the feeding habits
of Orthotylus species occurring together on Scotch broom. This had been
predicted earlier by Waloff and Southwood (1960) upon noting contrasting
rostral lengths among the different species. A similar anatomical difference
does not occur between congeners on C. crassifolius. Preliminary obser-
vation of feeding behavior in nymphs of P. breviceps and M. bicinctus does
not suggest different feeding sites. Both feed commonly on the underside
of leaves (see above).

The plant bugs on C. crassifolius occur during the coolest time of year
and during or directly after months of maximum precipitation (Fig. 3). This

VOLUME 84, NUMBER 1 109

Table |. Synchronic relationships of the Miridae on Ceanothus crassifolius .
Degree of Seasonal Overlap®” No.
— Synchronic
: i b califor- crocegut- Relation- _ Index of
Species ancorifer ursinus breviceps sp. bicinctus _ nicus tatus ships Synchrony‘
69 56 5 25 5 SS)
P. ancorifer xX = a ly p22. Bu = 2/3
67 55 22 0 0 0 3) 24.0
73 80 80 6 53
Py. ursinus a. xX e us ELE
60 80 60 40 23 6 43.3
75 100 83 67 5 5 3.
P. breviceps Te XxX S = =
3 100 63 38 13 5 46.2
54 92 Wal 5 23 A
Ea: Pus ey an X 8 Vi 6 68.0
25) 62 63 qs 50 0 5 45.8
42 75 67 92 83 33 Se
M. bicinctus ra aii = — X ae = 2 ee)
0 51 43 86 71 0) 4 41.8
33 67 58 83 83 42 3
L. californicus Cy x — sd Sho
0 33 ily 67 83 16 5 36.0
0 12 0 38 50 63 4 Die
Ph. nr. croceguttatus —_ a x= ex = oF X oa
0 0 0 0 0 25 l 4.2

4 Degree to which species X (horizontal) is overlapped seasonally by species Y (vertical)
determined by dividing the number of days both were collected together by the total number
of days X was collected multiplied by 100.

> Figures above horizontal lines in body of table refer to overlap regardless of instar; those
below horizontal line refer to adult overlap only.

© Index of synchrony for a given species determined by adding individual scores and dividing
by 6, the maximum number of synchronic interactions.

is the time of maximum growth and flowering for most chaparral plants. C.
crassifolius is among the first plants to bloom in the chaparral and its as-
sociated mirids are the first of the season to occur. C. crassifolius commonly
begins to bloom in January, two months or so before the other dominant
perennials, and before the appearance of most annuals.

The degree of host specificity for the mirids collected here is unknown,
nor is it possible to profitably use species distribution data as a means of
speculation since the taxonomy and biology of the species are so poorly
known. In any case, near the study site, adults of M. bicinctus have been
taken in low numbers from Quercus agrifolia Neé, and a series of P. brev-
iceps was collected from Rhamnus crocea Nuttall. Both plants are relatively
uncommon at or near the sampling site. The type series of M. bicinctus was
collected from an unidentified Ceanothus (Van Duzee, 1914). Psallus an-
corifer, a supposedly widespread species, was recorded from “‘clover”’ by
Knight (1927) and onions by Thompson (1945). Lepidopsallus californicus
is known from southern California and Fresno, Calif. The latter locale is out
of the range of C. crassifolius but is close to areas harboring congeners.

110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pycnocoris ursinus, so far, has been taken only on Ceanothus. I have col-
lected it on at least three species of Ceanothus, and two specimens in the
P. H. Timberlake collection (UC Riverside) were collected in Ensenada,
Baja California on Ceanothus verrucosus Nuttall.

ACKNOWLEDGMENTS

I thank E. M. Fisher for the data on plant cover at the study site and T.
J. Henry of the Systematic Entomology Laboratory, USDA, for the iden-
tification of the six species of Phylini. Figures 1-3 were prepared by P.
Mote.

LITERATURE CITED

Dempster, J. P. 1964. The feeding habits of Miridae (Heteroptera) living on broom (Saro-
thamnus scoparius (L.) Wimmer). Entomol. Exp. Appl. 6: 149-155.

Knight, H. H. 1927. Notes on the distribution and host plants of some North American
Miridae (Hemiptera). Can. Entomol. 59: 34-44.

—— —. 1968. Taxonomic review: Miridae of the Nevada Test Site and the Western United
States. Brigham Young Univ. Sci. Bull., Biol. Ser. 9(3): 1-282.

Munz, P. A. 1959. A California Flora. Univ. Calif. Press. 1681 pp.

Southwood, T. R. E. 1960. The flight activity of Heteroptera. Trans. R. Entomol. Soc. Lond.
112: 173-220.

Thompson, B. G. 1945. DDT to control Psallus ancorifer in onions. J. Econ. Entomol. 38:
Pia

Van Duzee, E. P. 1914. A preliminary list of the Hemiptera of San Diego County, California.
Trans. San Diego Soc. Nat. Hist. 2(1): 1-57.

Waloff, N. 1968. Studies on the insect fauna on Scotch Broom Sarothamnus scoparius (L.)
Wimmer. Adv. Ecol. Res. 5: 87-208.

Waloff, N. and T. R. E. Southwood. 1960. The immature stages of mirids (Heteroptera)
occurring on broom (Sarothamnus scoparius (L.) Wimmer). Proc. R. Entomol. Soc.
Lond. (A) 35: 39-46.

|

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 111-116

DESCRIPTIONS OF THE MATURE LARVA AND PUPA OF
HYPOMECIS UMBROSARIA (HUBNER)
(LEPIDOPTERA: GEOMETRIDAE)!

ROGERT L. HEITZMAN?

Maryland Center for Systematic Entomology, Department of Entomolo-
gy, University of Maryland, College Park, Maryland 20742.

Abstract.—The mature larva and pupa of Hypomecis umbrosaria (Hiib-
ner) are described and illustrated. A comparison is made with related gen-
era. What appear to be the first known larval defensive glands in a geometrid
are also described, and brief internal morphological studies are included.

The bivoltine Hypomecis umbrosaria (Hiibner) (Fletcher, 1979), formerly
known as Pseudoboarmia umbrosaria (Hubner), ranges from coastal Mas-
sachusetts to central Florida, west to eastern Texas and north through Ar-
kansas to central Missouri (Rindge, 1973; Heitzman, 1973). Hypomecis gno-
pharia (Guenée) is a sibling species only distinguishable by slight differences
in wing pattern and genitalia. The brief larval description of H. gnopharia
by Guenée (1857), revealed at least no superficial differences from H. um-
brosaria. The synonymy of these two species is confused, resulting in nu-
merous questionable host records, i.e., horse chestnut, birch, elm, and var-
ious conifers (Rindge, 1973).

Hypomecis is a Holarctic genus represented by five species. A single
Palearctic species H. punctalis (Scopoli), is found from Japan to western
Europe, and four Nearctic species occur in eastern and southeastern United
States (Rindge, 1973). This genus has affinities to the more primitive mem-
bers of the Boarmiini, a large worldwide tribe with 34 Nearctic genera. The
larval and pupal stages have been studied in many of the related genera,
e.g., Anavitrinella, Anacamptodes, Iridopsis, Glena, Stenoporpia, and
Cleora (McGuffin, 1977). The purpose of this paper is to describe the di-
agnostic morphological characters of the mature larva and pupa of Hypo-

1 Supported in part by the Systematic Entomology Laboratory, IIBIII, ARS, U.S. Dep.
Agric. Research Agreement No. 58-32UA-9-57. Scientific Article No. A-2917, Contribution
No. 5975, of the Maryland Agricultural Experiment Station, Department of Entomology.

* Graduate Research Assistant.

112 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

BAS

Figs. 1-5. Hypomecis umbrosaria, mature larva, 25x. 1, Frontal view of head. 2, Lateral
view of head, pro- and mesothorax. 3, Lateral view of abdominal segment |. 4, Lateral view
of abdominal segment 2. 5, Lateral view of abdominal segment 3.

mecis umbrosaria (Hubner) and compare these characters with those of
related genera.

MATERIALS AND METHODS

Four mature larvae and two pupae of Hypomecis umbrosaria were ex-
amined. These were reared on a leaf diet of mixed Quercus species from a
female (voucher specimen number: GVS-77-25) collected at the Ashland
Wildlife Area, near Columbia, Boone Co., Missouri, June 2, 1977. All spec-
imens were preserved on August 7, 1977. Descriptions and drawings are
based on these specimens. A WILD M5 microscope and drawing tube at-
tachment were used in making the illustrations. Measurements are based on
the average of the available specimens.

DESCRIPTION OF MATURE LARVA AND PUPA

Larva.—Head: Height, 2.0 mm; width, 2.7 mm; color reddish brown with
areas of dark brown composed of irregular spots located principally above
level of ocelli; cuticle heavily granular; ocelli 1-3 largest, 4-6 smaller (Fig.

————

VOLUME 84, NUMBER 1 113

| 2); mandibular teeth rudimentary, upper ental surface deeply concave (Fig.
9); labrum moderately bilobed (Fig. 1); epipharynx with inner pair of heli
_ smallest; spinneret short (Fig. 11); postmentum setae short (Fig. 11); maxilla
with proximalmost seta longest (Fig. 11).

Body: Length, 35 mm; width 3 mm; mottled in shades of reddish brown:
integument finely granular with small, variable, 5- or 6-sided sclerotized,
miniature volcano-shaped structures (Fig. 8); most setae short, dark brown,
arising from small, black chalazae. Dorsal view: Large areas of light color
variably occur at intersegmental folds; prothoracic shield concave on margin
above SD setae (Fig. 2); pair of large, wrinkled, invaginating glands on A2
located between D, and SD, setae (Fig. 4), examination of internal mor-
phology of gland reveals attachment of 5 circular and 2 longitudinal muscles,
| branching tracheal trunk, and 3 glandular ducts (Figs. 12, 13); A8 with D1
setae on small, black tubercles; anal plate triangular (Fig. 10). Lateral view:
Large areas of light color variably occur about spiracles; A6 proleg paler
than body; spiracles vary in size, in the following order of decreasing mag-
nitude—T1, A8, A7, Al, and A6, but those on 2-5 of uniform size; peritreme
black, spiracular valve pink; hypoproct and paraprocts of about equal
length, extending beyond tip of anal plate. Ventral view: Elongate patches
of light color centrally located on abdominal segments; thoracic leg bases
increasing in size by twice that of preceding segment; thoracic leg bases
increasing in size by twice that of preceding segment; thoracic leg claw dark
brown (Fig. 14); crochets biordinal and complete (not reduced in size in
middle), 38—42 in number on ventral proleg, 49-51 in number on anal proleg.

Chaetotaxy: 5 SV setae on A6 proleg; L1 of Al at level of top of spiracle
(Fig. 3); D1 of A3 (Fig. 5) further anterior than on Al or A2 (Figs. 3,4); L1
of A7 at level of top spiracle; SV1 seta 3 x length of L1 on A7-9; anal
proleg with CD2 closer to level of LG3 than LG2; CPI and CP2 just above
levels of LG2 and LGI1, respectively.

Pupa.—Body: Height, 15 mm; width, 5 mm.

Head: Color dark reddish brown; epicranial suture absent.

Thorax: Wings dark brown, rest variable shades of reddish brown; pro-
thoracic femur not exposed; large, slightly elliptical, pubescent callosities
present, 2 size of eyes.

Abdomen: Color dark reddish brown, paler at interfolds; cuticle densely
punctate (Fig. 7), setae arising from sunken pits (Fig. 7); AS with ridged,
prespiracular groove (Fig. 7); dorsum of A9 with 4 small, widely spaced
teeth on caudal edge. cremaster elongate, blunt-tipped with 4 small, lateral
flanges (Fig. 6).

DISCUSSION

The pair of glands on A2, which almost certainly play a defensive role,
are of special interest because they apparently have never before been re-

114 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Desf
ROG

Figs. 6-14. Hypomecis umbrosaria. 6-7, Pupa, 50x. 6, Ventral view of cremaster. 7, Right
prespiracular furrow on abdominal segment 5. 8-14, Mature larva. 8, Enlargement of cuticle,

50x. 9, Inner view of left mandible, 50x. 10, Anal plate, 25x. 11, Ventral view of mentum,
hypopharynx, labial palpi, spinneret and maxilla, 25x. 12-13, Gland on abdominal segment 2

VOLUME 84, NUMBER 1 115

ported to occur in the Geometridae. The closely related H. gnopharia pos-
sesses the glands also, and Forbes (1948) noted that the penultimate larva
of H. buchholzaria (Lemmer) has much reduced ‘‘warts’’ at these gland
sites. Of the related genera treated by McGuffin (1977), all but Cleora in-
clude species with one or more protuberances on each side of the second
abdominal segment. Except for those of Anavitrinella pampinaria Guenée,
however, all of these protuberances are tipped with setae, usually the L1
or D2 seta. On A. pampinaria the protuberances appear as considerably
smaller versions of the glands found on Hypomecis umbrosaria as they also
lie between the D1 and SD1 setae; however, they are yet to be examined.

In all related genera (based on McGuffin, 1977), the SV4 and SV1 setae
on the first abdominal segment are above or above and posterior to the V1
seta, and the LI seta is at the bottom of the spiracle except in Anavitrinella,
Anacamptodes, and Iridopsis (three closely related genera). In Hypomecis
umbrosaria the SV4 and SV1 setae are above and anterior to the V1 seta
and the LI seta is near the top of the spiracle. Only the L1 of Anavitrinella
does not correspond to these character states, though it is higher in Hy-
pomecis umbrosaria than in the other two genera. In H. umbrosaria on the
first abdominal segment the L2 seta, second segment the SV4 seta, and third
segment the D1 seta are all located further anterior on their respective seg-
ments than in any related genera.

The anal plate of H. umbrosaria is unusual in lacking a median notch or
furrow on its anterior edge. The shape of the plate is similar to many genera
but identical to none. The tip of the plate is strongly truncate and the setae
are short.

The pupal cremaster is unique for the tribe. Nearly all species have an
obvious bifurcation resulting from a deep median groove on the cremasteral
tip and lack lateral flanges. The cremaster of Anavitrinella pampinaria re-
sembles a distorted, shortened variant of that found on Hypomecis umbro-
saria in that it possesses three pair of semi-circular ridges that appear ho-
mologous to the basal humps and lateral flanges of H. umbrosaria. Some
species of Anacamptodes have a single pair of lateral flanges but the cre-
master is rarely as elongate.

Anavitrinella is the most closely related genus to Hypomecis on the basis
of larval and pupal characters. This is supported in the larva by the location
and type of protuberances on the second abdominal segment and the chae-
totaxy, and in the pupa by the structure of the cremaster and the type of

ee

showing internal morphology. 12, Lateral view showing circular muscles and tracheal trunk,
25x. 13, Ental view showing longitudinal muscles and glandular ducts, x50. 14, Thoracic leg
claw, 50x.

116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

prespiracular furrow on the fifth abdominal segment. Anacamptodes and
Iridopsis are also genera closely related to Hypomecis, but the second ab-
dominal protuberances have different locations, and the cremaster has only
two flanges at most; however, the chaetotaxy in some species is more similar
to Hypomecis umbrosaria than to Anavitrinella. The Hypomecis adult,
however, shows no clear affinity to any one particular genus.

ACKNOWLEDGMENTS

I thank Douglas C. Ferguson, Systematic Entomology Laboratory,
USDA, and John Davidson, Department of Entomology, University of
Maryland, College Park, for reviewing the manuscript.

LITERATURE CITED

Fletcher, D. S. 1979. In Nye, I. W. B. The generic names of moths of the world. Vol. 3:
Geometridae. Trustees of the British Mus. (Nat. Hist.), London. xx + 243 pp.

Forbes, W. T. M. 1948. Lepidoptera of New York and neighboring states. Part II]. Mem.
Cornell Univ. Agrc. Exp. Stn. no. 274, 263 pp., 255 figs.

Guenée, A. 1857. Spécies général des lépidopteres. Uranides et Phalenites. Tome 9, Paris.
514 pp.

Heitzman, R. L. 1973 [1974]. An annotated checklist of the Missouri Geometridae (Lepidop-
tera). J. Res. Lepid. 12: 169-179.

McGuffin, W. C. 1977. Guide to the Geometridae of Canada (Lepidoptera). II. Subfamily
Ennominae. 2. Mem. Entomol. Soc. Can. 191 pp., 206 figs.

Rindge, F. H. 1973. A revision of the North American species of the genus Pseudoboarmia
(Lepidoptera, Geometridae). Am. Mus. Novit. no. 2514, pp. 1-27, 25 figs.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 117-127

SYMPHYTA (HYMENOPTERA) OF SRI LANKA
DAvID R. SMITH

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
_ vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Seven species of Symphyta in three families, Tenthredinidae,
_ Xiphydriidae, and Orussidae, are known from Sri Lanka. One new species,
_ Nesoselandria minuta, is described. Senoclia bilanga Rohwer is a new syn-
— onym of Anisoarthra coerulea Cameron; Netrocerus nigriceps Enderlein is
anew synonym of Eusunoxa ceylonica Malaise; Nesoselandria ceylonensis
Rohwer is a new synonym of Nesoselandria thwaitesii (Kirby), new com-
bination; and Eusunoxa nigriceps (Rohwer) is a new combination. Keys to
families and genera and to species of Nesoselandria are provided.

The sawfly fauna of Sri Lanka has never been reviewed, only descriptions
of species are scattered in the literature. Nine species have been described,
but because of new synonymy and the possibility that one species, Ani-
soarthra cyanella Cameron, may not occur on the island, seven species in
three families (Tenthredinidae, Xiphydriidae, and Orussidae) are actually
known.

The data presented here are based on specimens collected during the
Smithsonian Ceylonese Insect Project (SC), on a study of the types of
species described from Sri Lanka, and on specimens from Sri Lankan col-
lections at the University of Lund, Lund, Sweden (LUND), and the Natur-
historisches Museum Basel, Switzerland (NHMB). During the Smithsonian
Ceylonese Insect Project, five species of Tenthredinidae were collected, but
the representatives of Orussidae and Xiphyriidae recorded from Sri Lanka
were not found. Two species, Anisoarthra coerulea Cameron and Mocsarya
metallica Mocsary, are also found outside of Sri Lanka; the former occur-
ring in southern India, which has a similar sawfly fauna. Hosts are not
known for the species treated here.

KEY TO FAMILIES AND GENERA

1. Antennae inserted on ventral aspect of head, below lower margin of
eyes and below clypeus; hindwing without closed cells .... Orussidae

118 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

— Antennae inserted on anterior aspect of head, above clypeus and

between eyes; hindwing with one or more closed cells ............. 2
2. Foretibia with 1 apical spur; antenna setaceous, with 15 or more
SEEMING SO er he ny vege hae eee oh eee rene Xiphydriidae
— Foretibia with 2 apical spurs; antenna filiform, with 9-10 segments
CPrenthredimidac). 25 ov: coynevgs oo see elon 9 habe pe ee oe ce ee 3
3. Anal cell of forewing without a crossvein (Fig. 10); small black
species 3—o mm long (Selandrumac) .. 2... a2. oes eo Nesoselandria

— Anal cell of forewing with crossvein or petiolate (Figs. 11, 12); usu-
ally larger species 7-11 mm long, if about 6 mm, then mostly orange

4. Anal cell of forewing complete and with anal crossvein (Fig. 11);
tarsal claw with 2 teeth (Fig. 14) (Allantinae) ............. Eusunoxa
— Anal cell of forewing petiolate, base of vein 2A + 3A atrophied (Fig.
12); tarsal claw comblike, with 4—S teeth (Fig. 13) (Blennocampinae)
ie Cita af abs ql NSM AR eS end oo PR Ps ere are ck, fee a Anisoarthra

Family Tenthredinidae
Subfamily Blennocampinae
Anisoarthra coerulea Cameron
Bigs: 13 9515

Anisoarthra coerulea Cameron, 1876: 462 (2, d, ‘‘Ceylon’’; lectotype 2, by
present designation, in British Museum (Natural History), labeled ““Type,
H.T.,’’ “‘B.M. Type Hym. 1.363,’ “‘B.M. Type, Hym. Anisoarthra coe-
rulea (Cameron, 1876),’’ “‘Ceylon,”’ “‘Kby. p. 8, f. 21’’; other specimens
were not found).

Senoclia bilanga Rohwer, 1921: 106(@, 3, ‘‘Kollegal, 2,000 feet (about 606
meters), Coimbatore, S. India’’; holotype 2 in USNM, examined). New
synonymy.

Remarks.—Anisoarthra coerulea is large, 9-11 mm long, and is entirely
purplish black, commonly with a purplish iridescent tinge. The tarsal claws
are comblike, having 4—S teeth plus a short basal lobe (Fig. 13). The female
ovipositor and male genitalia are as in Figs. 1, 9.

Malaise (1937) treated bilanga Rohwer from southern India as a variety
of coerulea; however, since I did not find differences between them, I be-
lieve that they are the same. Four or five other species of Anisoarthra are
known from Southeast Asia.

In Sri Lanka, this species has been collected in the wet zone, rain forest
hill country, with an annual rainfall of 1952 mm as well as in drier zones
where the annual rainfall is 1500 mm.

Records.—SRI LANKA: North Central Province: Anuradhapura Dis-
trict, Padaviya, 2-8-XI-1970, O. S. Flint, Jr. (1 2; SC). North Western

VOLUME 84, NUMBER 1 119

i
i
"i
Ld
al
;

TAO

HL rym Pee
fi

Mas OT
esuill
ha
aes Ts)
<
Pa
WA
ie ass
a ae
t
ww
G d

AT SAM Lae

4
ADE rT
TMT

Figs. 1-4. Female lancets. 1, Anisoarthra coerulea. 2, Eusunoxa ceylonica. 3, Nesoselan-
dria anthracina. 4, N. thwaitesii. Figs. 3 and 4 drawn to same scale, but not same scale as
Figs: 1/2.

Province: Puttalam District, Tabbowa, 17-X-1973, black light, sea level, M.
and B. Robinson (1 ¢; SC). Central Province: Kandy District, Udawattak-
ele, 1—3-X-1973, K. V. Krombein, P. B. Karunaratne, P. Fernando (1 6;
SC); Kandy District, Udawattakele, 27—28-X-1972, P. B. Karunaratne (1
2; SC); Kandy District, Waharawata, 1-IX-1975, D. M. Davies, S. Karu-
naratne, D. W. Balasooriya (1 6; SC); Kandy District, Kandy, X-07, 9-07,
X-09, XI-06, [X-1911, 10-03 (series; USNM); Kandy District, Kandy, 30-IX-
1953, F. Keiser (1 6; NHMB); Kandy District, Kandy, Roseneath, 29-IX-
53, F. Keiser (1 2; NHMB): Kandy District, Kandy, Deiyannewela, 17-X-
53, F. Keiser (3 2, 2 6; NHMB); Dambulla, 21-XI-53, F. Keiser (3 2;
NHMB); Matele District, Nalanda, X-07 (1 2; USNM). INDIA: Karnataka:
Kollegal, 2000 ft., Coimbatore, 1-IX-17 (type-series; USNM). Tamil Nadu:
Nilgiri Hills, Kallar, 1250 ft., X-1955 (1 2; USNM).

120 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Anisoarthra cyanella Cameron

Anisoarthra cyanella Cameron, 1876: 462 (2, ¢, “‘Ceylon’’; types not
found).

Remarks.—The types should be in the British Museum, but because they
cannot be found (J. Quinlan, personal correspondence) the status of cy-
anella is uncertain. Kirby (1882) synonymized cyanella under Senoclia pur-
purata Smith (currently in the genus Senoclidea) and gave the distribution
as ‘‘Celebes; New Guinea’’ but did not mention Ceylon. The type-locality
given by Cameron may be an error. I have not seen specimens from Sri
Lanka that agree with the description of cyanella and have not seen spec-
imens of purpurata from Sri Lanka. This species, if it is the same as pur-
purata as Kirby indicated, can be distinguished from coerulea by the tarsal
claws, which have two teeth and a basal lobe, by the antenna which is
‘‘shorter, more thickened towards the apex,’ and by the whitish apex of
the foretarsus; otherwise, the size and coloration are similar in the two
species.

Subfamily Allantinae
Eusunoxa ceylonica Malaise
Figs. 2, 8, 14

Netrocerus nigriceps Enderlein, 1920: 371 (2, “‘Ceylon’’; type in Polish
Academy of Sciences, Warsaw, examined). New synonymy. Preoccupied
in Eusunoxa by Eusunoxa nigriceps (Rohwer), new combination (de-
scribed in genus Beleses, 1915).

Eusunoxa ceylonica Malaise, 1932: 147 (2, ¢, “Colombo, Ceylon’’; holo-
type in British Museum (Natural History), paratype from Swedish Mu-
seum of Natural History examined).

Remarks.—Eusunoxa ceylonica was one of the most commonly collected
sawflies in Sri Lanka and may be distinguished as follows: Orange color
with antenna, head, apex of tibia and tarsus of fore- and midlegs, and most
of femur, tibia, and tarsus of hindleg black; uniformly blackish infuscated
wings, sometimes blackish abdominal apex in male; two teeth of the tarsal
claws (Fig. 14); and anal cell of forewing with an anal crossvein (Fig. 11).
The female ovipositor and male genitalia are as in Figs. 2, 8.

Other species of Eusunoxa are found in southeastern Asia. Eusunoxa
nigriceps (Rohwer), described from southern India, is entirely black, but it
and others are separated from ceylonica primarily by differences in the
genitalia.

Eusunoxa ceylonica occurs in both open and wooded areas in both dry
and wet zones with annual rainfall ranging from 660 to 1952 mm.

Records.—SRI LANKA: Western Province: Colombo District, Colombo,

VOLUME 84, NUMBER 1 121

50 ft. elev., Museum gardens, 15-I-1977, K. V. Krombein, P. Fernando, D.
W. Balasooriya, V. Gunawardane (1 2; SC); Colombo, Museum gardens,
50 ft., 18-11-1977, P. B. Karunaratne (1 ¢; SC); Colombo, collected in
Museum gardens, 22-VI-1977 (10 6; SC), 14-VI-1977 (1 2, 1 6; SO), T.
Wijesinhe; Colombo District, Labugama Reservoir, 16-II-1975, K. V. Krom-
bein, P. B. Karunaratne, P. Fernando, S. Karunaratne (1 2; SC); Colombo
District, Labugama, Reservoir Jungle, 2-4-II-1977, K. V. Krombein, P. B.
Karunaratne, D. W. Balasooriya, V. Gunawardane (1 2; SC); Colombo
District, Ratmalana Airport, Ratmalana, 13-I-1977, K. V. Krombein, P. Fer-
nando, D. W. Balasooriya, V. Gunawardane (1 ¢; SC); Colombo District,
Kalatuwawa, Malaise trap, 7-8-VIII-1975, Y.-M. Huang, E. L. Peyton, S.
Karunaratne, D. W. Balasooriya (1 2; SC); Yakkala, 18 mls NE Colombo,
13-31-I-1962, Loc. 11, swept on veg. at ditches in paddy fields, Lund
University Ceylon Expedition 1962, Brinck-Andersson-Cederholm (1 9;
LUND). Central Province: Kandy District, Peradeniya Botanical Gardens,
I-1971, Piyadasa and Somapala (1 2; SC). Southern Province: Hambantota
District, Palatupana Tank, 15-20 ft., 18-20 Jan. 1979, Malaise trap, K. V.
Krombein, P. B. Karunaratne, T. Wijesinhe, S. Siriwardane, T. Gu-
nawardane (1 °, 1 6; SC). Eastern Province: Amparai District, Ekgal Aru
Sanctuary Jungle, 9-11 March 1979, K. V. Krombein, T. Wijesinhe, Y. L.
Jayawickrema, S. Sirwardane (2 6; SC), same data, Malaise trap added
after date (1 d6; SC). North Western Province: Rajakadaluwa, 23—VIII-1953,
F. Keiser (2 6; NHMB). Northern Province: small stream, 2 mls E Man-
kulam, 14-II-1962, Loc. 75, grassy ground, Lund University Ceylon Expe-
dition 1962, Brinck-Andersson-Cederholm (1 6; LUND).

For Loc. 11 and 75 from Lund University specimens, see Brinck et al.,
1971.

Subfamily Selandriinae
Genus Nesoselandria Rohwer

Nesoselandria is a large genus in Asia. Most species are small, 3-6 mm
long, and mostly black. Few good external characters are evident for sep-
arating species, and the small, fragile female lancet is very similar through-
out the genus. The best characters for species separation seem to be in the
male genitalia where good differences were found in the Sri Lankan speci-
mens. Malaise (1944) gave a key to species of Nesoselandria, but many
described species were omitted, and only one species, anthracina, was
recorded from Sri Lanka. Malaise also proposed two subgenera, the typical
subgenus and Corrugia. These subgenera were separated by the sculptur-
ation of the head, Corrugia having a transverse furrow and a number of
transverse carinae between the eyes on the frons and Nesoselandria lacking
these. All of the species I have seen from Sri Lanka would belong in the

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 84, NUMBER 1 123

subgenus Corrugia. Nesoselandria is also found in southern India, but all
of the specimens I examined from southern India belong in the typical sub-
genus.

KEY TO SPECIES

1. Entirely black; usually larger species, 5~6 mm long; female lancet

as in Fig. 3, with annuli indicated; male genitalia as in Fig. 7
oe Re ee ee TS oe eee Ce Soe anthracina Malaise

— Black with tarsi, at least basitarsi, and sometimes apex of tibiae
whitishusually smaller species, 3=S mm long, 3.6: sssackh ss aasee ws 2

. Distance between hindocelli, at least in male, nearly 3x diameter of

an ocellus; male genitalia as in Fig. 6; female lancet as in Fig. 4
RNAs tiie Aiea Se. bos Gan, WETMORE As A eee thwaitesii (Kirby)

— Distance between hindocelli less than 2 diameter of an ocellus;:
Ailes OSU AS BUN IG 5) n as ge ce a es oe oe canals minuta, new species

i)

Nesoselandria anthracina Malaise
Figs3)7

Nesoselandria anthracina Malaise, 1944: 14 (6, “‘Ceylon (Ohiya)’’; holo-
type in Swedish Museum of Natural History, Stockholm, examined).

Remarks.—The entirely black coloration, slightly larger size than other
species, and genitalia as illustrated are adequate for identifying anthracina.

This species is clearly endemic, being restricted to the wet zone of the
Central Highlands at altitudes of over 1700 m. The annual rainfall at Nuwara
Eliya is 2162 mm.

Records.—SRI LANKA: Central Province: Nuwara Eliya District, Hak-
gala, 6000 ft., 2-VI-1976, K. V. Krombein, S. Karunaratne, D. W. Balasoo-
riya (1 2,2 6; SC); Nuwara Eliya District, Kanda-ela Reservoir, 6200’, 1-
5-X-1970, O. S. Flint, Jr. (1 6; SC); Nuwara Eliya District, Kanda-ela, 2-
VI-1975, S. L. Wood and J. L. Petty (5 ¢; SC); Nuwara Eliya District,
Kanda-ela Reservoir, 5.6 mi SW Nuwara Eliya, 6200 ft., 10—21-II-1970,
Davis and Rowe (2 2, 2 6; SC); Nuwara Eliya District, Mt. Pidurutalagala,
6500-7500 ft., 8-X-1976, G. F. Hevel, R. E. Dietz IV, S. Karunaratne, D.
W. Balasooriya (1 2; SC); Nuwara Eliya District, Galway Nat. Reserve,
Nuwara Eliya, 6200 ft., 10 June 1978, K. V. Krombein, P. B. Karunaratne,

—

Figs. 5-9. Male genitalia; left, capsule, ventral aspect of left half; right, valve, lateral
aspect with ventral side to left. 5, Nesoselandria minuta. 6, N. thwaitesii. 7, N. anthracina.
8, Eusunoxa ceylonica. 9, Anisoarthra coerulea. Figs. 10-12, Anal cell of forewing. 10, Ne-
soselandria. 11, Eusunoxa. 12, Anisoarthra. Figs. 13-14. Tarsal claws, 13, A. coerulea. 14,
E. ceylonica.

124 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

T. Wijesinhe, L. Jayawickrema (16 2, 2 ¢ SC); Nuwara Eliya District,
Nuwara Eliya, Galway Nat. Res., 1790-1990 m, 22-23-X-1977, K. V. Krom-
bein, T. Wijesinhe, M. Jayaweera, P. A. Panawatta (1 2; SC); Pidrutalagala,
2000-2460 m, 19-XI-1953, F. Keiser (1 2, 3 6; NHMB); Hakgala, 1800-
1900 m. 31-V-1953, F. Keiser (1 6; NHMB). Province of Uva: Badulla
District, Ohiya (type and paratype).

Nesoselandria thwaitesii (Kirby), new combination
Figs. 4, 6

Aneugmenus thwaitesii Kirby, 1883: 203 (d, “‘Ceylon’’; type in British Mu-
seum (Natural History), examined).

Nesoselandria ceylonensis Rohwer, 1912: 234 (¢, ‘‘Peradeniya, Ceylon”’;
type in USNM, examined). New synonymy.

Remarks.—This species has partly white tarsi, sometimes whitish only
on the basitarsi, as does minuta, and is 4-5 mm long. Male genitalia should
be used to separate thwaitesii from minuta, although the ocellar triangle
seems to be a good character for separating males of the two species. The
hindocelli in thwaitesii are nearly three ocellus diameters apart, forming a
low triangle, whereas in minuta the hindocelli are less than two ocellus
diameters apart, producing a higher triangle. The value of the ocellar triangle
for separation of females of these two species is unknown since only the
female of thwaitesii is available for study. The types of thwaitesii and cey-
lonensis proved to be opposite sexes of the same species. Kirby (1883)
stated that thwaitesii had 10-segmented antennae, but antennae of the type
are actually 9-segmented. The type of thwaitesii is a male; this was not
stated in the original description. Malaise (1944) indicated that thwaitesii
belonged in Nesoselandria, but he was not certain.

Nesoselandria thwaitesii is also a wet zone species, but it occurs at lower
altitudes than does anthracina. The average annual rainfall for collection
sites is 2400-2700 mm.

Records.—SRI LANKA: Central Province: Kandy District, Udawatta-
kele Sanctuary, 26-30-VII-1978, Malaise trap, K. V. Krombein, T. Wije-
sinhe, V. Kulasekare, L. Jayawickrema (2 2; SC); Udawattakele, Kandy,
510-580 m, 8-10-IX-1977, K. V. Krombein, P. B. Karunaratne, T. Wije-
sinhe, M. Jayaweera (1 2; SC); Kandy District, Peradeniya (type of cey-
lonensis); Kandy, 29-X-1953, F. Keiser (1 36), 30-IX-53 (1 3), 15-X-53 (1
2), 15-XI-53 (1 2) (NHMB). Western Province: Colombo District, Labu-
gama Reservoir, 400 ft., 1-XII-1976, G. F. Hevel, R. E. Dietz IV, P. B. and
S. Karunaratne, D. W. Balasooriya (1 2; SC); Kalutara District, Agala-
watta, black light, 13-14-X-1976, G. F. Hevel R.E. Dietz IV, S. Karuna-
ratne, D. W. Balasooriya (3 2, 1 6; SC). Province of Sabaragamuwa:
Kegalla District, Kitulgala, Bandarakele Jungle, 17-18 March 1979, K. V.

VOLUME 84, NUMBER 1 125

Krombein, P. B. Karunaratne, T. Wijesinhe, L. Jayawickrema, T. Guna-
wardane (1 2; SC); Ratnapura District, Rakwana, 1500’, 20-X-1970, O. S.
Flint, Jr. (1 ¢6; SC); Ratnapura District, Ratnapura, Pompekele, 10-X-1980,
P. B. Karunaratne, T. Wijesinhe, L. Jayawickrema, V. Gunawardane (1
2; SC). Province of Uva: Lunugala, 25-X-1953, F. Keiser (2; NHMB).

Nesoselandria minuta Smith, new species
Fig. 5

Female.—Unknown.

Male.—Length, 3.7 mm. Black with tarsi whitish, apical 2 or 3 segments
of each tarsus infuscated blackish. Wings lightly, uniformly infuscated
black; veins and stigma black. Third antennal segment 1'4 length of 4th
segment. Head with transverse carinae on frons between eyes; clypeus trun-
cate; malar space linear; postocellar area nearly 5x broader than long; dis-
tance between hindocelli less than 2x diameter of an ocellus and ocellar
triangle high, nearly equilateral. Genitalia as in Fig. 5.

Holotype.—6, Province of Sabaragamuwa, Ratnapura District, labeled
‘‘Ceylon, Rat. Dist., Uggalkaltota, 350 ft., Irrigation Bungalow, 31 Jan.-8
Feb. 1970, Davis and Rowe.’’ USNM type no. 76875.

Paratype.—Central Province, Kandy District, Thawalamtenne, 2200 ft.,
7-8-IX-1980, K. V. Krombein, P. B. Karunaratne, T. Wijesinhe, L. Ja-
yawickrema, V. Gunawardane (1 ¢).

Remarks.—The genitalia of minuta are very different from those of the
other two species from Sri Lanka, and the specimens do not agree with the
other species treated by Malaise in his key (1944). The distance between the
hindocelli may be used to separate minuta from the males of thwaitesii, as
given in the key and discussed under thwaitesii, but the two species are
best separated by comparing Figs. 5—7 of the genitalia. As with the other
species of Nesoselandria, this is apparently also a wet zone species.

Family Xiphydriidae
Cingalixiphia striatifrons (Cameron)
Xiphydria striatifrons Cameron, 1905: 70, pl. A, fig. 1 (2, **Pundalu-oya”’;
type in British Museum (Natural History), examined).

Remarks.—This is the only species of Xiphydriidae known from Sri Lan-
ka. I have not seen specimens other than the type. The type-locality is in
Central Province, Nuwara Eliya District.

Family Orussidae
Mocsarya metallica (Mocsary)

Oryssus metallicus Mocsary, 1896: 1 (2, *“Sumbawa in Insulis Sundaicis’’;
type probably in the Hungarian Museum of Natural History, Budapest).

126 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Remarks.—I have not seen specimens of Orussidae from Sri Lanka. Moc-
sarya metallica was described from Indonesia, but it was recorded from Sri
Lanka by Bingham (1908) and later by Benson (1935). Bingham stated that
his specimen from Sri Lanka corresponded closely with Mocsary’s descrip-
tion of metallica, but he noted several differences in the amount of infus-
cation of the forewings and the coloration of the legs. There is no doubt that
an orussid does occur in Sri Lanka, but a comparison of Bingham’s speci-
men with the type of metallica will be necessary to determine if Bingham’s
specimen is actually that species.

ACKNOWLEDGMENTS

I thank the following for allowing study of specimens: K. V. Krombein,
Principal Investigator, Smithsonian Ceylonese Insect Project, Department
of Entomology, Smithsonian Institution, Washington, D.C.; E. Kierych,
Polska Akademia Nauk, Instytut Zoologii, Warszawa, Poland; Per Inge
Persson, Naturhistoriska Riksmuseet, Stockholm, Sweden; J. Quinlan, Brit-
ish Museum (Natural History), London; M. Brancucci, Naturhistorisches
Museum Basel, Switzerland; and Per Brinck, University of Lund, Sweden.
Other specimens are in the National Museum of Natural History, Smith-
sonian Institution, Washington, D.C. (USNM).

LITERATURE CITED

Benson, R. B. 1935. On the genera of the Orussidae (with an account of the African species
(Hymenoptera Symphyta)). Occas. Pap. Rhodesian Mus. No. 4, 10 pp.

Bingham, C. T. 1908. Appendix to Mr. Wickwar’s paper. Spolia Zeylan. 5: 122-123.

Brinck, P., H. Andersson, and L. Cederholm. 1971. Report No. 1 from the Lund University
Ceylon Expedition in 1962. Entomol. Scand. Suppl. 1: II-XXXVI.

Cameron, P. 1876. Descriptions of new genera and species of Tenthredinidae and Siricidae,
chiefly from the East Indies, in the collection of the British Museum. Trans. Entomol.
Soc. Lond. 1876: 459-471.

———.. 1905. On the phytophagous and parasitic Hymenoptera collected Mr. E. Ernest Green
in Ceylon (first paper). Spolia Zeylan. 3: 67-97.

Enderlein, G. 1920(1919). Symphytologica II. Zur Kenntniss der Tenthredininen. Sitzb. Ge-
sell. Naturf. Freunde Berl., No. 9, pp. 347-374.

Kirby, W. F. 1882. List of Hymenoptera in the British Museum, Vol. I. London. 450 pp.

. 1883. Notes on new or little-known species of Hymenoptera, chiefly from New Zea-

land. Trans. Entomol. Soc. Lond. 1883: 199-203.

Malaise, R. 1932. A new sawfly from Ceylon (Hym. Tenthredinidae). Ceylon J. Sci., Sect. B,

17: 147-148.

. 1937. New Tenthredinidae mainly from the Paris Museum. Rev. Fr. Entomol. 4: 43-

54.

———.. 1944. Entomological results from the Swedish Expedition 1934 to Burma and British
India, Hymenoptera: Tenthredinoidea collected by René Malaise, the Tenthredinoidea
of South-Eastern Asia. Ark. Zool. 35A: 1-58.

Mocsary, A. 1896. Species Hymenopterorum magnificae novae in collectione Musaei Natio-
nalis Hungarici. Természetr. Fiizetek 19: 1-8.

VOLUME 84, NUMBER | 127

Rohwer, S. A. 1912. Notes on sawflies, with descriptions of new species. Proc. U.S. Natl.

Mus. 43: 205-251.

. 1915. Some Oriental sawflies in the Indian Museum. Rec. Indian Mus. 11: 39-53.

—. 1921. Notes on sawflies, with descriptions of new genera and species. Proc. U.S.
Natl. Mus. 59: 83-109.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, p. 127

NOTE

Authorship of the Family Name Erotylidae
(Coleoptera)

Boyle (1956. Bull. Am. Mus. Nat. Hist. 110) in his monograph of the
Erotylidae of North America cited Lacordaire (1842. Monographie des Er-
otyliens) as the author of the family group name Erotylidae. Sen Gupta
(1969. Proc. Zool. Soc. Calcutta 22) cited Guerin-Méneville based on the
usage in 1841 (Rev. Zool. 1841).

The earliest usage of a family group name for Erotylidae I have seen,
however, is by Leach in the Entomology Section of Brewster's Edinburgh
Encyclopaedia published in 1815. He used the heading “‘Tribe X. Erotyl-
ides,’ and beneath it ‘‘Family I. Erotylida.’’ His usage of the vernacular
and subsequent latinization and acceptance by later authors is consistent
with Article lle, International Code of Zoological Nomenclature. The name
Erotylidae should therefore be credited to Leach, 1815.

John M. Kingsolver, Systematic Entomology Laboratory, IIBIII, Agri-
cultural Research Service, USDA, % National Museum of Natural History,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 128-133

SEASONAL HISTORY OF THE WHITE PEACH SCALE,
PSEUDAULACASPIS PENTAGONA (TARG.-TOZZ.)
(HOMOPTERA: DIASPIDIDAE), IN
NORTHEASTERN PENNSYLVANIA

JAMES F. STIMMEL

Bureau of Plant Industry, Pennsylvania Department of Agriculture, Har-
risburg, Pennsylvania 17110.

Abstract.—The seasonal history of the white peach scale, Pseudaulacas-
pis pentagona (Targioni-Tozzetti), was studied on Kwanzan cherry, Prunus
serrulata Lindl., during 1980. In northeastern Pennsylvania the scale was
bivoltine with fertilized adult females overwintering. Oviposition began 15
May for the first generation and 22 July for the second generation; females
produced an average of 27.2 eggs and 78 eggs per generation, respectively.
Crawler emergence began 20 May and 28 July; adults appeared 8 July and
3 September. Natural enemies encountered in this study were the twospot-
ted lady beetle, Adalia bipunctata (L.) (Coccinellidae), and the wasp par-
asite Encarsia berleseii (Howard) (Aphelinidae).

The white peach scale, Pseudaulacaspis pentagona (Targioni-Tozzetti),
probably native to China (Gossard, 1902), was described from Italy in 1886.
It has been introduced repeatedly into the Western Hemisphere, particularly
the United States. The apparent first record of this diaspidid in the conti-
nental U.S. was from Florida in 1889 (Van Duyn and Murphey, 1971), or
Thomasville, Ga. in the late 1800’s (Yonce and Jacklin, 1974). It was also
imported into Georgia in 1902 (Bobb et al., 1973). The scale became a major
pest of oleander in Bermuda after its introduction in 1917 (Bennett, 1956),
and there was concern that oleander might be eliminated. So great was this
fear that the Bermudian government passed the White Scale Act in 1921 in
an attempt to control the pest. In Florida the demise of the peach industry
in the early 1900’s can be attributed partially to the white peach scale. In
orchard situations it can be as destructive as San José scale, Quadraspidi-
otus perniciosus (Comstock) (Bobb et al., 1973). Since white peach scale is
a polyphagous feeder it is a pest not only of fruit trees but many ornamen-
tals. Ferris (1937) noted that it is capable of attacking nearly any nonconi-
ferous host. While this may seem too broad a statement, Borchsenius (1966)

VOLUME 84, NUMBER 1 129

recorded it from 54 families and 121 genera of hosts, and Dekle (1976) lists
278 hosts in Florida, including juniper, a conifer. In Pennsylvania the most
common hosts are flowering cherry (Prunus spp.) and privet (Ligustrum
spp.).

The life history of white peach scale is well known in the southern states.
It is a multivoltine species having as many as four generations per year in
Florida (Kuitert, 1968) and three in North Carolina (Smith, 1969). The lit-
erature, however, contains little on its life history in the northern states.

In November 1979, my attention was called to a heavy infestation of white
peach scale on Kwanzan cherry (Prunus serrulata Lindl.) in a park at
Wilkes-Barre, Luzerne Co., Pa. Most of the nearly 100 trees in the park
were severely infested and had suffered various degrees of dieback. Ac-
cording to the county extension agent, most of the ornamental cherries in
the Wyoming Valley were infested with the scale and control attempts with
dormant oil had been unsuccessful. In attempting to make a control rec-
ommendation, I could find no exact dates of crawler emergence. For that
reason, I initiated a seasonal history study of this diaspidid in northeast
Pennsylvania.

METHODS

Seasonal history data were compiled from weekly samples of scale-in-
fested Kwanzan cherry taken in Wilkes-Barre, Pa. Sampling began in late
March while overwintering scales were still dormant and continued into the
following October when the scales ceased development. Throughout the
winter, monthly samples were taken to insure that the females were not
developing. In the laboratory, specimens were removed and slidemounted
in Hoyer’s medium and determined to stage with a phase-contrast micro-
scope. Specimens were selected by starting at a random point, then slide-
mounting all nearby individuals, including eggs, until a minimum of 100
specimens were mounted. Approximately 3000 scales were mounted and
identified. Parasites were collected by rearing them from scale-infested twigs
in large, covered plastic dishes and placing them into 70% ethanol as they
emerged. No predators were encountered in the laboratory samples.

Exact length of stages could not be determined by my sampling proce-
dure, but the precise length of stadia is known (Ball, 1980). My study fo-
cused on monitoring a population under field conditions to obtain actual
dates of oviposition and crawler emergence. Egg counts were made by lifting
the scale’s armor and counting the eggs beneath. Empty chorions were not
counted. Length of the egg stage was determined by observing newly de-
posited eggs until they hatched. These eggs were collected by excising an
ovipositing female from the host, removing eggs individually, and placing
them in separate plastic dishes for observation. Eggs were checked several
times daily until they hatched.

130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

SEASONAL History (Fig. 1)

Pseudaulacaspis pentagona overwinters as fertilized adult females. Eggs
could be seen in cleared females by 14 April. Shortly before depositing eggs,
females produced crawler emergence flaps on their armor at the end op-
posite the exuviae. Oviposition began 15 May, when the eggs of 20 females
were counted, and the average number of eggs held per female was 27.2
(range 7-46). Crawlers began to emerge about 20 May, as field observations
revealed only three active crawlers on that date. Settled crawlers appeared
in the 28 May sample, but most certainly were present by 22 May, owing
to the short life of active crawlers. By 28 May, settled crawlers composed
more than 50% of the sample. By 5 June, most females had finished ovi-
positing and had died. Settled crawlers were dominant until 19 June when
the entire sample consisted of second-instars. Prepupal and pupal males
were present in the 25 June sample. Adult males and females of this first
generation appeared 8 July, and mated shortly after attaining maturity. By
17 July, cleared females again contained eggs. Second generation eggs were
present in the 22 July sample; each ovipositing female had laid 12.4 eggs by
this time. The following week (31 July) eggs, active crawlers, and settled
crawlers could be found. At this time each of 18 females held an average of
78 unhatched eggs. Second-instars of both sexes appeared 14 August, with
males more abundant. Prepupal and pupal males were present in the 27
August sample, and adults of both sexes were found the next week (3 Sep-
tember). By 10 September all males had died, and only fertilized adult fe-
males were found in the remainder of the year’s samples. By late September
almost 33% of the females were parasitized; until this time the rate of par-
asitism had been low.

Two species of parasites, both Aphelinidae, were reared from the weekly
samples: Encarsia berleseii (Howard), a primary parasite, and Marietta
carnesi (Howard), a hyperparasite. The only predator observed was the
twospotted lady beetle, Adalia bipunctata (L.) (Coccinellidae). Both adults
and larvae were observed feeding on scales.

DISCUSSION

While eggs and crawlers of white peach scale are known to exhibit sexual
dimorphism in color (Bennett and Brown, 1958; Simmonds, 1958; Kuitert,
1968; Van Duyn and Murphey, 1971), all eggs encountered during this study
were coral, although of slightly various shades. Crawlers were light yellow
and sexes could not be separated solely on the basis of color. Heavy con-
centrations of males could be seen on the undersides of limbs and branches,
whereas most females settled on the upper portions. The males were so
thick that their armor made the undersides of the infested limbs look as
though they had been plastered with snow.

VOLUME 84, NUMBER 1 131

Percent of Individuals in Population

FIGURE 1

50
25
Active ik
Crawlers
25
50
25
Settled
aS Crawlers ye
50
25 Second
Instar
25 Females

Second
Instar
Males

Adult Ow.
Females

14°21 28,5 752128 ,5 9 19 °25);2 “s 17) 22 (Sin 7 14. 20) 277 3) 1019 as
ee fe May ay te Sais. July eae August September

Date

Fig. 1. Seasonal history of white peach scale on Prunus serrulata in northeastern Penn-

sylvania (0.w. = overwinters).

132 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Generations of this species were well defined, unlike the overlapping gen-
erations of the multivoltine scale Fiorinia externa Ferris (Stimmel, 1980).
There are several reasons for this. White peach scale overwintered in only
one form, fertilized females, so that in spring all individuals continued to
develop from the same stage. The oviposition period was relatively short,
so that almost all eggs hatched within a period of a few days. Finally, the
females died upon completing oviposition and could not be confused with
newly maturing females of the following generation.

Peak oviposition took place during the week of 15 May for the first gen-
eration and 22 July for the second. Eggs hatched in an average of four days
at laboratory temperatures (23°C). Periods of greatest crawler emergence
occurred one week after oviposition peaks, 28 May for generation one and
31 July for generation two.

Populations build quickly because this species is so prolific and has two
generations per year. Natural enemies simply cannot keep up with an ex-
ploding white peach scale population. Damage by this insect is typical of
the family: dieback of smaller twigs, progressing to larger branches and then
entire trees, if left unchecked.

White peach scale is uncommon, though not rare, in Pennsylvania. It has
been taken primarily on Ligustrum and Prunus in various locations through-
out the state. It has been taken only in ornamental plantings, never in or-
chards. Infestations appear to be sporadic throughout Pa., with the excep-
tion of the Wilkes-Barre population, which covers the entire Wyoming
Valley.

Recently, Kawai (1980) split Pseudaulacaspis pentagona into two
species, pentagona (Targ.-Tozz.) and prunicola (Maskell). Based on illus-
trations presented by Kawai, the two can be separated by the structure of
the pygidial gland spines, which are fringed in pentagona, simple and spike-
like in prunicola. Some speculation exists among U.S. workers concerning
the validity of the species separation, and the presence of prunicola in the
United States has not been established'. Specimens from the Wilkes-Barre
population appear to be the prunicola type with simple gland spines. How-
ever, because of the uncertainty surrounding the status of these species, I
have treated the scale in this paper as P. pentagona. Slidemounted voucher
specimens from the Wilkes-Barre population have been deposited in the
Coccoidea collection of the U.S. National Museum at the Beltsville Agri-
cultural Research Center—West, Beltsville, Maryland.

ACKNOWLEDGMENTS

I am grateful to Paul R. Heller, extension entomologist at the Pennsyl-
vania State University, for bringing the Wilkes-Barre infestation of white

" Personal communication: Douglass R. Miller and Sueo Nakahara.

VOLUME 84, NUMBER 1 133

peach scale to my attention and for his assistance during the study. I also
thank the PDA-BPI nursery inspectors from Region III who periodically
submitted samples. Tova Rivnay, Cooperating Scientist, Systematic Ento-
mology Lab., USDA, BARC-East, Beltsville, Maryland, kindly identified
the parasitic Hymenoptera collected during this study. Douglass R. Miller
and Sueo Nakahara, Systematic Entomology Lab. and APHIS—PPQ, re-
spectively, USDA, BARC-West, Beltsville, Maryland, provided helpful in-
formation concerning the taxonomic status of the species dealt with in this
paper. Finally, thanks are due my colleagues Karl Valley and A. G. Wheel-
er, Jr., who reviewed the manuscript and offered many suggestions for its
improvement.

LITERATURE CITED

Ball, J. C. 1980. Development and fecundity of the white peach scale at two constant tem-
peratures. Fla. Entomol. 63: 188-194.

Bennett, F. D. 1956. Some parasites and predators of Pseudaulacaspis pentagona (Targ.) in
Trinidad, B.W.I. Can. Entomol. 88: 704-705.

Bennett, F. D. and S. W. Brown. 1958. Life history and sex determination in the diaspine
scale, Pseudaulacaspis pentagona (Targ.) (Coccoidea). Can. Entomol. 90: 317-324.

Bobb, M. L., J. A. Weidhaas, Jr., and L. F. Ponton. 1973. White peach scale: life history
and control studies. J. Econ. Entomol. 66: 1290-1292.

Borchsenius, N. S. 1966. A catalogue of the armoured scale insects (Diaspidoidea) of the
world. Akad. Nauk SSSR Zool. Inst. Leningrad. 449 pp.

Dekle, G. W. 1976. Florida armored scale insects. Arthropods Fla. Neighboring Land Areas,
Vol. 3, 343 pp.

Ferris, G. F. 1937. Atlas of the scale insects of North America. Series I. Serial Nos. SI-1 to
SI-136. Stanford University Press, California.

Gossard, H. A. 1902. Two peach scales. Fla. Agric. Exp. Stn. Bull. 61: 492-498.

Kawai, S. 1980. Scale insects of Japan in colors (Illus.). Nat. Soc. for Agricultural Educ.
TaiNippon Printing Co., Inc., Tokyo, Japan. 455 pp.

Kuitert, L. C. 1968. Observations on the biology, bionomics, and control of white peach
scale, Pseudaulacaspis pentagona (Targ.). Proc. Fla. State Hort. Soc., 1967 80: 376-
381.

Simmonds, F. J. 1958. The oleander scale, Pseudaulacaspis pentagona (Targ.) (Homoptera,
Coccidae) in Bermuda. Dep. Agric. Bermuda. Bull. No. 31. 44 pp.

Smith, C. F. 1969. Controlling peach scale. Res. Farming (N.C. Agric. Exp. Stn.) 28: 12.

Stimmel, J. F. 1980. Seasonal history and occurrence of Fiorinia externa Ferris in Pennsyl-
vania (Homoptera: Diaspididae). Proc. Entomol. Soc. Wash. 82: 700-706.

Van Duyn, J. and M. Murphey. 1971. Life history and control of white peach scale, Pseu-
daulacaspis pentagona (Homoptera: Coccoidea). Fla. Entomol. 54: 91-95.

Yonce, C. E. and S. W. Jacklin. 1974. Life history of the white peach scale in central Georgia.
J. Ga. Entomol. Soc. 9: 213-216.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 134-137

A NEW SPECIES OF ZALE HUBNER FROM TEXAS AND NEW MEXICO
(LEPIDOPTERA: NOCTUIDAE: CATOCALINAE)

ANDRE BLANCHARD AND JOHN G. FRANCLEMONT

(AB) 3023 Underwood Street, Houston, Texas 77025; (JGF) Department
of Entomology, Cornell University, Ithaca, New York 14853.

Abstract.—A new species of Zale, chisosensis, related to Zale corvus
(Schaus) from Mexico, is described from Texas and New Mexico. The
species is uncommon in collections, a reflection of its limited range in the
southwestern United States.

The following species is poorly represented in collections, and it has usu-
ally been misidentified as either Zale undularis (Drury) or Zale coracias
(Guenée). When the junior author first saw it, he thought it was Zale corvus
(Schaus).

Zale chisosensis Blanchard and Franclemont, New Species

Diagnosis.—Among the species of Zale that occur north of the Mexican
Border this species is superficially most similar to undularis (Drury, 1770
[1773], 1: 19), but it differs in the course of the subterminal line which is
produced into sharp points filled with black between veins R; and M, and
between veins Mg and Cu,. It lacks the conspicuous pale spot in the sub-
terminal line between veins M, and Mg, but it has a line of white scales on
the outer side of the reniform. The most closely allied species is the Mexican
species corvus (Schaus, 1901: 78) with which it agrees in all details of pat-
tern. The male genitalia of the two species differ in many ways; those of
corvus have the left valve with a large, subquadrate projection before the
apex which is drawn out into an acute point, and the right valve broad
throughout, tapering gradually to a pointed apex, and with a small, pointed
lobe on the costal margin before the apex. The aedoeagus of corvus is more
sharply angled, forming almost a right-angle. The female genitalia of the two
species are similar, but the differences are conspicuous; the pointed angle
of the ventral plate of corvus is on the right side not the left, and the narrow,
lateral plate of corvus is truncate at the apex not pointed as in chisosensis.

Description.—Black with a dull luster and a dusting of yellowish white

VOLUME 84, NUMBER 1 135

es)
A

Figs. 1-3. Zale chisosensis. 1, Paratype, Mt. Locke, 6700 feet, Davis Mountains, Jeff Davis
Co., Texas, 10 June 1969, A. & M. E. Blanchard; collection A. Blanchard. 2, Male genitalia,
aedoeagus in situ; paratype; genitalia slide AB 3712. 3, Female genitalia; paratype; genitalia
slide AB 3711.

136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

scales, area beyond subterminal line appearing smoother and with a slight
bluish luster; transverse lines more intense black; costa of forewing with
white spots near inception of lines; basal line double, curved, undulate,
reaching base in fold; antemedial line curved, undulate, starting on costa 4
from base, reaching inner margin at '/s from base, double, elements widely
spaced, incurved on veins R and 2nd A; median line angled outward below
costa, somewhat undulate, appearing triple on lower 2 of wing; postmedial
line angled below costa, incurved toward middle of reniform, undulate to
inner margin, starting on costa %4 from base, reaching inner margin %4 from
base, double, elements fine; subterminal line conspicuous, undulate, more
or less parallel to outer margin, sharply angled outward between veins R,
and M, and between veins M; and Cu,, points filled with intense black,
some white or yellowish white scaling on outside of line between the two
points; a series of vague, fine, vertical striae in terminal area; terminal line
represented by pale scales and by pale points with black scales on inner
sides between veins; outer margin evenly scalloped, incurved between
veins; orbicular punctiform, black; reniform, narrow, elliptical, vaguely de-
fined by black, a line of white scales on outer side. Hindwing with markings
of forewing continued; antemedial line and median line represented by broad
shades; postmedial line double; subterminal line sharply angled outward
between veins M; and Cu,, point black filled; terminal area and terminal
line much as on forewing. Some specimens show a brownish flush on the
forewing between the median and postmedial lines above vein 2nd A and
below Cu,; the type is such a specimen.

Expanse: Forewing length 23-25 mm.

Male genitalia: As figured.

Female genitalia: As figured.

Type.—d; Mt. Locke, 6700 feet, Davis Mountains, Jeff Davis County,
Texas, 10 June 1969, A. & M. E. Blanchard; collection of the U.S. National
Museum of Natural History.

Paratypes.—6 3, 3 2; Mt. Locke, 6700 feet, Davis Mountains, Jeff Davis
County, Texas, 10 June 1969 (3), 5 July 1969 (1), 19 July 1971 (1), A. & M.
E. Blanchard; Green Gulch, Big Bend National Park, Brewster County,
Texas, 27 June 1965 (1), 12 May 1972 (1), A. & M. E. Blanchard, 10 May
1972 (2), J. G. Franclemont; 5 in collection A. Blanchard; 4 in collection J.
G. Franclemont.

In addition to the type-series from Texas, there are three specimens from
New Mexico, a male from Beartrap Canyon Camp Ground, San Mateo
Mountains, 8000 feet, 11 April 1971, collected by Richard Holland (in the
Canadian National Collection), and two females from Near Hot Springs,
Las Vegas [San Miguel County], 7000 feet, August 1882, collected by F. H.
Snow (in the Cornell University Collection). These two females lack the

VOLUME 84, NUMBER 1 137

white scaling and appear entirely black; they were purchased from Snow by
J. H. Comstock as coracias and stood under that name until recently.

ACKNOWLEDGMENTS

We wish to thank J. D. Lafontaine for the loan of the specimen from the
Canadian National Collection and R. W. Poole of the Systematic Entomol-
ogy Laboratory, U.S. Department of Agriculture, Washington, D.C., for
reviewing the manuscript.

The photographs are by the senior author.

LITERATURE CITED

Drury, D. 1770 [1773]. Illustrations of Natural History, Volume 1. B. White, London.
Schaus, W. 1901. New Species of Noctuidae from Tropical America. Ann. Mag. Nat. Hist.
(7) 8: 77-99.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 138-148

FIRST-INSTAR LARVAE OF THE UNIFORMA GROUP OF THE
GENUS EPICAUTA (COLEOPTERA, MELOIDAE)

RICHARD B. SELANDER AND Nancy J. AGAFITEI

(RBS) Professor, Department of Genetics and Development, University
of Illinois at Urbana-Champaign, Urbana, Illinois 61801; (NJA) 418 Jackson
Avenue, Saint Charles, Illinois 60174.

Abstract.—The Uniforma Group of the subgenus Macrobasis of Epicauta
is characterized on the basis of the first-instar larvae of Epicauta uniforma
Werner, E. leoni Duges, E. bipunctata Werner, and E. niveolineata (Haag-
Rutenberg), which are described and figured from samples from Texas, State
of Mexico, Sinaloa, and Oaxaca, respectively. Epicauta niveolineata is set
apart from the other species of the group by a number of distinctive larval
characteristics, including larger body size, marked elongation of the head,
and apical branching of the major body setae. In the larval stage, the Uni-
forma Group is more similar to the Albida Group than to the Fabricii Group
of Macrobasis. The larva of nominal E. uniforma described previously by
Werner is not separable from that of E. bispinosa Werner in the nominate
subgenus of Epicauta.

The Uniforma Group, as defined by Werner (1954, 1958), contains 14
species of blister beetles of the subgenus Macrobasis LeConte of the genus
Epicauta Dejean. So far as known, the group is confined to North America,
where it ranges from Arizona and Colorado south to Chiapas, Mexico.

In this, the first in a series of articles on the taxonomy and bionomics of
the Uniforma Group, we describe and key the first-instar larvae of Epicauta
uniforma Werner, E. leoni Dugés, E. bipunctata Werner, and E. niveoli-
neata (Haag-Rutenberg) and assess the taxonomic implications of the data
provided by them.

MATERIALS AND METHODS

Larvae of the Uniforma Group used in this study eclosed from eggs laid
in the laboratory by female beetles collected in the field. Epicauta uniforma
larvae were reared from six egg masses, E. leoni from seven, E. bipunctata
from 17, and E. niveolineata from three. Each species is represented by the
progeny of several females. Although several hundred larvae of each species

VOLUME 84, NUMBER 1 139

were examined, only a small proportion was mounted on slides for detailed
study.

Egg masses were incubated individually in cotton-stoppered 3-dram glass
vials in darkness at 27°C and 100% RH in a commercial environmental
chamber. First-instar larvae were killed in 70% ethyl alcohol (in water) a
few days after eclosion. Those selected for detailed study were dehydrated
through an alcohol series, cleared in oil of wintergreen and toluene, and
mounted on slides in Harleco Synthetic Resin. Measurements and propor-
tions specified in the descriptions refer to mounted specimens. Length of
the head is measured from the posterior margin of the head capsule to the
posterior margin of the labrum. Color is described from unmounted speci-
mens in alcohol.

In addition to the larval materials described above, we have examined 38
first-instar larvae from Cochise County, Arizona, representing the bulk of
the material on which Werner’s (1962) description of nominal E. uniforma
was based and several lots of larvae of E. bispinosa Werner obtained from
adults collected in West Texas by the senior author and associates.

LARVA OF EPICAUTA UNIFORMA

The larva described by Werner (1962) as that of Epicauta uniforma from
specimens obtained from eggs laid by a female or females collected 20 miles
east of Pearce, Cochise County, Arizona, on August 27, 1958, differs from
the larvae of E. leoni, E. bipunctata, and E. niveolineata and from the larva
described herein as that of E. uniforma in the majority of characters that
have proved to be taxonomically significant in the genus Epicauta. In Wer-
ner’s larva the cuticle is pale and weakly sclerotized, the head is strongly
constricted basally, the mandibles are unusually slender and elongate and
set with numerous minute, truncate teeth (about 12 visible in outline), the
maxillary and labial palpi are slender and elongate, the sensory area of
segment III of the maxillary palpus is confined to the distal “3 of the segment
and bears only a few papillae, the antenna is elongate, spinelike evaginations
at the bases of the posterior marginal setae of abdominal terga I—VIII are
only weakly developed, the coxae are short, and the number of setae on the
head and tibiae is reduced. The posterior marginal row of setae on the
abdominal terga, containing 10 setae, is unusual in that the third and fifth
setae on each side, counting from the midline, are only about 2 as long as
the others.

Selander (1981b) recently advanced the hypothesis that several char-
acteristics of Werner’s E. uniforma larva represent adaptations associated
with an evolutionary change from larval predation on grasshopper eggs,
which is the norm for the genus Epicauta, to predation on the eggs of blister
beetles. In this connection, Selander called attention to the similarity be-

140 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tween Werner’s larva and the larva of E. bispinosa recently described by
Pinto (1975). Further study of these larvae indicates that minor anatomical
differences between them, previously regarded by us as diagnostic, do not
hold consistently. As a result, we are presently unable to distinguish Wer-
ner’s larva from the larva of E. bispinosa. Whether we are dealing with a
remarkable case of convergence between species in separate subgenera or
an instance of mistaken association of larvae of E. bispinosa with adults of
E. uniforma remains to be seen.

DESCRIPTION OF THE GROUP

The following description of the Uniforma Group is based on larvae of
E. uniforma, E. leoni, E. bipunctata, and E. niveolineata, as we have iden-
tified them. Because of the questionable specific identity of the larva de-
scribed as that of E. uniforma by Werner (1962), we omit that larva from
further consideration. In any event, it would be virtually impossible to frame
a succinct description of the group that would include Werner’s larva and
still have diagnostic value.

Body surface (cuticle) finely reticulate. Sides of head parallel or subpar-
allel except at immediate base, where they are abruptly convergent; 13 (9
major, 4 minor) setae on each side of dorsum of head outside frontal area.
Antenna moderately robust; segment II at least 142x as long as IIi, with
anteroventral seta much shorter than posteroventral seta; sensory organ at
least as wide and as long as segment III; terminal seta of III at least 14x
as long as segment II. Mandible robust; proximal seta much shorter than
distal one. Maxillary palpus with lateral margin of segment III not expanded
basally; sensory area of segment III overlapping at least slightly onto ventral
surface at apex; papillae of sensory area of III moderately stout, numerous.
Labial palpus with segment II as least 2x as long as greatest width, bearing
one seta, which reaches apex of segment. Thorax less than % as long as
body; pronotum about as long as meso- and metanotum combined; line of
dehiscence well developed and complete on pro- and mesonotum, weakly
indicated and incomplete on metanotum. Abdomen with pleurites ventral;
pleurite V somewhat wider than long, with spiracle in lateral 4; spiracle I
slightly larger than II, II-VIII nearly equal in size; sterna I-VII weakly
sclerotized, VIII-IX strongly so; spinelike evaginations absent at bases of
setae of median transverse row on terga I-VIII; posterior marginal setae less
than 2* as long as those of median transverse row; tergum V more than
342 as wide as long. Legs with coxae 22x as long as wide; tibia with 7
setae in anterodorsal row, 6-7 in posterodorsal row, 4—5 in anteromedian
row; tarsungulus I with longer seta reaching at least apical 4.

SYSTEMATIC RELATIONSHIPS

Epicauta uniforma, E. leoni, and E. bipunctata form a homogeneous
subgroup which differs from E. niveolineata in nearly two-thirds of the

VOLUME 84, NUMBER 1 141

characters of larval anatomy that we have studied. Some of the more ob-
vious differences between the subgroups involve the number of mandibular
teeth, conformation of the maxillary palpus, number of posterior marginal
setae on the abdominal terga, and form (branched or not) of the major body
setae. The number of posterior marginal setae on the abdominal terga is
compelling evidence of a close phylogenetic relationship between the
species of the first subgroup. Epicauta uniforma and E. bipunctata each
has 12 setae, E. /eoni usually 12 but occasionally 14. Elsewhere in the genus
Epicauta only E. conferta (Say), E. leopardina (Haag-Rutenberg), and E.
floydwerneri Martinez, in the nominate subgenus, have more than 10 setae
(12-14 in the first two species, 14 in the last) (Agafitei and Selander, 1980;
Selander, 1981a). Consequently, it is reasonable to regard the presence of 10
setae in E. niveolineata as the primitive condition for the Uniforma Group
and the presence of 12 and 14 in the first subgroup as progressively derived
character states.

Within the first subgroup, E. uniforma and E. leoni are especially similar
in appearance in that they lack a distinct abdominal color pattern. Epicauta
bipunctata has a well developed pattern produced by darkening of the
pronotum, metanotum, abdominal terga I and II, and the caudal terga and
in this regard closely resembles E. niveolineata. However, E. bipunctata
shares with E. uniforma the development of toothlike posterior extensions
of the cuticular reticulations of the abdominal pleurites not evident in either
E. leoni or E. niveolineata. These evaginations are weakly and inconsis-
tently developed in E. uniforma. In E. bipunctata they are strongly devel-
oped not only on the pleurites but on the sides of the terga as well. In
other characters varying within the first subgroup FE. uniforma tends to
agree more frequently with E. bipunctata than with E. leoni and therefore
is perhaps more closely related to the former species (Selander and Agafitei,
in press).

The shape of the head in E. niveolineata is unique in the genus Epicauta
and undoubtedly a derived characteristic. Branching of the body setae is
assuredly a derived characteristic also. It is known to occur elsewhere in
the genus only in two New World species of the Vittata Group [E. temexa
Adams and Selander and E. monachica (Berg)] and in the African E. al-
bovittata (Gestro) (Cros, 1938; MacSwain, 1956; Agafitei and Selander,
1980).

Of the several groups of Macrobasis currently recognized (Werner, 1954;
Selander and Mathieu, 1969), only two, the Albida and Fabricii groups, have
been described previously in the larval stage. Larvae of the Uniforma Group
more closely resemble those of the Albida Group. No single larval char-
acteristic is unique to the Uniforma and Albida groups, but the following
combination of characteristics is diagnostic: Head not or only slightly con-
stricted behind middle; mandibles robust, with relatively few large round
teeth visible in outline; segment I of labial palpus with one seta; spiracles

142 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Hatching time of eggs (at 27°C) and number of eggs per mass.
Hatching Time (days)* Eggs per Mass?
Species Mean SE N Mean SE N
E. uniforma LS7. 49 6 96.8 11.44 5
E. leoni 24.1 9 7 111.2 5.44 5)
E. bipunctata 19.5 3) 12 356.8 14.95 9
E. niveolineata 40.0 1.00 3 Sife3} 18.28 3

@ MSE = 1.72, F(G,24) = 256.33, P < .00001.
> MSE = 1183.77, F(3,18) = 88.59, P < .00001.

ventral; femur I with seven lanceolate setae; spinelike evaginations absent
at bases of setae in median transverse row of abdominal terga I—VIII; ab-
dominal terga uniform in color or, if patterned, with at least terga VII and
VIII dark; abdominal sternum VII weakly sclerotized.

Larvae of the Albida Group are easily distinguished from those of the
Uniforma Group by the length of the setae on the abdominal terga. In par-
ticular, the setae in the posterior marginal row of tergum V are about as
long as the tergum in the Albida Group and never more than % as long in
the Uniforma Group.

BIONOMICS

Data on the hatching time (at 27°C) of eggs and number of eggs per mass
for our samples of species of the Uniforma Group are summarized in Table
|. Hatching times for E. uniforma, E. leoni, and E. bipunctata are com-
parable in length to those of species of the Albida Group (Selander,
1981b). Epicauta niveolineata, in contrast, has an unusually lengthy incuba-
tion period, equaled elsewhere in Macrobasis only by that of E. fabricii
(LeConte) of the Fabricii Group (Selander, 1981b).

Hatching time in Table 1 refers to the number of days from oviposition
until hatching began in an egg mass. In E. uniforma, E. leoni, and E. bi-
punctata the eggs in a mass hatched more or less synchronously. In E.
niveolineata the hatching period extended over a period of several days.

Proper interpretation of the data on number of eggs per mass in meloids
requires that the size of the adult females producing the eggs be taken
into consideration (Adams and Selander, 1979). The eggs (and first-instar
larvae) of E. uniforma, E. leoni, and E. bipunctata are nearly equal in size.
Consequently, one might expect the ratio of number of eggs per mass to
adult body volume in these species to be constant or nearly so. In our
samples the adults of E. uniforma and E. leoni were similar in size and, as
shown in Table 1, their egg masses contained, on the average, roughly the
same number of eggs. Females of E. bipunctata in our samples were 3-4
times as large (in terms of volume) as those of E. uniforma and E. leoni

VOLUME 84, NUMBER 1 143

and, as expected, their egg masses contained more than three times the
mean number of eggs. The relationship between adult body size and number
of eggs per mass is quite different, however, for E. niveolineata. Adults of
E. niveolineata, although nearly as large as those of E. bipunctata, pro-
duced egg masses containing less than half the number of eggs. The differ-
ence reflects the fact that the volume of an egg in E. niveolineata is much
greater (about three times) than in E. bipunctata.

Larvae of several species of the Albida Group are known to prey on the
eggs of grasshoppers of the genus Melanoplus in nature (Horsfall, 1943;
Parker and Wakeland, 1957). Judging from the anatomical similarity of the
larvae of the Albida and Uniforma groups, one would infer that the latter
are grasshopper egg predators also and, in fact, both E. /eoni and E. niveo-
lineata have been reared in our laboratory on the eggs of Melanoplus dif-
ferentialis (Thomas).

KEY TO SPECIES OF THE UNIFORMA GROUP BASED ON THE
First LARVAL INSTAR

1. Head longer than wide, with sides, except at immediate base,
straight and subparallel (Fig. 1b); mandible with 5—6 teeth visible in
outline; long setae of thorax and abdomen branched several times
Neartipsate see Ao! BREE ee niveolineata (Haag-Rutenberg)

— Head wider than long, with sides rounded and weakly convergent

behind middle (Fig. la); mandible with 34 teeth visible in outline;

lone setae 'ofbody hot: branehed : 62/009 86 Se I 2
. Lanceolate setae on femora relatively slender, especially on femur

III (Fig. 2c); major seta on coxa III heavy; abdominal terga I-II and

VI-VIII much darker in color than HJ—V; cuticular reticulations

near lateral margins of terga I-VIII prolonged posteriad into con-

spicuous toothlike evaginations .................. bipunctata Werner

— Lanceolate setae on femora broad, heavy (Figs. 2a, b); major seta
on coxa III slender; abdominal terga usually uniform in color; cu-

ie)

ticular reticulations of terga not prolonged posteriad ............... 3
3. Mandibular teeth more or less V-shaped in outline; abdominal terga
Hehe Drown to rowan: POL ee See leoni Duges

— Mandibular teeth rounded; abdominal terga light yellow brown
uniforma Werner

Bea eerie a ee eee ete) a ow. ot ee ee wre 6 ate sere Ne C's! a ew VIET eTe Vere jerete "ere

DESCRIPTIONS OF THE SPECIES

Epicauta uniforma Werner
Figs. la, 2a

Description.—Color of head, pro- and metanotum, abdominal tergum I,
and, sometimes, terga VI-VIII light brown; rest light yellow brown.

144 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Head of first-instar larva, ventral view. a, Epicauta uniforma. b, E. niveolineata.

Head shorter than pro- and mesonotum combined, nearly 1!/10 as wide as
long; sides straight, parallel before middle, rounded and weakly convergent
behind (except at immediate base); gula about % as long as greatest width
of head; gular setae not reaching anterior margin of gula. Antenna with
segment II more than 142 as long as III; sensory organ as long as segment
III, extending beyond apex of segment. Mandibles less than 3/s as long as
head, % as wide as long, bearing 5—6 teeth (3-4 visible in outline); teeth
rounded. Maxillary palpus with segment III more than 12x as long as wide;
sensory area of segment III extending more than 2 length of dorsal surface;
papillae of sensory area moderately dense, partially obscuring one another
in dorsal view; length of 2-segmented sensory appendix more than 4% width
of segment II of labial palpus. Labial palpus with segment I about % as long
as II; Il more than 2 as long as wide. Abdomen with cuticular reticulations
prolonged posteriad into inconspicuous toothlike evaginations on median 4
of pleurites I-VIII in some specimens; 12 (rarely 10) setae in posterior
marginal row on terga I-VIII; setae not divided apically; spinelike evagin-
ations at bases of marginal setae moderately developed on terga I-IV, weak-
ly so on V or V and VI; tergum V about 4x as wide as long, with central
pair of posterior marginal setae less than 2/s as long as tergum; central pair
of setae of median transverse row of tergum V 3/10 as long as tergum; spi-
racle | slightly more than 4 as large as mesothoracic spiracle; caudal setae
at most as long as combined length of terga VII-IX. Femur I with 7 (rarely
6) lanceolate setae; all lanceolate setae heavy; tarsungulus I with longer seta
reaching distal 4. Length of body 2.2 mm. Length of caudal setae 0.3 mm.

VOLUME 84, NUMBER 1 145

Material studied.—25 larvae, adults from Ft. Davis, Jeff Davis Co., Tex-
as, 7/14-VII-63, R. B. Selander and J. M. Mathieu (Notes 24-63 RBS); 15
larvae, adults from Ft. Davis, Texas, 12-VII-63, R. B. Selander and J. M.
Mathieu (Notes 28-63 RBS); 10 larvae, adults from Ft. Davis, Texas, 10/12-
VIII-67, R. B. Selander (Notes 8-67 RBS).

Remarks.—This species, E. leoni, and E. bipunctata run to E. conferta
(Say) in couplet | of MacSwain’s (1956) key to the larvae of Epicauta. They
are immediately distinguished from that species by the shape of the head,
the presence of fewer mandibular teeth, and the fact that there are only
seven, rather than nine, lanceolate femoral setae.

Epicauta leoni Duges
Fig. 2b

Description.—Color of head and pronotum brown to dark brown; meso-
and metanotum and abdominal terga I-IX light brown to brown.

Head about as long as pro- and mesonotum combined, 1|'/10 as wide as
long; sides straight, parallel before middle, rounded and weakly convergent
behind middle (except at immediate base); gula about % as long as greatest
width of head; gular setae not reaching anterior margin of gula. Antenna
with segment II about 2 as long as III; sensory organ longer than segment
III, extending well beyond apex of segment. Mandibles about 3/s as long as
head, about /% as wide as long, bearing 5—6 teeth (3-4 visible in outline);
teeth more or less V-shaped in outline. Maxillary palpus with segment III
about 112 x as long as wide; sensory area of segment III extending more than 2
length of dorsal surface; papillae of sensory area moderately dense, partially
obscuring one another in dorsal view; length of 2-segmented sensory ap-
pendix about % width of segment II of labial palpus. Labial palpus with
segment I about % as long as II; IJ 2x as long as wide. Abdomen with
cuticular reticulations not prolonged into toothlike evaginations; 12 (rarely
14) setae in posterior marginal row on terga I-VIII; setae not divided api-
cally; spinelike evaginations at bases of marginal setae moderately devel-
oped on terga I-IV, weakly so on V; tergum V more than 4x as wide as
long, with central pair of posterior marginal setae about 2 as long as tergum;
central pair of setae of median transverse row of tergum V 3/10 as long as
tergum; spiracle I less than 74 as large as mesothoracic spiracle; caudal
setae longer than combined length of terga VII-IX. Femur I with 7 (rarely
6 or 8) lanceolate setae; all lanceolate setae heavy; tarsungulus I with longer
seta reaching distal '/6. Length of body 2.0 mm. Length of caudal setae 0.4
mm.

Material studied.—32 larvae, adults from Chapingo, Mexico, Mexico, 10-
VIII-63, R. B. Selander and R. H. Storch (Notes 3-63 RBS).

Remarks.—See remarks for E. uniforma.

146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Leg III of first-instar larva, anterior view (tibia and tarsungulus omitted except in
Fig. 2a). a, Epicauta uniforma. b, E. leoni. c, E. bipunctata. d, E. niveolineata.

Epicauta bipunctata Werner
Fig. 2c

Description.—Color of head, pro- and metanotum, and abdominal terga
I, Il, and VI-VIII brown; rest light yellow brown.

Head shorter than pro- and mesonotum combined, nearly 1'/10 as wide as
long; sides straight, parallel before middle, rounded and weakly convergent
behind (except at immediate base); gula % as long as greatest width of head;
gular setae nearly reaching anterior margin of gula. Antenna with segment
II more than 12x as long as III; sensory organ longer than segment III,
extending beyond apex of segment. Mandibles nearly 3/s as long as head,
about */s as wide as long, bearing 5—6 teeth (3-4 visible in outline); teeth
rounded. Maxillary palpus with segment III more than 112 as long as wide;
sensory area of segment III extending % length of dorsal surface; papillae
of sensory area moderately dense, partially obscuring one another in dorsal
view; length of 2-segmented sensory appendix more than 4 width of seg-
ment II of labial palpus. Labial palpus with segment I about 4 as long as
II; If 2x as long as wide. Abdomen with cuticular evaginations near lateral
margins of terga and on median % of pleurites of segments I-VIII prolonged
posteriad into conspicuous toothlike evaginations; 12 setae in posterior mar-

VOLUME 84, NUMBER 1 147

ginal row on terga I—-VIII; setae not divided apically; spinelike evaginations
at bases of marginal setae moderately developed on terga I-IV, very weakly
so on V; tergum V about 4x as wide as long, with central pair of posterior
marginal setae 7/; as long as tergum; central pair of setae of median transverse
row of tergum V almost ?/s as long as tergum; spiracle I 7/10 as large as
mesothoracic spiracle; caudal setae °/10 as long as combined length of terga
VII-IX. Major seta on coxa III unusually heavy; femur I with 7 lanceolate
setae; lanceolate setae relatively slender (especially noticeable on leg III):
tarsungulus I with longer seta reaching distal 4. Length of body 2.1 mm.
Length of caudal setae 0.3 mm.

Material studied.—27 larvae, adults from Culiacan, Sinaloa, Mexico, 28-
IX-72, R. B. Selander and A. Berrios-Ortiz (Notes 599-72 RBS).

Remarks.—See remarks for E. uniforma.

Epicauta niveolineata (Haag-Rutenberg)
Figs. 1b, 2d

Description.—Color of head amber; pro- and metanotum and abdominal
terga I, Il, and VI-IX dark brown; mesonotum and abdominal terga HI-—V
light yellow brown.

Head about as long as pro- and mesonotum combined, about 4/s as wide
as long; sides straight, slightly divergent posteriad to near base; gula about
3/5 as long as greatest width of head; gular setae extending well beyond
anterior margin of gula. Antenna with segment II at least 1*4x as long as
III; sensory organ reaching apex of segment III. Mandibles robust, 3/s as
long as head, about % as wide as long, bearing 7—9 large teeth (4-6 visible
in outline); teeth rounded. Maxillary palpus with segment III about 134 as
long as wide; sensory area of segment III extending at most % length of
dorsal surface, more clearly overlapping onto ventral surface than in other
species; papillae of sensory area sparse, not obscuring one another in dorsal
view; length of 2-segmented sensory appendix more than *% width of seg-
ment II of labial palpus. Labial palpus with segment I more than % as long
as II; II more than 2 as long as wide. Abdomen with cuticular reticulations
not prolonged into toothlike evaginations; 10 setae in posterior marginal row
on terga I-VIII; setae divided several times apically; spinelike evaginations
at bases of marginal setae strongly developed on terga I-VI, weakly so on
VII; tergum V more than 32x as wide as long, with central pair of posterior
marginal setae about % as long as tergum; central pair of setae of median
transverse row of tergum V 2/s as long as tergum; spiracle I about 74 as
large as mesothoracic spiracle; caudal setae longer than combined length of
terga VII-IX. Femur I with 7 lanceolate setae; all lanceolate setae very
heavy, broad; tarsungulus I with longer seta reaching distal '/s. Length of
body 3.3 mm. Length of caudal setae 0.8 mm.

Material studied.—10 larvae, adults from between Tequesistlan and To-
tolapan, Oaxaca, Mexico, 17-VIII-74, M. Morgan (Notes I-74 RBS).

148 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Remarks.—This species runs as far as couplet 5 in the key to the larvae
of Epicauta developed by MacSwain (1956). It can be distinguished from
the two alternatives in that couplet by the following combination of char-
acteristics: setae of abdominal tergum V about *% as long as tergum, prox-
imal mandibular seta much shorter than distal one, femur I with seven (or
fewer) lanceolate setae.

ACKNOWLEDGMENTS

We are indebted to F. G. Werner for a gift of larvae of nominal Epicauta
uniforma from Arizona; to A. Berrios-Ortiz, J. M. Mathieu, and R. H.
Storch for assistance and companionship in the field; to M. Morgan for
collecting adults of E. niveolineata; and to A. Prickett for inking the draw-
ings for this article.

Field work in Texas and Mexico was supported by a grant from the Na-
tional Science Foundation (GB-35526, R. B. Selander, Principal Investi-
gator).

LITERATURE CITED

Adams, C. L. and R. B. Selander. 1979. The biology of blister beetles of the Vittata Group
of the genus Epicauta (Coleoptera, Meloidae). Bull. Am. Mus. Nat. Hist. 162(4): 137-
266.

Agafitei, N. J. and R. B. Selander. 1980. First instar larvae of the Vittata Group of the genus
Epicauta (Coleoptera: Meloidae). J. Kans. Entomol. Soc. 53: 1-26.

Cros, A. 1938. Considérations générales sur le genre Epicauta Redtenbacher. Etude biolo-
gique sur Epicauta albovittata Gestro. Mem. Soc. Entomol. Ital. 16: 129-144.

Horsfall, W. R. 1943. Biology and control of common blister beetles in Arkansas. Arkansas
Agric. Exp. Stn. Bull. 456, 55 pp.

MacSwain, J. M. 1956. A classification of the first instar larvae of the Meloidae (Coleoptera).
Univ. Calif. Publ. Entomol. 12, 182 pp.

Parker, J. R. and C. Wakeland. 1957. Grasshopper egg pods destroyed by larvae of bee flies,
blister beetles, and ground beetles. U.S. Dep. Agric. Tech. Bull. 1165, 29 pp.

Pinto, J. D. 1975. The taxonomy of three species of maculate Epicauta (Coleoptera: Meloi-
dae), with descriptions of courtship behavior and first instar larvae. J. Kans. Entomol.
Soc. 48: 429-452.

Selander, R. B. 198la. The Caustica Group of the genus Epicauta (Coleoptera, Meloidae).
Proc. Entomol. Soc. Wash. 83: 573-591.

——. 1981b. Evidence for a third type of larval prey in blister beetles (Coleoptera:
Meloidae). J. Kans. Entomol. Soc. 54: 757-783.

Selander, R. B. and N. J. Agafitei. In press. The first instar larva of Epicauta funesta (Chev-
rolat) (Coleoptera, Meloidae). Coleopt. Bull.

Selander, R. B. and J. M. Mathieu. 1969. Ecology, behavior, and adult anatomy of the Albida
Group of the genus Epicauta (Coleoptera, Meloidae). Illinois Biol. Monogr. 42, 168 pp.

Werner, F. G. 1954. A review of the subgenus Gnathospasta of the genus Epicauta (Meloi-
dae). Coleopt. Bull. 8: 25-27.

——.. 1958. Studies in the genus Epicauta of the North American continent (Meloidae).
Il. The Uniforma Group. Coleopt. Bull. 12: 1-20.

——. 1962. The first instar larva of Epicauta, subgenus Gnathospasta (Coleoptera: Meloi-
dae). Proc. 11th Int. Congr. Entomol. 1: 106-109.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 149-156

NESTING BEHAVIOR OF LYRODA SUBITA (SAY)
(HYMENOPTERA: SPHECIDAE)

FRANK E. KURCZEWSKI AND DAVID J. PECKHAM

(FEK) Department of Environmental and Forest Biology, S.U.N.Y. Col-
lege of Environmental Science and Forestry, Syracuse, New York 13210;
(DJP) Upstate Medical Center, Syracuse, New York 13210.

Abstract.—Observations made on 17 females of Lyroda subita (Say) nest-
ing in upstate New York extend the range of variation in the behavioral
components and nesting parameters of this species. Characteristic features
include nesting in a variety of friable soils, beginning nests from pre-existing
depressions, leaving entrances open during provisioning activities, practic-
ing mandibular prey transport, usually in low flights, preying upon mostly
immature gryllids which are incompletely paralyzed and stored in multicel-
lular nests, and attaching the egg to the venter of the prey’s thorax between
the fore- and midlegs. Several of the cells were parasitized by satellite-flies
of the genus Senotainia (Sarcophagidae: Miltogramminae). The components
of L. subita behavior are discussed and compared with those of other (ex-
otic) species in the genus.

The genus Lyroda contains only two species, subita (Say) and triloba
(Say), in North America north of Mexico (Krombein, 1979). Lyroda subita
occurs transcontinentally in southern Canada and the U.S., whereas L.
triloba has been reported from Canada, D.C., Illinois, Indiana, Kansas,
Louisiana and Texas (Krombein, 1979). Using larval characters, Evans
(1964) placed the genus in the tribe Miscophini. A study of adult characters
supports this placement (Bohart and Menke, 1976).

Nothing is known about the biology of L. triloba. Patton (1892) reported
on the provisions and prey transport of L. subita. The Peckhams (1898,
1905) observed a female of L. subita nesting in Wisconsin and described
prey transport, nest entry, cell depth, prey paralysis, and larval develop-
ment. Williams (1914) mentioned the searching activity of L. subita females.
Evans (1964) reported on nest-site selection, cell depth, prey transport, nest
entry, provisions, number of prey per cell, egg placement, and parasitism.
Our observations add to the knowledge of the nesting behavior of this
species, particularly nest structure, dimensions, and cell contents.

150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ETHOLOGY

Nest-site selection.—Females of L. subita were observed nesting at three
localities in Cayuga County, N.Y. during August and September 1969-71.
Selection of a nesting site involved much searching activity on the surface
and exploration below ground level. Active nests were located in the center
and at the edge of a sand pit, border of a field, and slope of an erosion ditch.
Wasps utilized for nesting sites deep cracks and crevices in the sand as well
as both emergence and provisioning burrows of the cicada-killer Sphecius
speciosus (Say), provisioning burrows of Tachytes validus Cresson (Spe-
cidae), and resting burrows of larval tiger beetles, Cicindela spp. In each
case the insect had abandoned the site before Lyroda began using it.

Hunting behavior—Females searched for prey on and in the ground,
exploring holes, cracks, and crevices and foraging near the bases of vege-
tation. The distance of this activity from the nest varied with the success
of the excursion; the wasp usually began nearby and gradually moved far-
ther from the entrance. Prey capture was observed twice. In both cases the
wasp pounced upon a cricket and inserted her sting in the throat for several
seconds. She then reinserted the sting in the throat and, later, after repo-
sitioning herself, stung it near the base of a hindleg. The wasp then malax-
ated the cricket and/or rested on the sand and cleaned.

Prey transport, nest entry and exit.—Following prey paralysis, the female
positioned the cricket ventral side upward or on its side (once dorsal side
upward) and head forward, and grasped its antennae with her mandibles.
She then proceeded forward to the nest either on the ground or in flight.
The usual manner of transport involved a series of short, low flights, during
which the legs assisted in holding the prey. Some provisioning females re-
turned directly to their nests, almost in a straight line, and plunged inside
without hesitating. These wasps left head first several s or min later and
resumed hunting activities. One wasp spent one and one-half h provisioning
a cell with five crickets, spending 6 to 55 ( = 18) min between successful
hunting excursions.

Nest structure and dimensions.—The entrance to a nest of L. subita was
either that of the original inhabitant, sometimes slightly modified, or simply
an opening in the ground. The tumulus often had been eliminated entirely
by weathering. The diameter of the entrance and proximal burrow usually
divulged the first inhabitant. For example, burrows of renovated T. validus
nests leading to burrows and cells of L. subita were 6-7 mm wide, those of
the cicada-killer, 20-24 mm wide, except for one which narrowed to 6 mm
at the entrance, and those of larval tiger-beetles, ca. 6 mm at the entrance,
narrowing to 4 mm farther down. One nest had three species, S. speciosus,

T. validus and L. subita nesting in succession, all utilizing a common en-
trance and proximal burrow.

VOLUME 84, NUMBER | 151

20cm

Fig. 1. Six nests of Lyroda subita, as viewed from the side, showing structure of burrow
and placement of cells. Abbreviations: S = Sphecius speciosus burrow; T = Tachytes validus
cell. L. subita cellular contents: e = egg; 1 = larva; m = maggot(s); a = ants. Stippling indi-
cates burrow filled with sand. Scale at lower left refers to all six nests.

The L. subita portion of the nest exhibited a basic pattern, which included
a main burrow and branching side-burrows leading to rearing cells (Fig. 1).
The number of completed cells per nest ranged from two to five, the wasp
finishing ca. one per day. Cells were located at distances of from 12.5 to

152 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

29.5 cm (* = 19.5; n = 11) from the surface entrances. However, the bi-
furcating L. subita burrows themselves were only 3 to 19 cm (* = Il; n =
6) long. Upon completion of the nest the wasp filled that portion of the
burrow directly in front of the cell and all or much of the remaining portion
of the burrow with sand, but three finished nests remained partly open near
the surface.

Cells of L. subita were never built in series but always singly at the ends
of short side-burrows. This arrangement, as viewed from above, appeared
palmate or fan-shaped. The distance between adjacent cells ranged from 2.5
to 7.0 cm in three nests. Cell depth from the sand surface varied with the
dimensions of the pre-existing burrow, and ranged from 6.5 to 26.5 cm (% =
14.0; n = 20). Cells in a nest were more nearly equivalent in depth than
cells from different nests; e.g., four cells in one nest begun from a shallow
subterranean crevice were only 6.5—12.5 cm deep, whereas three cells in a
nest modified from a tiger beetle burrow were 17.5—26.5 cm deep.

Cells (19) of L. subita were 5 to 8 mm (x = 6.4) in diameter, and 10 to
16 mm (x = 13.2) long. There appeared to be a correlation between cell size
and number of prey per cell in 12 of 17 fully-provisioned cells, i.e., seven
‘large’? cells, 13-16 mm long, contained 3-7 prey per cell, and five ‘‘small”’
ones, 10-12 mm long, only |-2 prey per cell. Some larger cells were nearly
perfectly ovoidal-elliptical, whereas a few smaller ones were rather asym-
metrical. Cells were slanted either upward or downward in the sand, or
rarely horizontally, to coincide with the course of the burrow at the end.
With one exception, all or most cells in a nest sloped in one direction.

Provisions.—L. subita preyed on small, mostly nymphal ‘‘field-crickets”’

(Gryllidae). Prey collected from cells consisted of four adults and four
nymphs of Allonemobius carolinus Scudder, 7 adults and 41 nymphs of A.
fasciatus (DeGeer), and 11 nymphs of Gryllus pennsylvanicus Burmeister.
Two nests at one locality contained only A. fasciatus. A third nest at the
same locality held this species and G. pennsylvanicus. A nest in another
area contained all A. carolinus.

Prey recovered from recently provisioned cells were incompletely para-
lyzed and moved only the appendages and abdominal segments. Up to one-
half of the crickets in a nest had a single hindleg missing; in two nests, the
missing leg on four crickets was the left, and in one nest with two prey, the
right.

The number of prey stored in fully provisioned cells ranged from 1-7 (% =
3; n = 17). The weights of the individual crickets varied from 11-73 mg
(t = 35; n = 48). Female wasps weighed, on the average, 21 mg (n = 4).
The weight of all prey in a completed cell averaged 107 mg (39-173; n =

17). Cells with a greater biomass usually contained more crickets rather
than fewer, larger prey.

VOLUME 84, NUMBER 1 153

Forty-four of 49 crickets were placed in cells head inward and ventral
side upward. Three individuals were positioned head inward and on the side
with their venters facing the cell walls, and two were put head inward and
dorsal side upward. In cells containing several prey, crickets were often
piled atop one another; however in some cells with only two prey, the
crickets were laid in tandem. In one unfinished nest a paralyzed cricket was
located outside of the cell in an open burrow.

Egg.—The wasp did not lay her egg on a cricket until the full complement
of prey had been put in the cell. Three cells were unearthed, evidently in
early stages of provisioning, with only one or a few crickets and no egg or
larva. The cricket on which the egg had been laid was invariably placed
head inward and ventral side upward (n = 12). In five of six cases, this prey
was positioned innermost in the cell. Once, the egg-bearing prey was placed
atop others in the cell. The cricket containing the wasp’s egg was the largest
prey in the cell in 4 of 12 examples. The egg-bearing individual averaged 36
mg (18-73; n = 13) in weight.

The egg of L. subita was 2.0—2.5 mm long, 0.6—0.7 mm in diameter, white,
elastic, elongate and slightly curved. It was attached by the cephalic end to
the forecoxal corium of the prey, the caudal end extending laterally to the
other side between the first and second pairs of legs (Fig. 2). In four nests,
eggs or small larvae were attached to either the left or right sides of the
crickets in equal numbers.

Mortality.—Of the 21 completed cells, 10 contained up to three maggots
feeding on the innermost cricket. At least one cell in every multicellular
nest was afflicted and in one nest, all three cells were destroyed. Two mag-
gots were reared and identified as Senotainia trilineata (Wulp) (Sarcophag-
idae). In several instances, female S. trilineata were observed actively pur-
suing provisioning wasps. Two additional cells were destroyed by ants,
probably Solenopis molesta (Say), and one nest contained cells with moldy

prey.
DISCUSSION

Evans (1964, 1973) found L. subita renovating pre-existing burrows and
cavities in friable soils in Massachusetts and New York. The utilization of
such depressions was reflected in the diverse architecture and dimensions
of the several nests we excavated. Lyroda formosa (F. Smith) also nests in
the ground in similar situations, utilizing deep burrows of large sand wasps
and crevices for nesting sites (Iwata, 1938, 1964).

The Nearctic L. subita stores its nests with Gryllidae (see Patton, 1892;
Peckham and Peckham, 1898; Evans, 1974), whereas some Asiatic and Af-
rican species (japonica Iwata, formosa, madecassa Arnold) prey upon Te-
trigidae (Iwata, 1938, 1963, 1964; Tsuneki and lida, 1969). The fact that

154 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Egg of Lyroda subita attached to gryllid prey at right forecoxal corium and extending
laterally between fore- and midlegs. Note missing left hindleg.

some of the L. subita nests we excavated contained only Allonemobius
fasciatus or A. carolinus suggests either conditioning on the part of the
female wasp during hunting or that individual wasps hunted in areas where
each of these species was prevalent. Similar conclusions have been drawn
regarding the hunting behavior of Chlorion aerarium Patton (Peckham and

VOLUME 84, NUMBER 1 155

Kurcezewski, 1978) and Liris argentata (Palisot de Beauvois) (O’Brien and
Kurczewski, unpublished data).

The transport of prey to the nest in L. subita is typically ‘‘larrine’’, i.e.,
on the ground or in flight depending on the size and weight of the prey, and
grasping the prey’s antennae with the mandibles. Similar details of prey
transport for members of this genus have been recorded by Patton (1892),
Iwata (1963, 1964), and Evans (1964). Tsuneki and lida (1969), on the other
hand, reported that females of L. japonica carried their prey with the middle
legs.

The incomplete paralysis of the prey of L. subita coincides with that of
most larrine wasps that utilize orthopterous prey (see especially Krombein
and Kurczewski, 1963; Iwata, 1942, 1976; Evans, 1966, for summaries).
Tsuneki and lida (1969) reported only partial paralysis with eventual com-
plete recovery of the prey in L. japonica.

Gryllid prey missing one or two hindlegs are common among the Spheci-
dae, e.g., Chlorion aerarium (Peckham and Kurczewski, 1978), Liris
nigra V.d.L. (Steiner, 1968), so that finding a number of prey of L. subita
missing one or more legs was not unusual. This appendage loss may reflect
the manipulation of the prey during the stinging or attempted stinging pro-
cess (see Steiner, 1968, Plate 22).

We found that the egg of L. subita may be laid on the first, last or inter-
mediate prey taken into the nest. Tsuneki and lida (1969) noted that the egg
of L. japonica was laid invariably on the last provision taken into the nest.
In L. subita (Evans, 1964), L. formosa (Iwata, 1964) and L. japonica (Tsu-
neki and lida, 1969) the egg is laid in a rather typically larrine position
between the first and second pairs of legs and is affixed to the forecoxal
corium of one of the prey. In these species the egg is not laid until the full
complement of prey had been placed in the cell despite Peckham and Peck-
ham’s (1898) account of progressive provisioning in L. subita.

The rather high incidence of miltogrammine parasitism we found in cells
of L. subita substantiated work by Evans (1964), who noted many L. subita
nests parasitized by miltogrammine flies, in two cases, Metopia argyro-
cephala Meigen.

ACKNOWLEDGMENTS

We thank A. B. Gurney, Systematic Entomology Laboratory, USDA, for
determining the prey gryllids and R. J. Gagné, Systematic Entomology Lab-
oratory, USDA, for naming the parasitic miltogrammine flies.

LITERATURE CITED

Bohart, R. M. and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. Univ.
Calif. Press, Berkeley. ix + 695 pp.

156 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Evans. H. E. 1962. The evolution of prey-carrying mechanisms in wasps. Evolution 16: 468-
483.

—__. 1964. The classification and evolution of digger wasps as suggested by larval char-
acters. Entomol. News 75: 225-237.

———. 1966. The behavior patterns of solitary wasps. Annu. Rev. Entomol. 11: 123-154.

——. 1973. Digger wasps as colonizers of new habitat (Hymenoptera: Aculeata). J. N.Y.
Entomol. Soc. 82: 259-267.

Iwata, K. 1938. On the habits of some Larridae in Japan. Kontyd 12: 1-13.

. 1942. Comparative studies on the habits of solitary wasps. Tenthredo 6: 1-146.

———. 1963. Miscellaneous biological notes on aculeate Hymenoptera in Kagawa in the years
of 1948 and 1949. Trans. Shikoku Entomol. Soc. 7: 114-118.

—. 1964. Bionomics of non-social wasps in Thailand. Nat. Life Southeast Asia 3: 323-
383.

—. 1976. Evolution of Instinct. Comparative Ethology of Hymenoptera. Amerind Publ.
Co., New Delhi. xi + 535 pp.

Krombein, K. V. 1979. Larridae, pp. 1617-1650. In Krombein, K. V. et al., eds., Catalog of
Hymenoptera in America North of Mexico, Vol. 2, Smithsonian Institution Press, Wash-
ington, D.C.

Krombein, K. V. and F. E. Kurczewski. 1963. Biological notes on three Floridian wasps.
Proc. Biol. Soc. Wash. 76: 139-152.

Patton, W. H. 1892. Notes upon Larradae. Entomol. News 3: 89-90.

Peckham, D. J. and F. E. Kurczewski. 1978. Nesting behavior of Chlorion aerarium. Ann.
Entomol. Soc. Am. 71: 758-761.

Peckham, G. W. and E. G. Peckham. 1898. On the instincts and habits of the solitary wasps.

Wisconsin Geol. Nat. Hist. Surv., Sci. Ser., Bull. 2: 1-245.

. 1905. Wasps social and solitary. Houghton, Mifflin and Co., Boston. xv + 311 pp.

Steiner, A. L. 1968. Behavioral interactions between Liris nigra Van der Linden and Gryllus
domesticus L. Psyche (Camb. Mass.) 75: 256-273.

Tsuneki, K. and T. lida. 1969. The biology of some species of the Formosan Sphecidae, with
descriptions of their larvae. Etizenia 37: 1-21.

Williams, F. X. 1914. The Larridae of Kansas. Univ. Kans. Sci. Bull. 18: 121-213.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 157-164

TWO NEW SPECIES OF WATER STRIDERS OF THE GENUS
TREPOBATES UHLER (HEMIPTERA: GERRIDAE)

PAUL D. KITTLE

Department of Biology, Southeast Missouri State University, Cape Gi-
rardeau, Missouri 63701.

Abstract.—Trepobates carri, new species, is described from Guatemala
and is also known from Belize, Cuba, Honduras, Jamaica, Mexico, and
Texas. Trepobates polhemi, new species, is described from Mexico.

In the course of my studies on Trepobates, I have discovered two new
taxa which are described below.

Trepobates carri Kittle, New Species
Figs. 1-7

Trepobates pictus (Herrich-Schaeffer) [misidentification]; Drake and Harris,
1932- We

Diagnosis.—Trepobates carri can be separated from all other species of
Trepobates except T. pictus by the presence of a posterior mesonotal pro-
jection in the female. Although very similar to 7. pictus, T. carri differs
from it in having (1) a more tumid, shorter, and hairier posterior mesonotal
projection in the female; (2) a shorter first antennal segment, especially in
the male; (3) a slightly different color pattern; and (4) a different shaped
paramere (compare Figs. 7 and 8).

Description.—Length: Male 3.09-3.66 mm; female 3.27-4.11 mm. Width:
Male 1.22—1.45 mm; female 1.53—1.94 mm.

Head: Marked with black and light yellow (Figs. 2-4). Antennal segment
1 slightly bowed at base, 0.76-0.99 mm long in male, 0.63—0.84 mm long in
female. Antennal segments 2-4 straight, 3 without long hairs.

Thorax: Pronotum marked with black and light yellow (Figs. 2-4). Prono-
tum in alate form prolonged posteriorly, with light yellow lateral, longitu-
dinal stripes usually continuous with light band along posterior margin,
sometimes with an anterior median yellow stripe. Anterior femur moder-
ately bowed just beyond middle in male, slightly bowed near base in female,
not constricted at apex. Anterior femur 1.19—1.46 mm long in male, 0.99-
1.27 mm long in female. Anterior tibia slightly bowed in male, straight in

158 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ITY

a sais Ne
Se

Fig. 1. Distribution of Trepobates carri (circles) and T. polhemi (triangles).

female. Mesonotum with black and light yellow markings (Figs. 2-4); pro-
longed posteriorly in female, this projection with long hairs and variable in
shape (Figs. 5 and 6); a patch of very short, dark hairs usually present along
median portion of mesonotal-mesopleural line in female. A light yellow me-
sopleural stripe present posterior to black postocular stripe of pronotum.
Mesosternum with a pair of small, oval, widely separated brown spots often
present posteriorly, especially in female. Middle femur of male with a thick
ventral fringe of short, dark brown hairs; these hairs, at maximum length,
approximately 20-40% of diameter of femur. Middle femur 1.45—1.88 mm
long in male, 1.59-1.95 mm long in female. Middle tibia of male with a single
row of short hairs along basal % which gradually decrease in length poste-
riorly; these hairs, at maximum length, approximately 40% of diameter of
tibia. Middle tibia 2.71-3.76 mm long in male, 2.85—3.59 mm long in female.
Metanotum black with paired, light yellow markings, without a median
patch of hairs. Posterior femur without long hairs at base. Posterior femur
1.92—2.60 mm long in male, 1.97-2.48 mm long in female.

Abdomen: Abdominal terga mostly black, some marked with light yellow,
especially in female, almost entirely black in dark individuals; some gray-
blue bloom present, especially in male. Ventrites mostly pale yellow, dark

VOLUME 84, NUMBER I 159

brown along anterior margins and laterally in some individuals; ventrite 7
in female with a row of moderately long, dark brown hairs along posterior
margin, this row interrupted at middle. Connexival segments mostly light
yellow with black anterior margins; lateral margins of segments 5 and 6 in
male and 5-7 in female with moderately long, dark hairs. Connexiva with
moderately long, dark hairs at apex and very slightly produced posteriorly
but without long spines in female. Abdominal segment 8 of male with scat-
tered, short, brown hairs ventrally. Pygophore with several long, dark
brown hairs at middle; short, pale pubescence posteriorly. Paramere sharply
pointed (Fig. 7) and shaped different from that of 7. pictus (Fig. 8).

Holotype.—Guatemala: Apterous 2, pond, 3 mi. S Tikal, Peten, CL 653,
2 Jan 1973, J. T. Polhemus, in J. T. Polhemus Collection, Englewood, Col-
orado.

Paratypes.—BELIZE: Beaver Dam Cr., 15 mi. E Roaring Creek, 27 Dec
1973 (1 6, 2 2; JTP). CUBA: Santiago de Las Vegas, 13 Nov 1922 (1 9;
UK); Consolacion del Sur, 16 Apr 1930 (1 ¢6, 1 2; NN). GUATEMALA:
pond, 3 mi. S Tikal, Peten, 2 Jan 1973 (17 6, 30 2; JTP, PDK). HONDU-
RAS: Tela, 15 Mar 1936 (16 3, 31 2; UK). JAMAICA: Port Antonio (1
2; AMNH); pond, St. Ann, Feb. 1928 (1 9; UK). MEXICO: Campeche:
Champoton Janataya, 9 Jul 1932 (1 2; USNM). Nuevo Leon: Rio Cabezano,
17 mi. N Linares, 14 Dec 1969 (1 6; JTP). San Luis Potosi: 3 mi. W El
Naranjo, 5 Jun 1965 (1 6, 1 2; JCS). Tamaulipas: stream, El Salto, 14 Dec
1969 (2 2; JTP). Veracruz: 3 mi. W Paso de Ovejas, 17 Aug. 1959 (1 9;
USNM); stream, 3 mi. N Conejos, 6 Jan 1971 (4 3; JTP); borrow pit, 14
mi. N Nautla, 7 Jan 1971 (1 2; JTP); stream, 17 mi. S Tuxpan, 7 Jan 1971
(2 6; JTP); stream, 19 mi. N Conejos, 6 Jan 1971 (2 6; JIP); stream, S
Gutierrez Zamora, 7 Jan 1971 (1 6, 2 2; JTP); 15.8 mi. S Tampico, 19 Apr
1974 (4 6, 10 2; TAMU); 16 mi. S La Tinaja, 4 Jan 1971 (3 2; JTP); 19 mi.
SE Tantoyuca, 8 Jan 1971 (7 6, 8 2; JTP, PDK). Yucatan: Carlos Morales
Escuela Cenote, Merida, 23 Jul 1932 (3 ¢, 2 2; UK, USNM); Uki Cenote,
Motul, 26 Jul 1932 (6 2; UM, USNM); Manzanilla Cenote, Merida, 23 Jul
1932 (6 2; UM, USNM); Chapultapec Cenote, Merida, 20 Jul 1932 (6 2;
UK, UM, USNM); Geiser Cenote, Merida, 19 Jul 1932 (3 6, 3 2; UM,
USNM); Huntun Cenote, Piste, 19 Jun 1932 (1 2; UM), 20 Jun 1932 (1 6,
4 2; UK, UM, USNM); Choch Cenote, Piste, 21 Jun 1932 (2 6,5 2; UK,
UM, USNM); Ciruak Cenote, Piste, 22 Jun 1932 (8 2; UK, UM, USNM);
Ixil Cenote, Chichen Itza, 15 Jun 1932(1 d, 1 2; UK, UM); Xanaba Cenote,
Chichen Itza, 25 Jun 1932 (2 2; UM); Xcan Yui Cenote, Chichen Itza, 16
Jun 1932 (1 2; UM), 17 Jun 1932 (1 6, 9 29; UK, UM, USNM). UNITED
STATES: Texas: Cameron Co.: 28 Dec 1967 (1 36, 1 2; NTSU). La Salle
Co.: Nueces River, | mi. S Cotulla, 13 Jun 1976 (1 6, 1 2; PDK). Starr
Co., Arroyo Salado, 14 mi. E Rio Grande City, 11 Jun 1975 (2 6, 1 2;
CLS). Zapata Co.: pond, Zapata, 13 Jun 1976 (2 2; PDK).

160 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

0.1mm

7 8

Figs. 2-7. Trepobates carri. 2-4, Variation of color pattern in female. 5-6, Variation of
posterior mesonotal projection. 7, Paramere. Fig. 8, T. pictus, paramere. (Scale lines = 0.5
mm except as noted.)

VOLUME 84, NUMBER 1 161

Distribution.—Trepobates carri has been collected in southern Texas,
Mexico, Belize, Guatemala, Cuba, Honduras, and Jamaica (Fig. 1).

Variation.—The color pattern is highly variable (Figs. 2-4), and individ-
uals examined from Yucatan, Mexico, are almost entirely dark dorsally.
The posterior mesonotal projection in the female is somewhat variable in
length, tumidity, and hairiness (Figs. 5, 6), and Yucatan specimens have the
most tumid and hairy projections. Hairs along the median portion of the
mesonotal-mesopleural line usually are present in the female, but are absent
in some individuals. Brown mesosternal spots were often present, especially
in the female and darker specimens. Thirty-seven percent of the specimens
examined were alate.

Biology.—Trepobates carri has been taken from both lentic and lotic hab-
itats, including creeks, rivers, ponds, borrow pits, and sinkholes. I collected
T. carri in association with 7. subnitidus Esaki and Rheumatobates hun-
gerfordi Wiley in Texas. Collection records are known from November
through April and June through August.

Etymology.—This species is named in honor of the late Professor Lloyd
G. K. Carr, Rio Grande College, Rio Grande, Ohio, who first stimulated my
interest in entomology.

Remarks.—I have examined the specimen upon which Drake and Harris
(1932) based their record of 7. pictus from Jamaica and determined that it
is T. carri. Hungerford (1936) referred to this taxon as 7. sp.

Trepobates polhemi Kittle, New Species
Figs. 1, 9-14

Diagnosis.—Trepobates polhemi can be easily separated from its conge-
ners by the patch of long hairs at the posterior end of the mesonotal-me-
sopleural line, the long hairs along the basal inner margin of the posterior
femur, and the short hairs on the seventh ventrite of the female, and the
relatively large size and long appendages in both sexes.

Description.—Length: Male 3.61-4.11 mm; female 4.16-4.80 mm. Width:
Male 1.36—1.52 mm; female 1.62—1.98 mm.

Head: Marked with black and light yellow (Figs. 9-11). Antennal segment
1 slightly bowed at base, 0.96-1.09 mm long in male, 0.96-1.12 mm long in
female. Antennal segments 2-4 straight, 3 without long hairs.

Thorax: Pronotum marked with black and light yellow (Figs. 9-11). Prono-
tum in alate form prolonged posteriorly, black with a light yellow band along
posterior margin, with light yellow lateral stripes usually continuous with
band along posterior margin, and sometimes with an anterior median yellow
stripe. Anterior femur strongly bowed just beyond middle in male, only
slightly bowed at base in female; apex not constricted but slightly swollen
in male. Anterior femur 1.32—1.55 mm long in male, 1.40—1.72 mm long in

162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VOLUME 84, NUMBER 1 163

female. Anterior tibia very slightly bowed in male, straight in female. Meso-
notum with black and light yellow markings (Figs. 9-11) and not prolonged
posteriorly; a patch of long, dark hairs at posterior end of mesonotal-me-
sopleural line in female (Fig. 12). A light yellow mesopleural spot absent
posterior to black postocular stripe of pronotum. Middle femur of male with
a thick ventral fringe of long, dark brown hairs, these hairs, at maximum
length, approximately 55—75% of diameter of femur. Middle femur 2.06-
2.41 mm long in male, 2.32—2.83 mm long in female. Middle tibia of male
with a single row of hairs along basal ’% which gradually decrease in length
distally, these hairs, at maximum length, approximately 85-125% of diam-
eter of tibia. Middle tibia 3.89-4.58 mm long in male, 4.46-4.98 mm long in
female. Metanotum black, with paired, light yellow spots; rarely with a
median patch of long hairs in female. Posterior femur with a dense fringe
of very long hairs along basal inner margin in female (Fig. 13). Posterior
femur 2.93—3.37 mm long in male, 3.29-3.91 mm long in female.

Abdomen: Abdominal terga mostly black, some marked with light yellow,
especially in female; mostly covered with a gray-blue bloom, more extensive
in male. Ventrites mostly pale yellow with some black along sides, espe-
cially in male; ventrite 7 in female with a row of short, mostly light hairs
along posterior margin, these hairs not readily visible. Connexival segments
light yellow with black margins; lateral margins of segment 5 in male with
moderately long, dark hairs, segment 5 and sometimes 4 with long, dark
hairs and segments 2 and 3 sometimes with moderately long hairs in female;
connexiva not produced at apex into long spines. Abdominal segment 8 of
male with very short, thin, pale pubescence ventrally. Pygophore with very
short, pale pubescence posteriorly. Paramere as in Fig. 14.

Holotype.—Mexico: Apterous ¢, stream, 2 mi. S Cuautla, Morelos, CL
1036, 25 Apr 1964, J. T. and M. S. Polhemus, in J. T. Polhemus Collection,
Englewood, Colorado.

Paratypes.—MEXICO: Distrito Federal: Mexico City, 7 Jul 1937 (4 6;
UK). Guerrero: stream, 40 mi. N Acapulco, 26 Apr 1964 (1 ¢,2 2; JTP); La
Sabana, 20 Oct 1936 (1 36, 11 2; UK). Morelos: stream, 20 mi. S Cuerna-
vaca, 27 Apr 1964 (7 6, 6 2; JTP, PDK); stream, 5 mi. W Zacatepec, 25
Apr 1964 (2 6, 2 2; JTP); river, 4 mi. N Amacuzac, 25 Apr 1964 (2 6, 3
2; JTP); river, 8 mi. E Zacatepec, 25 Apr 1964 (1 6; JTP); stream, 2 mi.
S Cuautla, 25 Apr 1964 (21 6, 27 2; JTP, PDK); Acatlipa, 5 May 1944 (1
2; UK); Mazatepec, 29 Apr 1944 (11 ¢, 10 2; UK). Nayarit: river, Santa
Cruz, 8 Jun 1975 (1 6, 1 9; JTP); stream, 5 mi. E Tuxpan, 21 Apr 1964 (11

<—

Figs. 9-14. Trepobates polhemi. 9-11, Variation of color pattern in female. 12, Mesonotum
of female. 13, Posterior femur of female. 14, Paramere. (Scale lines = 0.5 mm except as noted.)

164 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

é.-12 9: JTP, PDK); 23 miz:NW Tepic, 10 Sep 1972; 1) 25 Use)
stream, 15 mi. E San Blas, 21 Apr 1964 (2 6; JTP); river, San Blas, 7 Jun
1966 (1 3, 1 2; JTP), 28 Nov 1968 (1 ¢6; PDK); San Blas, 17-21 Sep 1953
(1 2; JTP); spring, San Blas, 3 Jun 1966 (1 2; JTP). Sinaloa: Rio Presidio,
Villa Union Presidio, 21 Jul 1952 (1 6, 1 2; JTP); Arroyo Sonolona, 18.5
mi. E Culiacan, 2 Apr 1955 (7 6, 11 2; UM); Los Mayos, 24 Jul 1952 (5
6,11 9: JTP; PDK); 10:mi. N Los Mochis, 23°’ Apr 1977.4. 6, 2:22US0);
Sonora: Arroyo Cuchujaqui, 9 mi. E Alamos, 17 Feb 1957 (1 6, 1 2; UM);
Rio Cuchujaqui, 5 mi. E Alamos, 11 Jun 1974 (2 6; PDK); Arroyo El Cajou,
Cajou, 16 Feb 1957 (2 2, UM); Arroyo Cuchujaqui, Alamos, 21 Mar 1967(1 6;
JTP); Arroyo Cuchujaqui, 5 mi. ESE Alamos, 29 May 1966 (3 d, 2 2; JTP).

Distribution.—Trepobates polhemi has been collected in the Distrito Fed-
eral and five states of Mexico (Fig. 1).

Variation.—T7repobates polhemi is variable in color pattern (Figs. 9-11).
Metanotal hairs may be present or absent in the female. The alate form
apparently is uncommon and represented 14% of all specimens examined.

Biology.—This species has been collected from lotic habitats, including
creeks, rivers, and a large spring. Collection records are known for each
month from February through July and September through November.

Etymology.—This species is named in honor of John T. Polhemus, En-
glewood, Colorado, for his many contributions to our knowledge of the
systematics of aquatic and semiaquatic Hemiptera.

ACKNOWLEDGMENTS

I express appreciation to the following individuals and institutions who
loaned material for this study: Peter D. Ashlock, University of Kansas,
Lawrence (UK); Richard C. Froeschner, National Museum of Natural His-
tory, Washington, D.C. (USNM); Wilford J. Hanson, Utah State Univer-
sity, Logan (USU); Thomas E. Moore, University of Michigan, Ann Arbor
(UM); Nico Nieser, Rijksuniversiteit Utrecht, Netherlands (NN); John T.
Polhemus, Englewood, Colorado (JTP); Joseph C. Schaffner, Texas A&M
University, College Station (JCS, TAMU); Randall T. Schuh, American
Museum of Natural History, New York (AMNH); Cecil L. Smith, Univer-
sity of Georgia, Athens (CLS); and Kenneth W. Stewart, North Texas State
University, Denton (NTSU).

LITERATURE CITED

Drake, C. J. and H. M. Harris. 1932. Some miscellaneous Gerridae in the collection of the
Museum of Comparative Zoology (Hemiptera). Psyche (Camb. Mass.) 39: 107-112.

Hungerford, H. B. 1936. Aquatic and semiaquatic Hemiptera collected in Yucatan and Cam-
peche. Carnegie Inst. Wash. Publ. No. 457, pp. 145-150.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 165-166

A SECOND SPECIES OF CERATITIS (DIPTERA: TEPHRITIDAE)
ADVENTIVE IN THE NEW WORLD

GEORGE C. STEYSKAL

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Ceratitis malgassa Munro, a species described from Malagasy
(Madagascar), was reared in Mexico from nutmegs obtained from Puerto
Rico. A key to distinguish the species from its nearest relatives, including
C. capitata, is given.

In 1968 the Systematic Entomology Laboratory of the U.S. Department
of Agriculture received three adult specimens of a species of Ceratitis from
Chapingo, Mexico that had been reared there from nutmeg seeds received
from Mayaguez, Puerto Rico. Information in USDA files indicated that the
seeds had been gathered from trees planted in Mayagtiez with seeds received
from ‘‘Granada.”’

The specimens belong to the typical subgenus of Ceratitis, the generitype
of which is the Mediterranean fruit fly, C. capitata (Wiedemann). The sub-
genus is characterized by the presence only in the males of elongated, api-
cally spatulate anterior upper fronto-orbital bristles. These bristles are in-
serted on small conical bases on the upper part of the head a little distance
before the ocellar triangle. There are four species known in the subgenus.
Our specimens compare well with a pair of C. malgassa Munro borrowed
from the Museum of Natural History of Paris, France. Paulian (1953) gave
a key to the subgenus and stated that C. malgassa (incorrectly referred to
as C. malagassa) had been reared from oranges, mandarins, guavas, and
fruit of Sclerocarya caffra Sond. (Anacardiaceae; as Poupartia caffra). To
this list of hosts may now be added nutmeg (Myristicaceae; Myristica fra-
grans Houtt.). The four known species of Ceratitis sensu stricto may be
distinguished as in the following key, translated with modifications from
Paulian (1953). Ceratitis malgassa was described by Munro (1939: 141-143).

KEY TO SPECIES OF CERATITIS S.S.

1(4). Distal part of wing with 2 transverse bands; male orbital seta with
black palette.

166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

2(3). Orbital seta with more or less lozenge-shaped palette borne on
stem less than twice as long as palette; basal yellow scutellar mark
SIMNIALE . ..2 eau qaeenanns ow ee oer aan See capitata (Wiedemann)

3(2). Orbital seta with palette nearly round, borne on stem several
times as long as palette; basal yellow scutellar mark with 2 long,
posteriorly directed pomts. . . «<1. tee. Gace eee caetrata Munro

4(1). Distal part of wing with 3 transverse bands; orbital palette more
or less oval, whitish to pale brownish.

5(6). Mesoscutum with median dark stripe not connected with pair of
posterior spots; stem of spatulate seta round, bristly, palette about
AS NOMS AS WIGS i coca eee eee es eee catoirei Guérin

6(5). Mesoscutum with median dark stripe connected posteriorly with
pair of large spots; orbital seta with tapering; ribbon-like stem,
palette loneer thaniwide*. 22-05 SoA oe Seah. malgassa Munro

LITERATURE CITED

Munro, H. K. 1939. Some new species of South African Trypetidae (Diptera), including one
from Madagascar. J. Entomol. Soc. South Afr. 2: 139-153.

Paulian, R. 1953. Recherches sur les insectes d’importance biologique a Madagascar. XII. Les
mouches des fruits. Mém. Inst. Sci. Madagascar, Ser. E, 3: 1-7.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 167-176

NEW SPECIES OF UROSIGALPHUS (HYMENOPTERA: BRACONIDAE)
FROM SOUTH AMERICA

LESTER P. GIBSON

U.S. Department of Agriculture, Forest Service, Northeastern Forest
Experiment Station, Delaware, Ohio 43015.

Abstract.—Five new species of Urosigalphus are described. One, di-
versus, is the first species of the subgenus Microurosigalphus to be collected
in South America. One, alius, belongs to the subgenus Urosigalphus and
three, flexus, porteri, and surinamensis, belong to the subgenus Neourosi-
galphus. Changes in existing keys to South American species are presented.

This paper describes the first species of the subgenus Microurosigalphus
collected in South America. A new species of the subgenus Urosigalphus
is also described along with three new species of the subgenus Neourosi-
galphus. Changes in the existing keys to species (Gibson, 1974) within these
subgenera are provided.

Urosigalphus (Microurosigalphus) diversus Gibson, New Species
Figsicls2

General features —Length, 2.8 mm. Head and thorax deep reddish black;
carapace dark red brown; abdomen brown; clypeus dark red brown apically,
labrum yellow tan; mandible yellow tan, with red base and apex; palpi pale
yellow; antenna dark brown distally, becoming yellow basally with yellow
scape and pedicel; fore- and midlegs yellow; hindcoxa and femur yellow,
hindtibia yellow, distal tip of tibia and tarsi brown. Tegula yellow tan. Wings
clear, hyaline, with nearly colorless hairs, costa tan basally and light brown
apically; stigma medium brown, remaining veins medium brown distally but
becoming nearly colorless basally.

Male.—Head: General anterior outline quadrate. Vertex, temples, and
genal regions sparsely finely punctured. Lower face nearly flat, evenly finely
punctate; fronto-clypeal groove with a small, deep, oval anterior tentorial
pit near each end. Clypeus impunctate, rather quadrate; lower central mar-
gin straight. Labrum punctate. Antennal scapes separated with area between
fossae narrow, finely punctate with a small central carina, extending dorsally
toward median ocellus, and a shallow, broad, nitidous, sparsely, finely punc-

168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tate antennal sculcus extending feebly around rear ocelli. Antenna 16-seg-
mented; scape long and relatively slender, inner margin slightly rounded;
pedicel almost half as long as scape; Ist flagellar segment longer than 2nd.
Ocelli circular, on the ocellar triangle. Occipital carina apparently incom-
plete dorsally and moderately developed.

Thorax: Unevenly punctate. Mesonotum fairly regularly, finely punctate,
the general surface uneven; notaulices narrow, deep, with a row of mod-
erately large punctures, area of convergence broad, nearly square, de-
pressed, bisected centrally with a carina, with moderately large punctures;
median lobe regularly, finely punctate; lateral lobes gently rounded, sparse-
ly, finely punctured, with a small lateral carina; posterolateral carina weakly
flangelike. Scutellar groove broad and deep with 7 dissecting carinae and 2
bordering lateral carinae. Scutellum roundly trigonal in dorsal profile, with
very small sparse punctures; lateral and posterior surface with a line of
moderate punctures. Propodeum gently rounded posteriorly, entirely ru-
goso-punctate without carinae. Mesopleuron impunctate except for a line
of moderate punctures around anterior, dorsal, and posterior borders; meso-
sternum smooth nearly impunctate with a line of small punctures along
center line. Lateral regions of pronotum centrally impunctate, rest irregu-
larly punctured with small punctures. Tegula impunctate. Wings hyaline;
submedian cell longer than median cell basally.

Abdomen: Carapace elongate oval in dorsal view and teardrop-shaped in
lateral view; general surface rugoso-punctate anteriorly, punctate and shiny
posteriorly; apex of carapace without apical spines. Parameres of external
genitalia very short.

Female.—Unknown.

Holotype.—d, Argentina, ‘‘ Villa Nouges,’’ Tucuman, XI-26/28-64; 1250
m, C. Porter. In Museum of Comparative Zoology.

Biology and host—Unknown.

Remarks.—This is the only member of the subgenus Microurosigalphus
known from South America. The apex of the carapace is without tubercles
so it would key neomexicanus Crawford in existing keys (Gibson, 1972). It
differs from neomexicanus by leg, antenna, palpi, and tegula color. These
being predominately yellow in diversus and medium to dark brown in neo-
mexicanus. It is also the only known species of Microurosigalphus with 16
antennal segments.

KEY TO SPECIES OF SUBGENUS UROSIGALPHUS OF SOUTH AMERICA

1. Apex of carapace without tubercles
- Apex of carapace with tubercles
20): ‘Thorax and’carapace red ~.: /)0%.. G18 see rubicorpus Gibson
_ Thorax black, carapace red black or black

VOLUME 84, NUMBER 1 169

Figs. 1-7. 1, Urosigalphus (Microurosigalphus) diversus, showing open radial cell. 2, U.
(M.) diversus, type. 3, U. (Urosigalphus) alius, type. 4, U. (Neurosigalphus) surinamensis,
type. 5, U. (N.) flexus, type. 6, U. (N.) flexus, allotype. 7, U. (N.) porteri, type.

Palpi maroon, fore- and midlegs medium brown basally ........

eT en ee ee er ee er eee re trinidadensis Gibson
Palpi light brown, fore- and midlegs orange tan ... alius, new species
Carapace with short, poorly formed tubercles .. sanguineus Gibson
Carapace with long, well-developed tubercles ............---+--- 5

170 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

5(4). Carapace tubercles sharp pointed; wings clear ................
Me OD Bak GR cuits free. ee eee chalcodermi Wilkinson
- Carapace tubercles blunt tipped; wings tinted brown...........
eee. ho Js eee venezualaensis Gibson

Urosigalphus (Urosigalphus) alius Gibson, New Species
Fig. 3

General features —Length, 4.9 to 5.1 mm. Head black; thorax black;
carapace black to red black; abdomen dark brown; clypeus maroon apically;
labrum red; mandible orange with red base and black apex; palpi light
brown; antennal scape and pedicel deep maroon, pedicel with narrow tan
apical band, rest of antenna dark red brown; fore- and midlegs orange tan;
hindleg light red, hindtibia apex and tarsi dark red brown. Tegula dark red
brown. Wings hyaline, tinted brown, with brown hairs, costa dark red
brown, stigma dark red brown, remaining veins dark brown but becoming
paler towards base of the wing.

Female.—Head: General anterior outline quadrate with small ocellar pyr-
amid. Vertex coarsely rugoso-punctate; temples and genal regions finely
punctate. Face finely punctate, fronto-clypeal groove well impressed with
small round anterior tentorial pits. Clypeus transverse; finely punctate; low-
er part wide, margin reflexed and gently concave. Labrum evenly punc-
tured. Mandible surface finely rugoso-punctate. Antennal scapes well sep-
arated, with 2 weak carinae dividing the intervening area into 3 equal rugoso-
punctate parts, extending dorsally and joining ocellar pyramid carinae; a
broad, rugoso-punctate antennal sulcus. Antenna 14-segmented; scape short
and broad, arcuate; pedicel short, nearly round; Ist and 2nd flagellar seg-
ments subequal. Ocelli small and ovoid. Occipital carina complete.

Thorax: Shorter than carapace, and irregularly punctate. Mesonotum ir-
regularly punctate; notaulices broad, fairly deep, with large punctures, area
of convergence quadrate, deeply, coarsely rugoso-punctate; median lobe
finely punctate with or without a poorly formed small central carina ante-
riorly and a transverse carina, coarsely rugoso-punctate, laterally; lateral
lobes flattish, finely punctured, with lateral carina bordered by large punc-
tures, posterolateral carina well developed and flangelike. Scutellar groove
rather narrow and deep with 3 strong central carinae. Scutellum roundly
trigonal in dorsal outline, with moderate to large irregular punctures and
moderate to strong rugae forming longitudinal carinae dorsally, coarsely
rugoso-punctate and slightly concave laterally. Propodeum flat posteriorly,
with a moderately strong transverse carina; dorsal area narrow in lateral
profile, the posterior surface abruptly dropping to point of abdominal artic-
ulation; central dorsal carina strongly developed, joining the transverse ca-
rina which proceeds down both sides in an arc past the point of abdominal
articulation; dorsal surface with strong carinae delimiting large, irregular

VOLUME 84, NUMBER 1 171

depressions; lateral areas rounded in cross-section and irregularly, strongly
rugoso-punctate. Mesopleuron smooth centrally and sparsely, finely punc-
tate; upper margin coarsely rugoso-punctate with a vertical carina; lower
central area with a few elongate punctures in the mesopleural groove; pos-
terior margin with a row of large punctures divided by short carinae and a
bordering carina; mesosternum rounded and finely, densely punctate, sec-
tioned by low carinae with a punctate median line. Lateral regions of prono-
tum moderately punctate; the posterior corner similar behind a carina. Teg-
ula broadly pear shaped. Wings hyaline; the stigma long and broad,
submedian cell longer than median.

Abdomen: Carapace elongate oval; general surface longitudinally rugoso-
punctate, tending to form carinae dorsoanteriorly; apex of carapace round-
ed, grooved ventrally, without a pair of tubercles. Ovipositor 1.0 mm long,
sheath brownish-black clothed with fine hairs.

Male.—Similar to female in general structure and color except for cus-
tomary sexual differences. Parameres of external genitalia moderately long
and narrow, dark brown.

Holotype.—@, Surinam-Lelydorp; Sumatra Road; 23—26-III-1964, D. C.
Geijskes, sandridges in savannah forest. In Rijksmuseum van Natuurlijke
Historie, Lieden, Netherlands.

Allotype.—d, 5 Paratypes. Surinam-Lelydorp; Sumatra Road. (1 ¢) 17-
21-IV-1964; (1 2) 23-26-II-1964; (1 2) 26-31-III-1964; (1 2) 4-7-I1V-1964;
D. C. Geijskes; sandridges in savannah forest. In Rijksmuseum van Na-
tuurlijke Historie. (2 2) 17-21-IV-1964. In National Museum of Natural
History, Washington, D.C.

Biology and host.—Collected from sand ridges in savannah forest in
March and April.

Remarks.—This species is close to trinidadensis but separated by the
light brown palpi and orange-tan fore- and midlegs. Urosigalphus trinida-
densis has maroon palpi and medium brown fore- and midlegs.

KEY TO SOUTH AMERICAN SPECIES OF SUBGENUS NEOUROSIGALPHUS

VB Carapace with apical tubercles, antenna 14-segmented ......... 2

- Carapace without apical tubercles, antenna 14- to 17-segmented
aries. rei. Leas uiriel . orowren ck Beet AE ie CM ee ee 9

2(1). Carapace with apical tubercles very small to obsolete ......... 3
Carapace with apical tubercles well developed ................ 4

3(2). Hindleg dark red-brown; body length over 4.5mm ..........
en ee 22 cian aking Meiecuol Saag inant pullatus Gibson

4) altindiemundark:red brow) ¢ tee snGasanidateh ligt. a acutus Gibson
_ Hindfenumnot darkired brown il.as ceo a0) .seweed. dank. foe 5

11(10).

12¢LL).
13(2):

14(13).

15(15)»

16(9).

17(16).

Hindfemur medium red brown ........... eulechriopis Cushman
Hindfemur' tan to'red' orange ()2% 222 ale ee 6
Head andithorax maroone +o) Sees ieee tuberculatus Gibson
Head: and! thorax black: «0311! 248 eae a Se q
Carapace black ornearly black”). -23.). 9u..902 9. I. Oe eee 8
Carapace red anteriorly, black posteriorly ....... bicolor Gibson
Legs yellow tan; ovipositor 0.5 mm long ....... spinatus Gibson
Legs red orange; ovipositor 1.2 mm long ...... bidentata Gibson
Antéenna14-seemented | 2.14.02. 2 eee 16
Antenna-16-or 17-segmented .. .. 2032-280 Ae ee 10
Antenna 17-segmented; interantennal area with a large central

carina; body length over 5.0mm ....... pseudochelonus Gibson
Antenna 16-segmented; interantennal area without a large cen-

tral.carina; ‘body length less ‘than’5.0 mmf! 2,440- 22 Peer i

Hindleg dark red brown to black; palpi dark brown..........
Pi Le A) NAA LAER, SO) ERR RS ASI braziliensis Gibson

Hindleg yellow to dark brown; palpi yellow or tan ........... 12
Hindcoxa orange; trochanters tan. 2 eee ee ornatus Gibson
Hindcoxa mediumto dark. brown) <2-0.2¢e6 22 22k eee eee 13
Fore- and midcoxa medium to dark brown .................- 14
Foré- andmidcoxa)tan( ‘2 4s. sate ee Fo eee 15

Trochanters tan; tegula dark brown, carapace apex smooth...
be SUA edna eae es Rik gd SRR Cae eer cautus Gibson
Trochanters medium brown; tegula medium brown, carapace
ape Pllnctate> agen eid ac aes See eee obscurus Gibson
Wings tinted brown; apex of carapace finely rugoso-punctate
Gastctrail. 1k Sovaes) Sle aa oo eas surinamensis, new species

Wings clear; apex of carapace impunctate ...... safflavus Gibson
With apical projections on ventral flange of carapace; palpi light
Red). 5.05 (see SUGy SR. Se ee paraguayensis Gibson
Without apical projections on carapace flange; palpi tan or yel-
DOW ni ak a ol oD acdin meee GERRI reg Gaon cs Seo 17
Palpi tan; hindcoxa dark red basally; hindfemur and tibia yellow
ordark red-brown (eee ee ee flexus, new species
Palpi yellow; hindcoxa brown; hindfemur brown; hindtibia
brown; darker dorsally. s35e-e 37202 eee porteri, new species

Urosigalphus (Neourosigalphus) flexus Gibson, New Species
Figs. 5, 6

General features—Length 2.2-3.3 mm. Head and thorax black; carapace
black; abdomen brown; clypeus red black to black apically, labrum deep
maroon; mandible rufo-testaceous with dark red base and apex; palpi tan,
antenna dark brown; forelegs yellow tan; midlegs brown or yellow tan;

VOLUME 84, NUMBER I 173

hindlegs with coxa dark red basally, hindfemur and tibia yellow or femur,
tibia, and tarsus dark red brown. Tegula dark brown to red brown. Wings
clear, hyaline, with nearly colorless hairs, costa light brown; stigma medium
to dark brown, remaining veins medium brown distally but becoming pale
basally.

Male.—Head: General anterior outline roundly quadrate. Vertex, temples
and genal regions finely punctured. Lower face rounded, evenly, finely
punctate; fronto-clypeal groove with a small, deep, oval anterior tentorial
pit near each end. Clypeus quadrate; lower central margin gently concave.
Mandible teeth long and narrow, outer surface punctate. Antennal scapes
well separated, area between fossae smoothly rounded and rugoso-punctate
without a small central carina and with a shallow, narrow, nitidous, sparse-
ly, finely punctate antennal sulcus extending to or slightly past rear ocelli.
Antenna 14-segmented; scape long and relatively slender, inner margin
slightly rounded; pedicel almost half as long as scape. Ocelli ovoid, inter-
ocellar area rugose, slightly grooved with slightly raised areas centrally be-
tween groove and rear ocelli. Occipital carina complete and well developed
throughout.

Thorax: Unevenly punctate. Mesonotum fairly regularly punctate, general
surface uneven; notaulices broad and shallow, with moderately large punc-
tures, area of convergence broad, quadrate, depressed, with moderately
large punctures; median lobe densely, finely punctate without a small, short
central carina anteriorly; lateral lobes gently rounded, sparsely, finely punc-
tured, with a small lateral carina; posterolateral carina weakly flangelike.
Scutellar groove rather broad and deep with 3 to 5 dissecting and 2 bordering
carinae. Scutellum roundly trigonal in dorsal profile, with irregular moderate
to large punctures; posterior surface gently rounded. Propodeum gently
sloped posteriorly, with a moderate, irregular transverse carina; central dor-
sal carina short and prominent, transverse carina becoming lost laterally in
the lateral rugosity; dorsal surface and posterior surfaces roughly and dense-
ly rugoso-punctate; lateral areas irregularly rugoso-punctate. Mesopleuron
punctate; middle central area impunctate, lower anterior area with moderate
punctures in a mesopleural groove; mesosternum rugoso-punctate or punc-
tate. Lateral regions of pronotum punctate, upper posterior corner with a
slight bend, the triangular area behind bend sparsely, finely punctate and
centrally carinate. Wings hyaline; submedian cell as long as median basally.

Abdomen: Carapace teardrop shaped in dorsal view and comma shaped
in lateral view; general surface longitudinally rugoso-punctate anteriorly,
densely punctate posteriorly; apex of carapace without apical spines. Par-
ameres of external genitalia very short.

Female.—Similar to ¢ but carapace oval in dorsal view and teardrop
shaped in lateral view. Ovipositor 0.75 mm long, 2 as long as carapace.
Ovipositor sheath brown.

174 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Holotype.—é, Argentina, ‘Villa Nouges,’”” Tucuman; X1-26/28/64 7250
m, C. Porter. In Museum of Comparative Zoology.

Allotype.—?, Same data as holotype.

Paratype.—¢, Same data as holotype. In National Museum of Natural
History.

Biology and host.—Unknown.

Remarks.—This species is close to paraguayensis by the presence of 14
antennal segments and lack of tubercles on carapace apex but differs in lack
of ventral flange projections and tan palpi color.

Urosigalphus (Neourosigalphus) porteri Gibson, New Species
Fig. 7

General features.—Length, 2.9 mm. Head and thorax black; carapace
black with maroon tint basally; abdomen rufo-testaceous; clypeus dark red
brown apically, labrum red; mandible yellow with dark red brown base and
apex; palpi yellow; antenna dark brown; fore- and midlegs yellow tan; hind-
legs with coxa and femur brown, tibia and tarsus dark brown dorsally.
Tegula brown. Wings clear, hyaline with colorless hairs, costa light brown
basally and medium brown apically; stigma dark brown, remaining veins
medium brown distally but becoming paler basally.

Female.—Differs from flexus in following: Head: Clypeus semi-circular.
Interantennal area with a shallow, finely punctate rugoso-punctate antennal
sulcus extending past rear ocelli. Interocellar area grooved with slightly
raised rugoso-punctate areas between groove and rear ocelli.

Thorax: Mesonotum notaulices broad and deep. Scutellar groove with 3
dissecting and 2 lateral carinae. Scutellum rugoso-punctate with irregular
large punctures. Propodeum with dorsal surface with irregular rugae delim-
iting irregular depressions; posterior surface roughly and densely rugoso-
punctate. Mesopleuron lower anterior area with large punctures in a me-
sopleural groove; mesosternum coarsely rugoso-punctate. Lateral regions
of pronotum punctate, with deep, moderate punctures.

Abdomen: Carapace oval in dorsal view and somewhat teardrop shaped
in lateral view. Ovipositor 1 mm long, 4/; as long as carapace, ovipositor
sheath brown.

Male.—Unknown.

Holotype.—@, Quebrada-Lules, Tucuman, Argentina 9-XII-64; C. C.
Porter. In Museum of Comparative Zoology.

Distribution.—Known only from type.

Biology and host.—Unknown.

Remarks.—Urosigalphus porteri is close to flexus, but flexus differs in
that its palpi color is yellow and the hindleg and coxa are brown.

VOLUME 84, NUMBER | 175

Urosigalphus (Neourosigalphus) surinamensis Gibson, New Species
Fig. 4

General features.—Length 4.2 mm. Head and thorax black; carapace dark
red black; abdomen dark brown; clypeus dark red brown apically; labrum
red tan; mandible red tan with red apex; palpi tan; antenna medium brown
with dark brown scape and pedicel; foreleg tan, midleg tan, hindleg dark
brown with tan trochanter; coxae same color as legs. Tegula red tan. Wings
hyaline with brownish tint, with sparse brown hairs, costa and stigma
brownish black, remaining veins pale brown.

Female.—Differs from flexus in following: Head: Clypeus rather quad-
rate; lower central margin gently curved. Mandible outer surface fairly
smooth. Interantennal area densely, finely punctate and with three small
carinae just anterior to the median ocellus, and a fairly deep, finely punctate
antennal sulcus extending almost to the rear ocelli. Antenna 16-segmented;
scape short and relatively slender; inner margin arcuate; pedicel amost half
as long as scape; apical segment of flagellum 4 longer than the penultimate
segment. Ocelli oval, interocellar area deeply, finely punctate, slightly
raised. Occipital carina incomplete, poorly developed laterally and absent
in dorsal region.

Thorax: Mesonotum irregularly punctate, with notaulices narrow and
shallow with large-sized punctures, area of convergence broad and round,
with large-sized punctures; median lobe finely punctate with a small central
carina; lateral lobes flattened, densely, finely punctured, with a small lateral
carina; posterolateral carina not flangelike. Scutellar groove deep, seg-
mented by 5 carinae. Scutellum coarsely, deeply, rugoso-punctate and car-
inate dorsally, with irregular moderate, carinate punctures laterally and pos-
teriorly. Propodeum nearly flat posteriorly, with a strong transverse carina;
the transverse carina proceeds irregularly down both sides to the point of
abdominal articulation; dorsal surface irregularly carinate and rugoso-punc-
tate; posterior surface roughly and densely rugoso-punctate and bicarinate.
Mesopleuron mostly impunctate; central area impunctate and nitidous, sur-
rounded by large, deep punctures; mesosternum bicarinate and rugoso-
punctate with large deep punctures. Lateral regions of pronotum irregularly
punctured with moderate, fairly deep punctures, except impunctate cen-
trally; upper posterior corner with a carinate bend, triangular area behind
carina with large deep punctures. Wings hyaline, with a brown tint; sub-
median cell longer than median basally.

Abdomen: Carapace elongate oval in dorsal view and teardrop shaped in
lateral view; general surface longitudinally finely carinate dorsally, and fine-
ly rugoso-punctate distally. Ovipositor 1.5 mm long, nearly as long as car-
apace; Ovipositor sheath reddish brown.

176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Male.—Unknown.

Holotype.—@, Surinam, Phedra, 7 to 14-XII-64; D. C. Geijskes, rain
forest in hilly interior. In Rijksmuseum van Natuurlijke Historie.

Biology and host.—Unknown.

Remarks.—This species differs from all known South American species
by the brown tinted wings. It is similar to braziliensis Gibson in size and
keys near safflavus Gibson but is easily separated by the brown tinted wings
and finely, rugoso-punctate carapace apex.

ACKNOWLEDGMENTS

I thank Dr. Kees van Achterberg for the loan of specimens from the
Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands, and Alfred F.
Newton, Jr. for the loan of specimens from the Museum of Comparative
Zoology, Harvard University, Cambridge, Massachusetts.

LITERATURE CITED

Gibson, L. P. 1972. Revision of the Genus Urosigalphus of the United States and Canada
(Hymenoptera: Braconidae). Misc. Publ. Entomol. Soc. Am. 8(3 & 4): 83-134.
—. 1974. South American Urosigalphus (Hymenoptera: Braconidae). Misc. Publ. Ento-

mol. Soc. Am. 9(4): 201-226.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 177-183

COCCOBAPHES SANGUINARIUS AND LYGOCORIS VITTICOLLIS
(HEMIPTERA: MIRIDAE): SEASONAL HISTORY AND DESCRIPTION
OF FIFTH-INSTAR, WITH NOTES ON OTHER MIRIDS
ASSOCIATED WITH MAPLE

A. G. WHEELER, JR.

Bureau of Plant Industry, Pennsylvania Department of Agriculture, Har-
risburg, Pennsylvania 17110.

Abstract.—Seasonal history of the mirids Coccobaphes sanguinarius
Uhler and Lygocoris vitticollis (Reuter) was followed on red maple, Acer
rubrum L., in southcentral Pennsylvania during 1979-80. Both are univol-
tine species whose nymphs feed on the undersides of expanding foliage.
Although overwintered eggs of both species hatched in early May, adults
of L. vitticollis first appeared in early June, about 7-10 days before those
of C. sanguinarius. Feeding by L. vitticollis produces translucent spots at
sites of stylet penetration on red maple leaves; when dead tissue tears or
drops from the feeding sites, the foliage appears ragged or perforated. A
diagnosis of the fifth-instar of both mirids is given, along with biological
notes on L. hirticulus (Knight) and other mirids found associated with Penn-
sylvania maples.

During studies on the mirid fauna of Pennsylvania, T. J. Henry and I
accumulated data on host plants and phenology of the species associated
with maple, Acer spp. I began a more intensive study of the fauna when
my colleague K. Valley brought to my attention the unusual injury that
Lygocoris vitticollis (Reuter) produces on leaves of red maple, A. rubrum L.
In this paper I summarize the seasonal history and host plants of the two
principal mirids occurring on Pennsylvania maples, L. vitticollis and Coc-
cobaphes sanguinarius Uhler, and give a diagnosis of the fifth-instar
nymphs. Notes on seasonal history are given for L. hirticulus (Knight) and
other mirids collected from but not restricted to breeding on maple.

StubDY SITES AND METHODS

The main study area near Harrisburg (Dauphin Co.), Pennsylvania, was
dominated by a dense stand of Virginia pine, Pinus virginiana P. Mill., with
an understory of grasses and forbs mixed with the rosaceous shrub ninebark,

178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Physocarpus opulifolius (L.) Maxim., and red maple. The maples were
young trees about 2-3 m high, plus smaller saplings.

I sampled red maple weekly from mid-May to late June 1979. In April
1980 I began to collect from red maple to determine first egg hatch of L.
vitticollis and C. sanguinarius and then made weekly collections through
July once eggs had begun to hatch (early May). Mirids were sampled by
beating branches over a small tray (Wheeler, 1980) or sweeping branches
with a beating net. All specimens taken in approximately 15 minutes of
collecting were identified to stage in the field (adults, late instars) and re-
turned to the host plants; instars I-III were placed in alcohol and sorted in
the laboratory under a binocular microscope. Populations of L. vitticollis
and C. sanguinarius were much smaller than those known for several or-
thotyline mirids, e.g., Diaphnocoris chlorionis (Say) (Wheeler and Henry,
1976), or Brachynotocoris puncticornis Reuter (Wheeler and Henry, 1980).
Thus, a typical sample consisted only of 3-7 specimens of each species
rather than the many hundreds of certain orthotylines that could be collected
in 15 minutes. To determine possible injury to host foliage by the two main
species, nymphs of different stages were placed in small plastic dishes con-
taining fresh sprigs of red maple, and any feeding damage was noted.

Lygocoris vitticollis (Reuter)

This mirid, widely distributed in eastern North America, is known from
Nova Scotia south to Mississippi and west to Kansas (Kelton, 1971). Ly-
gocoris vitticollis, along with the ubiquitous tarnished plant bug, Lygus
lineolaris (Palisot de Beauvois), and fourlined plant bug, Poecilocapsus lin-
eatus (F.), was one of the first North American mirids reported to damage
shade trees or ornamental plants. In the spring of 1882 Murtfeldt observed
injury to young foliage of silver maple, A. saccharinum L., at Kirkwood,
Missouri; unfolding leaves, initially stippled with transparent spots, soon
became perforated (Murtfeldt, 1887). Uhler (1887), unaware of Reuter’s
(1876) description published without a host association, described Murt-
feldt’s ‘“‘noxious capsid’’ as a new species, Lygus monachus. Uhler’s name
thus is a junior synonym of Reuter’s vitticollis (Reuter, 1909).

Since Murtfeldt’s time, there apparently have been no further reports of
L. vitticollis damaging maple, and little information on habits has been
added. Murtfeldt was perceptive to associate the tattered appearance of
maple leaves with a sucking, rather than chewing, insect, but some of her
observations, made before life histories of North American mirids were
generally known, have proved inaccurate.

Seasonal history and habits —Eggs that overwintered near dormant leaf
buds of red maple began to hatch shortly after leaf flush, about 7 May in
1980. Early instars fed on the undersides of tiny, expanding leaves; the pale,
whitish-green nymphs resembled the color of this young foliage. The pop-

VOLUME 84, NUMBER 1 179

Egg

1st Instar

2nd Instar

3rd Instar

4th Instar

5th Instar

Adult

1 8 15 22 29 5 12 19 26 3 10 17 24
Ken nl ee nea June ene eee July opens!

Fig. 1. Generalized seasonal history of Lygocoris vitticollis (shaded bar) and Coccobaphes
sanguinarius (open bar) in central Pennsylvania, 1979-80.

ulation consisted mainly of third-instars by mid-May and fifth-instars by the
last week of May in both years of study (Fig. 1). Adults began to appear
during the first week of June and were present only for 3-4 weeks, males
dying about a week before females. In the weekly collections, adults of this
univoltine mirid no longer were present after 21 June in 1979; two females
were collected on 30 June in 1980. The latest known collection of L. vitti-
collis in Pennsylvania is 22 July at Patton by the hemipterist Rev. Modestus
Wirtner (Wheeler and Henry, 1977).

Injury became noticeable when the population consisted mainly of third-
instars (mid-May) and appeared as translucent, irregularly shaped blotches
on curled, unfolding leaves (Fig. 2). In the laboratory nymphs provided with
fresh foliage produced this characteristic damage within 12-24 hours. As
leaves expand, the thin tissue remaining at the feeding site tears or drops
out (Fig. 3). The resulting ragged appearance of the foliage easily could be
blamed on feeding by lepidopteran larvae or some other chewing insect.

180 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 2, 3. Injury to red maple by Lygocoris vitticollis. 2, Translucent areas on expanding
foliage. 3, Ragged edges and holes resulting from dead tissue tearing and dropping from leaf.

Injury seemed disproportionate to the size of populations since fewer than
ten bugs were taken in any sample during 1979-80. Populations, however,
may fluctuate from year to year; Murtfeldt (1887) noted that populations
were greater in 1886 than in her previous seasons of observation. At only
two localities in New York, Knight (1917) was able to collect unusually
large numbers (250 and 118 adults) from sugar maple, A. saccharum H.
Marsh.

Host plants.—The principal hosts are silver maple (Murtfeldt, 1887) and
red and sugar maple (Knight, 1917). In addition to the population studied
on red maple, I collected L. vitticollis from this host in other old fields and
in hedgerows and nurseries, mostly on cut-over trees or saplings. I also
found this species breeding on silver maple in Pennsylvania and have seen
a series of specimens taken on silver maple in Berkeley Co., West Virginia.

Lygocoris vitticollis appears to be a specialist mirid confined to maple.
Records from other plants, e.g., Ulmus sp. (Akingbohungbe et al., 1972),
probably reflect dispersal of adults from maple. Adults of Lygocoris spp.
show a strong tendency to disperse to non-host plants (Kelton, 1971; Wheel-
er, unpublished data).

Description of fifth-instar—Length 4.68 mm. Elongate oval, uniformly
pale or whitish green, apex of labial segment IV darker. Sparsely clothed
with pale, recumbent setae, intermixed with a few semierect setae. Antenna:
I, length 0.48 mm; II, 1.46 mm; III, 1.22 mm; IV, 0.66 mm. Rostrum, length
1.20 mm, reaching bases of mesocoxae. Wing pads reaching 4th abdominal
segment; dorsal abdominal scent gland opening pale, indistinct.

On maple, nymphs of L. vitticollis may be distinguished from those of L.
hirticulus by their pale, uniformly whitish-green color. Knight (1917) was
the first to note that vitticollis nymphs differed from those of other species

VOLUME 84, NUMBER 1 18]

of Lygocoris (cited as the communis group or Group VI of the genus Lygus
Hahn).

Coccobaphes sanguinarius Uhler

Although the recorded distribution of C. sanguinarius is similar to that
of L. vitticollis—Ontario south to Florida and west to Missouri (Carvalho,
1959)—its habits are not as well known. Drake (1922) found this mirid on
striped maple, A. pennsylvanicum L.; Knight (1923) recorded the second
growth of young sugar, and less often, red maple as hosts; and Watson
(1928) added silver maple as a host. I am aware of only one reference to C.
sanguinarius in the economic literature. At Knoxville, Tennessee, a popu-
lation on ‘“‘hard maples’’ attracted attention when large numbers of this
brilliant red bug fell to the ground beneath host trees (Bentley, 1941).

Seasonal history and habits.—Overwintered eggs, like those of L. vitti-
collis, hatched during early May, and nymphs fed on the lower surfaces of
young, expanding foliage. Based on two seasons’ observations, nymphal
development appeared to take somewhat longer, so that populations of C.
sanguinarius consisted mainly of fifth- and a few fourth-instars when adults
and fifth-instars of L. vitticollis were present (Fig. 1). Adults of C. sangui-
narius first appeared in the 12 June collection and were present at the sample
site until mid-July. The latest Pennsylvania record of this single-brooded
species is | Aug., based on Rev. Wirtner’s collecting at Patton (Wheeler
and Henry, 1977).

Since Coccobaphes nymphs fed on new growth of red maple in company
with Lygocoris nymphs, it was not possible from field observations to de-
termine whether C. sanguinarius also injured host foliage. In the laboratory,
nymphs did not produce any visible damage no matter how long they were
confined with unblemished leaves of red maple.

The number of specimens taken in both seasons was small, with a typical
sample consisting of 4-6 specimens. Froeschner (1949) noted that in Mis-
sourl C. sanguinarius was ‘“‘rather scarce.”

Host plants.—Although adults have been collected from beech, Fagus
sp. (Van Duzee, 1889), serviceberry or Juneberry, Amelanchier sp. (Wheel-
er and Henry, 1977), and mountain ash, Sorbus sp. (Drake, 1922), there is
no evidence that plants other than Acer spp. serve as hosts. Like L. vitti-
collis, C. sanguinarius is most abundant on sapling and second-growth red
and sugar maple. In addition to Knight’s (1923) record of sanguinarius from
red and sugar maple, this mirid has been taken on the latter host at Cincin-
nati, Ohio (U.S. National Museum collection). I have seen a series of spec-
imens collected from striped maple at Cranberry Lake, New York (Cornell
University collection).

Description of fifth-instar.—Length 4.40 mm. Broadly oval, uniformly
bright red except antennal segment IJ, apex of labial segment IV, and tibiae

182 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

fusco-reddish to black; and antennal segments III-IV and tarsi white. Head,
pronotum, and wing pads with short erect and semierect pale setae; abdo-
men more sparsely set with pale setae, longer bristle-like setae along lateral
margins of segments IX—X. Antenna with black bristle-like setae, becoming
more fine and sparse on segments III-IV; I, length 0.88 mm; II, 2.06 mm;
Ill, 1.28 mm; IV, 1.40 mm. Rostrum, length 2.00 mm, reaching bases of
metacoxae. Wing pads reaching abdominal segment VI; dorsal abdominal
scent gland opening indistinct, surrounded by brighter red area.

Nymphs of C. sanguinarius may be separated easily from nymphs of
other mirid species breeding on maple. They are distinguished by their bright
red color with contrasting white on the last two antennal segments and the
tarsi.

OTHER SPECIES

Several mirids whose hosts include various hardwoods were found breed-
ing on maple in Pennsylvania. The mirine Lygocoris hirticulus was the most
common of these additional Acer inhabitants. This species was present on
red maple at the sample site in both years of study, with adults appearing
the first week of June along with those of L. vitticollis. Nymphs of the two
Lygocoris species often are distinguishable in the field; those of L. hirticulus
differ from those of vitticollis by being yellowish green rather than pale
whitish green. Adults of L. hirticulus were present at the main sample site
until mid-July. This phytophagous, univoltine mirid, collected previously on
maple (Hussey, 1922; Blatchley, 1926), also breeds on other deciduous
trees, especially birch (Betula spp.) and beech (Fagus spp.) in Pennsylva-
nia.

Other mirids found breeding on Pennsylvania maples include the preda-
tors Deraeocoris nebulosus (Uhler), Diaphnocoris provancheri (Burque)
(=pellucida (Uhler)), Hyaliodes sp. (nymphs only—harti Knight or vitri-
pennis (Say), or both), and Phytocoris spp. Deraeocoris nebulosus was
common on sycamore maple, A. pseudoplatanus L., heavily infested with
the aphid Drepanosiphum platanoides (Schrank) (Wheeler et al., 1975).
Diaphnocoris provancheri, a species of at least partially predacious habits,
is known to occur on maple (Hussey, 1922).

ACKNOWLEDGMENTS

I thank my colleagues J. F. Stimmel for the photographs and K. Valley
for calling my attention to damage inflicted by L. vitticollis and for reading
the manuscript. E. R. Hoebeke, Cornell University, Ithaca, N.Y., also read
the manuscript.

LITERATURE CITED

Akingbohungbe, A. E., J. L. Libby, and R. D. Shenefelt. 1972. Miridae of Wisconsin (He-
miptera: Heteroptera). Univ. Wis.-Madison Res. Div. R 2396. 24 pp.

VOLUME 84, NUMBER 1 183

Bentley, G. M. 1941. A mirid (Coccobaphes sanguinareus Uhl.). Insect Pest. Surv. Bull. 21:
205.

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America, with special
reference to the faunas of Indiana and Florida. Nature Publ. Co., Indianapolis. 1116 pp.

Carvalho, J. C. M. 1959. Catalogue of the Miridae of the world. Part IV. Arg. Mus. Nac., R.
de J. 48: 1-384.

Drake, C. J. 1922. Heteroptera in the vicinity of Cranberry Lake, pp. 54-86. Jn Osborn, H.
and C. J. Drake, An ecological study of the Hemiptera of the Cranberry Lake region,
New York. N.Y. State Coll. For. Tech. Publ. 16.

Froeschner, R. C. 1949. Contributions to a synopsis of the Hemiptera of Missouri, Pt. IV.
Am. Midl. Nat. 42: 123-188.

Hussey, R. F. 1922. Hemiptera from Berrien County, Michigan. Occas. Pap. Mus. Zool.
Univ. Mich. No. 118, 39 pp.

Kelton, L. A. 1971. Review of Lygocoris species found in Canada and Alaska (Heteroptera:
Miridae). Mem. Entomol. Soc. Can. 83: 1-87.

Knight, H. H. 1917. A revision of the genus Lygus as it occurs in America north of Mexico,

with biological data on the species from New York. Cornell Univ. Agric. Exp. Stn.

Bull. 391: 555-645.

. 1923. Family Miridae (Capsidae), pp. 422-658. In Britton, W. E., ed., Guide to the

insects of Connecticut. Part IV. The Hemiptera or sucking insects of Connecticut. Conn.

State Geol. Nat. Hist. Surv. Bull. 34.

Murtfeldt, M. E. 1887. Notes from Missouri for the season of 1886, pp. 59-65. Jn Reports of
observations and experiments in the practical work of the Division, made under the
direction of the Entomologist. U.S. Dep. Agric. Div. Entomol. Bull. 13.

Reuter, O. M. 1876. Capsinae ex America boreali in Museo Holmiensi asservatae, descriptae.
Ofv. Svenska Vet.-Ak. Forh. 32(9): 59-92(1875).

—. 1909. Bemerkungen liber nearktische Capsiden nebst Beschreibung neuer Arten. Acta
Soc. Sci. Fenn. 36(2): 1-86.

Uhler, P. R. 1887. A new noxious capsid. Can. Entomol. 18: 208-209 (1886).

Van Duzee, E. P. 1889. List of Hemiptera from the Muskoka Lake District, Canada. Can.
Entomol. 21: 1-11.

Watson, S. A. 1928. The Miridae of Ohio. Ohio Biol. Surv. Bull. 16: 1-44.

Wheeler, A. G., Jr. 1980. Life history of Plagiognathus albatus (Hemiptera: Miridae), with
a description of the fifth instar. Ann. Entomol. Soc. Am. 73: 354-356.

Wheeler, A. G., Jr. and T. J. Henry. 1976. Biology of the honeylocust plant bug, Diaphnocoris
chlorionis, and other mirids associated with ornamental honeylocust. Ann. Entomol.

Soc. Am. 69: 1095-1104.

. 1977. Rev. Modestus Wirtner: Biographical sketch and additions and corrections to

the Miridae in his 1904 list of western Pennsylvania Hemiptera. Great Lakes Entomol.

10: 145-157.

. 1980. Brachynotocoris heidemanni (Knight), a junior synonym of the Palearctic B.
punticornis Reuter and pest of European ash. Proc. Entomol. Soc. Wash. 82: 568-575.

Wheeler, A. G., Jr., B. R. Stinner, and T. J. Henry. 1975. Biology and nymphal stages of
Deraeocoris nebulosus (Hemiptera: Miridae), a predator of arthropod pests on orna-
mentals. Ann. Entomol. Soc. Am. 68: 1063-1068.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 184-190

NEW CENTRAL AND SOUTH AMERICAN LEAFHOPPERS OF THE
‘*BAHITA’? GROUP (HOMOPTERA: CICADELLIDAE:
DELTOCEPHALINAE)

DwiGHT M. DELONG

Department of Entomology, The Ohio State University, Columbus, Ohio
43210.

Abstract.—Seven new species and a new genus of the “‘Bahita’’ group of
Deltocephaline leafhoppers, Perubahita confusa n. sp., Angubahita atra, n.
gen.,n. sp., Megabahita patula, n. sp., Parabahita lamina, n. sp., P. thalla,
n. sp., Frequenamia particula, n. sp., and F. atrata, n. sp., are described.

The genus Bahita was described by Oman (1936). DeLong (1947) de-
scribed the genus Frequenamia from Mexican specimens. Several related
genera (including Parabahita) were described by Linnavuori (1959). The
‘‘Bahita’’ group of genera was treated by Linnavuori and DeLong (1978a,
b). A new genus and seven new species are described in this paper. All
types are in the DeLong collection.

Perubahita confusa DeLong, New Species
Figs. 1-6

Description.—Length of male 6 mm, female unknown. Crown longer at
middle than next to eyes, % as long at middle as wide at base between eyes.
Crown dull white, a dark brown spot at each side of apex which becomes
paler brown caudally, and both spots merge near middle of crown. With
dark brown spot, each side, between eye and middle line at base. Basal %
of crown mostly tinted with orange brown. Pronotum brown with irregular
black markings and a narrow whitish transverse band across middle of
pronotum. Scutellum with basal angles mostly dark brown, with a small
white triangle in each basal angle, apical ’% white. Forewing pale brownish
subhyaline with a few dark brown spots; veins mostly dark brown.

Male genital plates 3x as long as wide at middle, apex narrow, bluntly
pointed, styles with apophyses slender, curved outwardly. Aedeagus
curved, tapered from rather broad base to slender pointed apices. Apical

portion composed of 2 lateral, contiguous portions. Pygofer narrowed api-
cally and rounded.

VOLUME 84, NUMBER 1 185

Holotype.—d, Boa Vista, Mu Castanbal, Para, Brazil 13-I-1965, W.
France coll.

Comments.—Perubahita confusa is related to P. longifal Linnavuori and
DeLong and can be separated by the small, rather slender, pointed, curved
aedeagus.

Angubahita DeLong, New Genus

Description.—Head as wide as pronotum, crown produced, rounded api-
cally, almost as long at middle as wide at base between eyes, % longer at
middle than length at eyes, 34 as long as pronotum. Crown angled with face,
margin, thick. Forewings parallel sided, longer than abdomen, with 3 closed
apical cells. Crown, pronotum, and scutellum marked with bright red spots.
Style triangular, aedeagus bifid apically. Species small in size, male 4 mm
in length and slender.

Type-species.—Angubahita arta DeLong, new species.

A key to the known genera of the Bahita group was published by Lin-
navuori and DeLong (1978a). The genus Angubahita would key out to cou-
plet 6, Taperinha. The species of Taperinha are larger, more robust than
Angubahita with a more broadly angled crown. The eyes are proportion-
ately larger in Angubahita, occupying about 2 of the dorsal surface of the
head, while in species of Taperinha the eyes occupy about % of the head
area, and have a crown which is 1% to twice as wide at base as long at
middle.

Angubahita arta DeLong, New Species
Figs. 7-12

Description.—Length of male 4 mm, female unknown. Crown roundly
produced, almost as long at middle as wide between eyes at base. Ocelli
large. Crown whitish, a black transverse stripe between ocelli just above
marginal white band. The black stripe surrounding ocelli, narrowed at mid-
dle and thickened on basal side, each side of middle. A black triangular
area, with pointed apex basad, on middle of crown. A red area at each side
of apex of black triangle and a black area along basal portion of each eye.
Pronotum with a reddish area at each side of apical portion, and with 4
black spots, 2 behind each eye at base. Scutellum grayish brown, basal
angles mostly reddish. Forewing grayish subhyaline, veins brown, color
intensified on costal veinlets, claval veinlets at commissure, and at base of
apical cells.

Male genital plates 5x as long as wide at middle, apices narrowed, sharply
pointed. Style triangular, longer than wide, apical portion straight, narrow,
finger-like, apex rounded. Aedeagus narrow, except near base, curved, api-
cal % divided into 2 narrow divergent processes. Pygofer narrowed, rounded
apically.

186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

aiid
IG
Ga

Figs. 1-17. 1-6, Perubahita confusa. 7-12, Angubahita arta. 13-17, Megabahita patula.
1, 7, Head, pronotum, scutellum, dorsally. 2, 12, 17, Pygofer laterally; apical portion in Fig.

2. 3, 8, 13, Aedeagus ventrally. 4, 9, 14, Aedeagus laterally. 5, 10, 15, Style laterally. 6, 11, 16,
Plate ventrally.

VOLUME 84, NUMBER 1 187

Holotype.—d, Las Cumbres, Panama, at light, 23-VI-1975, H. Wolda
coll.

Megabahita patula DeLong, New Species
Figs. 13-17

Description.—Length of male 5.5 mm, female unknown. Crown 3/s as
long at middle as wide at base between eyes. Crown with apical “4 white,
with a black spot at each side of apex. Remainder of crown reddish brown
at each side of a median grayish area. Pronotum dark gray, anterior 4
brownish. Scutellum grayish, apical ’% white. Forewing grayish yellow, cos-
tal veinlets broadly brown, veins brown.

Male genital plates 2x as long as wide at middle, apex rather broad,
rounded. Style with apophysis narrow, curved dorsally. Aedeagus broad at
middle (in lateral view) narrow at base and with a slender apex bearing 2
short apical processes. Pygofer bearing a base-apical process, enlarged, and
bearing a ventrally angled tooth near base, extending dorsally beyond mar-
gin of pygofer, apical portion slender, tapered, sharp pointed.

Holotype.—<d, Panama, El Llano, Carte Rd., 8-X-1974, H. Wolda coll.

Comments.—Megabahita patula is related to M. irroratus (Osborn) and
can be separated by the more broadened (lateral view) and the more basally
broadened pygofer spine.

Parabahita lamina DeLong, New Species
Figs. 18-23

Description.—Length of male 5.5 mm, female unknown. Crown short,
broadly rounded, appearing parallel margined. Pale brownish, crown with
a marginal, narrow, black, transverse band. A straighter, narrow, transverse
band between ocelli, and portions of a basal, wider, broken, black band
next to eyes. Pronotum rather dark brownish with blackened areas. Scu-
tellum pale brown with darker brown angles. Forewing dull yellowish, sub-
hyaline, veins pale to dark brown, some broadly embrowned.

Male genital plates 22x as long as wide at middle, apices rounded. Style
triangular, with the basal portion broad, apical portion thumblike, apex nar-
row, rounded. Aedeagus curved, sickle-shaped bearing a nonsclerotized,
saucer-shaped disc attached to the basai portion, extending across opening
between base and pointed apex. Apical portion of aedeagus divided, apices
pointed. Connective with basal, divided portion twisted at middle. Pygofer
bearing a long, slender, tapered, apical process which arises basocaudally
and extends dorsally beyond dorsal portion of pygofer.

Holotype.—¢, Santa Cruz, Bolivia, Exper. Sta. Saavedra, 350 m, 12-IV-
1979, 131Bb, D. Foster, V. Gongalez, and I. Caballero colls., at trap light.
Paratype male same data as holotype.

Comments.—Parabahita lamina is related to P. vesenyii Linnavuori and

188 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

can be separated by the slender, smooth tip of the apical portion of the
pygofer appendage, the rounded, nonsclerotized vertical disc at the base of
the aedeagus, and the twisted, basal portion of the connective.

Parabahita thalla DeLong, New Species
Figs. 24-28

Description.—Length of male 6 mm, female unknown. Crown short,
broadly rounded, appearing parallel margined. Crown pale brown with a
curved. transverse, dark brown, narrow band, broadened at middle, just
above margin. A very narrow, straight, black, transverse band between
ocelli. A broader brown band between eyes, near base, slightly broken at
middle. Pronotum pale brown on anterior %, posterior %4 dark brown. Scu-
tellum with dark brown basal angles surrounded by white. Median longi-
tudinal portion pale brown.

Male genital plates 342x as long as wide at middle, apices narrowed,
bluntly pointed. Style elongate, narrow, apical 2 long, narrow, slightly
curved, apex rounded. Aedeagus sickle-shaped, apical portion divided into
2 pointed processes. Pygofer bearing a long curved apical process, the apical
'4 of which is broadened and pointed, spear-shaped.

Holotype.—¢, Santa Cruz, Bolivia, 2-I[V-1979, 350 m, D. Foster, V. Gon-
zalez, and I. Caballero colls, at trap light. Paratype male same data as
holotype.

Comments.—Parabahita thalla is related to P. vezenyii Linnavuori and
can be separated by the long slender apophysis of the style and the broad-
ened spearlike apical portion of the pygofer apical process.

Frequenamia particula DeLong, New Species
Figs. 29-33

Description.—Length of male 5.5 mm, female unknown. Crown produced
and rounded, 3 as wide at base between eyes as long at middle. Crown
white, a brown marginal line. A dark brown narrow transverse band just
distal to a narrow whitish band between the 2 brownish bands. Basal 2 of
crown, pale brown. Pronotum dark brown with paler brownish mottling.
Scutellum with basal 24 brown, apical % white. Forewing pale brownish
subhyaline, with numerous white areolar spots, 5 prominent spots on clavus,
veins mostly pale brown.

Male genital plates more than 3x as long as wide at middle, apex nar-
rowed, bluntly pointed. Style with apophyses narrow, rounded. Aedeagus
rather broad at base, narrowed to a curved, narrow, bluntly pointed apex,
with 2 straight, long, slender processes attached laterally, which extend

caudally beyond curved apex of aedeagus. Pygofer narrowed and rounded
apically.

VOLUME 84, NUMBER 1 189

22
DO 20 21
18
19
| = /| NS
24 26
27

AIR
Wa OA

Figs. 18-38. 18-23, Parabahita lamina. 24-28, P. thalla. 29-33, Frequenamia particula.

34-38, F. atrata. 18, 24, 30, 34, Aedeagus ventrally. 19, 25, 29, 35, Aedeagus laterally. 20,
Connective ventrally. 21, 26, 31, 36, Style laterally. 22, 33, 38, Pygofer laterally, apical portion.

28, Pygofer laterally. 23, 27, 32, 37, Plate ventrally.

Holotype.—¢ , Panama, Chiriqui, Fortuna, 1050 m, 26-IX-1976, H. Wolda

coll.
Comments.—Frequenamia particula is related to F. crassistylus Linna-
vuori and can be separated by the two long slender processes attached to

the aedeagus extending caudally beyond the apex of aedeagus.

190 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Frequenamia atrata DeLong, New Species
Figs. 34-38

Description.—Length of male 4 mm, female unknown. Crown broadly
rounded, twice as wide at base between eyes as long at middle. Crown
declivated, margin thick, rounded to front. Anterior ¥% of crown with deep
lateral striae. Crown brownish, apical 2 black, basal % brownish. Pronotum
black with brownish, widened punctate spots. Scutellum black with pale
brownish spots on central portion. Forewing appearing black, a few brown-
ish areas on corium, apical, and anteapical cells.

Male genital plates more than twice as long as wide at middle, apex nar-
rowed, bluntly rounded, style with apophyses narrow, slightly curved, ex-
tending almost caudally. Aedeagus rather narrow, curved, tapered from
base to slender apex, divided apically into 2 divergent processes. Pygofer
narrowed apically, rounded.

Holotype.—¢, Panama, Barro Colorado, at light, 22-IX-1975, H. Wolda
coll.

Comments.—Frequenamia atrata is related to F. lacerae (Signoret) and
can be separated by the unbranched apical process of the aedeagus.

LITERATURE CITED

DeLong, D. M. 1947. A new genus (Frequenamia) and species of Mexican leafhoppers related
to Mesamia (Homoptera: Cicadellidae). Bull. Brooklyn Entomol. Soc. 42: 63-64.
Linnavuori, R. 1959. Revision of the Neotropical Deltocephalinae and some related subfam-

ilies (Homoptera: Cicadellidae). Ann. Zool. Soc. Vanamo 20: 1-370.
Linnavuori, R. and D. M. DeLong. 1978a. Neotropical leafhoppers of the ‘‘Bahita’’ group
(Homoptera: Cicadellidae: Deltocephalinae), a contribution to the taxonomy. Brenesia
14/15: 109-169.
. 1978b. Some new or little known Neotropical Deltocephalinae (Homoptera: Cicadel-
lidae). Brenesia 14/15: 227-247.
Oman, P. H. 1936. A generic revision of American Bythoscopinae and South American Jas-
sinae. Univ. Kans. Sci. Bull. 24: 343-429.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 191-203

REPRODUCTIVE AND CALLING BEHAVIOR IN TWO CLOSELY
RELATED, SYMPATRIC LACEWING SPECIES,
CHRYSOPA OCULATA AND C. CHI
(NEUROPTERA: CHRYSOPIDAE)

CHARLES S. HENRY

Biological Sciences Group, Box U-43, University of Connecticut, Storrs,
Connecticut 06268.

Abstract.—The sister species Chrysopa oculata Say and C. chi Fitch,
like other green lacewings of eastern North America that have been studied,
jerk their abdomens vigorously while courting and mating. The substrate-
borne signals (calls) produced principally by male insects are simple and
repetitive, consisting of more or less extended trains of discrete volleys or
bursts of abdominal vibration. The calls of both species are similar but
distinctive, differing significantly and consistently from one another in their
volley spacing, volley duration, and frequency of abdominal vibration. Some
inter- and intra-individual variation in call parameters exists within each
species; for example, vibration frequency is particularly well correlated with
ambient temperature, at least in C. oculata. Acoustical communication in
these two Chrysopa s. str. species seems much less crucial to successful
courtship than it is in Chrysoperla species, indicating that calling patterns
probably do not serve a reproductive isolating function within species of the
oculata group. Alternative functions for jerking behavior are therefore con-
sidered, and an attempt is made to relate acoustical performance with other
aspects of lacewing reproductive biology and ecology.

A special form of acoustical communication, in which courting conspe-
cific individuals exchange substrate-borne low-frequency signals, has been
shown to be widespread in North American green lacewings of the family
Chrysopidae (Henry, 1979, 1980a, b, c; Smith, 1922; Toschi, 1965; Tauber,
1969). Transverse ‘‘sound’’ waves are generated in lightweight substrates
by vigorous jerking motions of a lacewing’s abdomen and are thereby trans-
mitted over short distances to other insects standing on those surfaces (Hen-
ry, 1980c). Calls consist of bursts or volleys (‘pulse trains’’—see Graf,
1970) of such abdominal jerking or vibration, usually modulated with respect
to frequency and amplitude structure and released over a period of time at

192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

species-characteristic intervals. Calling duets are usually established be-
tween two sexually receptive conspecifics of opposite sex, and often cop-
ulation will not occur without a period of such reciprocal signaling.

Chrysopa (broad sense) in North America includes at least three distinct
species-groups, called subgenera by Tjeder (1966) but recognized as full
genera by Séméria (1977). These, commonly known in the United States as
the oculata group, carnea group, and lineaticornis group, are defined by
features of the male terminalia (Bram and Bickley, 1963), but possess dis-
tinctive and consistent bionomic characteristics as well. For example, mem-
bers of the oculata group (Chrysopa s. str. S¢méria) produce strong-smell-
ing ‘‘repugnatorial’’ secretions, are predatory as adults, and overwinter as
prepupae within the cocoon, while those of the carnea group (Chrysoperla
Steinmann) are neither odoriferous nor obligatorily carnivorous as adults
and pass the winter as diapausing, reproductively immature imagos (Tauber
and Tauber, 1974; Séméria, 1977). This strong dichotomy between Chry-
sopa s. str. and Chrysoperla is substantiated by calling behavior patterns:
members of the latter taxon will not mate without first participating in pro-
longed, often elaborate calling duets, while those of the oculata group that
I have studied call in a simpler, more perfunctory, and less predictable
manner and will occasionally copulate without acoustical preliminaries. In
this paper, I describe abdominal jerking behavior and courtship and mating
patterns in two closely related species of Chrysopa s. str., C. oculata Say
and C. chi Fitch, confirming in this case that members of species-pairs not
dependent for mating success on acoustical communication can be more
similar in their calls than those for which such communication is a necessary
adjunct to copulation. That prediction is derived from the hypothesis that
calling in lacewings functions at least in part, or once functioned, to isolate
closely related species reproductively from one another; in two close rela-
tives which utilize acoustical signals extensively, selection against hybrids
would likely cause rapid divergence in call characteristics, as seen in Chry-
soperla species.

Chrysopa oculata and C. chi are common Nearctic green lacewings of
medium size. Chrysopa oculata is found throughout most of North America
north of Mexico and is the most abundant chrysopid species at most sites
(Smith, 1922; Bickley and MacLeod, 1956; Bram and Bickley, 1963). It is
a meadow species, confined to low vegetation, and passes through three
generations each summer in New England (Propp et al., 1969). Chrysopa
chi is thought, on morphological grounds, to be C. oculata’s closest relative
(Bram and Bickley, 1963), and similarly occupies a large but rather more
northerly range in the United States, Canada, and Alaska. It is described
as an “‘early’’ species (Smith, 1922), inhabiting the taller shrubs, like mead-
owsweet (Spiraea latifolia), of overgrown fields; little is known of its life
history and habits beyond the brief comments by Smith (1922) on hatching,

VOLUME 84, NUMBER 1 193

larval development, and eclosion. The reproductive behavior of neither
species has been described in detail, although Smith (1922) observed ab-
dominal jerking, courtship, and copulation in C. oculata on two separate
occasions.

METHODS AND MATERIALS

Specimens of the two species were collected during the summers of 1977
through part of 1981 from Mansfield, Connecticut; Rensselaerville, New
York; Echo Lake, Vermont; and North Conway, New Hampshire. Breeding
colonies in 2-gallon cardboard cartons were provided with water, artificial
WHEAST™ diet (Hagen et al., 1970), and black aphids from Nasturtium.
Eggs were removed daily and larvae reared on ether-killed Drosophila spp.,
as described in an earlier paper (Henry, 1979). All adults and larvae were
maintained under long day (16L:8D) light conditions at 28 + 1°C and 40-
80% relative humidity. Populations from the different collecting locales were
maintained separately only until calling patterns were shown to be identical
in lacewings from all four areas. First through third laboratory generations
were tested for reproductive and acoustical behavior.

Since lacewing signals are of low frequency and are transmitted through
the substrate, a simple piezoelectric transducer contacting the thin plastic
cover of the experimental arena sufficed as a detector of insect-induced
substrate vibrations, as in earlier experiments (Henry, 1979, 1980a, b, c).
The electrical output of that transducer was then fed to a Tektronix™ S511
storage oscilloscope through a matching 5A25 dual differential amplifier;
tracings of lacewing calls could then be viewed and photographed for future
analysis. Both Chrysopa oculata and C. chi displayed greater sexual activ-
ity under red rather than fluorescent illumination; a GE™ 60-watt BCJ in-
candescent bulb was used for that purpose.

The results and conclusions that follow are based upon 193 hours of ob-
servations, approximately equally divided between C. oculata and C. chi.
For C. oculata, 197 calls consisting of three or more volleys were analyzed;
these represent the recorded activities of 11 different males and 4 females,
including two heterosexual duets involving 4 different individuals. Six cop-
ulations (12 individuals) were also observed in that species. For C. chi, I
recorded and analyzed 158 calls of three or more volleys, representing the
activities of 8 males and one female. Additionally, one heterosexual duet
produced 7 male and 11 female recordable responses (calls). Nine C. chi
copulations (7 different males, 8 females) were tabulated. During observa-
tion sessions of C. oculata and C. chi, temperature averaged 26.46 + 1.62°C
(n = 53 sessions) and 25.94 + 3.01°C (n = 36), respectively.

Any reference in the Results or Discussion sections to “‘significant dif-
ferences’’ indicates that the means of two normally distributed samples were
demonstrated to differ from one another by a 2-tailed t-test using confidence

194 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

limits of 95% or better. Similar confidence limits were used in chi-square
evaluations. Values following a + sign are one standard deviation of the
mean.

RESULTS

My collecting records confirm the ecological observations of earlier work-
ers (Smith, 1922). In three successive years at the New York State site,
adult C. chi appeared at least as early in the spring as C. oculata, and in
one year (1980) were abundant on the 29th of May when no C. oculata
could be found. At all locales, populations of both species were emphatically
sympatric throughout the summer; moving one or two meters from a mead-
ow dominated by Solidago spp. to an open aspen grove with a thick Spiraea
understory resulted in complete replacement of C. oculata by C. chi in the
collecting samples. Adults of both species disappeared from all my sites by
15 September, with C. oculata persisting consistently longer (by a week or
two) than its relative in each of three consecutive years.

Reproductive behavior is basically similar in both Chrysopa oculata and
C. chi, although it differs significantly from that described for members of
the Chrysoperla group. Sexual receptivity in both sexes 1s first signaled by
lowered head and elevated abdomen, accompanied by vigorous antennal
movements. The abdomen is then waved slowly back and forth several (up
to 12-15) times at 1.5 second intervals and is flexed ventrally at its tip as it
approaches the left and right limits of its excursion. The wings on that side
of the body toward which the abdomen is swinging are partially raised and
lightly fluttered; simultaneously, one can detect the characteristic, unpleas-
ant odor of species in this taxon. The male may then produce a series of
volleys of abdominal vibration (Fig. 1A—D), either continuously for a minute
or more (C. oculata and highly receptive C. chi) or more sporadically (less
receptive C. chi). Females may also jerk their abdomens in a manner quite
similar to that of the males; however, female activity of this kind is rare in
either species. In the unlikely event of a heterosexual duet, the male and
female will reciprocally exchange bouts of abdominal jerking, exchanging
volley for volley in C. chi but alternating longer trains (series) of volleys in
C. oculata (Fig. 1E, G). More commonly, however, courtship will be brief
and copulation sudden, accompanied by little if any jerking activity: typi-
cally the female will approach and antennate the male, after which the two
insects will immediately swing their bodies into parallel positions and effect
copulation. Individuals of each species averaged 30.60 + 8.02 (C. oculata;
n = 5) and 14.00 + 4.76 (C. chi; n = 7) minutes in copulo (t = 2.62). In C.
oculata, three out of six copulations were preceded by at least one series

of volleys from the female, while in C. chi three of nine included such
participation.

VOLUME 84, NUMBER | 195

Fig. |. Oscillographs of abdominal vibration patterns produced by sexually receptive in-
dividuals of Chrysopa oculata (A, B, E) and C. chi (C, D, G). Heterosexual duets are recorded
in E and G at an oscilloscope writing speed of 2 seconds/major division; in C. oculata, whole
trains of volleys of abdominal jerking are exchanged between partners, while in C. chi each
insect exchanges single volleys with its partner. Note striking convergence between the duet
sequence of C. chi (G) and that C. (Chrysoperla) carnea (F), recorded at the same writing
speed. Traces B and D are 20x details of A and C (writing speed = 0.10 sec/div. vs. 2 sec
div.), showing volley structure characteristic of each species.

Calls of both Chrysopa oculata and C. chi are simpler in structure than
those of any Chrysoperla species yet studied, consisting of long trains of
identical volleys of abdominal vibration lacking pronounced frequency or
amplitude modulation (Fig. 1). Differences between the calls of the two
species include (a) average volley spacing, (b) frequency of abdominal vi-
bration within volleys, (c) volley duration, and (d, calculated from b and c)
number of abdominal jerks per volley (Table 1). In C. oculata (Fig. 1A, B),
volleys are spaced closely together (interval = 0.266 sec) and are charac-
terized by high rates of abdominal jerking (av. = 109.18 abdominal strokes
sec), short durations (av. = 0.082 sec), and relatively few total strokes
(jerks) per volley (av. = 8.95). On the other hand, C. chi produces volleys
(Fig. 1C, D) that are more distantly spaced (interval = 0.813 sec), lower in
vibration frequency (76.11 strokes/sec), longer by a factor of two (0.168
sec), and composed of considerably more abdominal strokes (12.79) than

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Jaye syuasaidai sed yous Jo oinsy 19d1e] oy} {A[feoNeyUosed poidjud oie sazis s[dweg “payeynqe) ov suOleIAap PAvpUe}s ITOY} PUB SoN[eA URI]
“1y9 “2 pure vinjns0 vdosduy+D Jo sayeway pue soyeul Jo (S9duaNbas 10 sures] AQTJOA) S][Bd By) JO SINSL9}JIeAeYS a[GvoinsPow juewiodwy “| Iqey

VOLUME 84, NUMBER 1 197

those of C. oculata. These differences are all significant to a high degree,
even when data are averaged for each individual insect and sample sizes are
accordingly reduced from total number of calls to number of different in-
dividuals (not tabulated). Both species, but particularly C. chi, display slight
but detectable frequency modulation of each volley: the rate of abdominal
jerking decreases somewhat toward the end of each volley (Fig. 1B, D).

Variation in call parameters exists both among individuals and within a
single individual of C. oculata and C. chi. Inter-individual variation was
not usually very pronounced in either species; however, one C. oculata
displayed consistently and significantly longer intervals between its volleys
than any other conspecific individual (other features of its call were less
deviant). Temperature was the major determinant of intra-individual varia-
tion: decreasing temperatures tended in both species to increase volley spac-
ing but lower the frequency of abdominal vibration, the latter in a highly
significant linear manner for C. oculata (Fig. 2; slope = 5.87, coefficient of
determination = 0.94, n = 6). A third type of variation, existing within a
call, was also identified. Here, it was noticed in both species that volley
spacing nearly always increased significantly during the course of an ex-
tended call. Table 2 documents this observation, for each individual studied,
and includes the highly significant results of a chi-square analysis; the trend
is also reflected in the average volley spacing of ‘‘early’’ vs. “‘late’’ portions
of calls listed in Table 1. It should be mentioned that, despite such multi-
faceted variation, no significant interspecific overlap existed in any call pa-
rameter between C. oculata and C. chi even when calls of the most extreme
(and therefore most similar) individuals, under unrealistically different tem-
perature conditions, were compared with each other.

As mentioned earlier, C. oculata and C. chi differ from one another in
their manner of alternating acoustical signals during heterosexual duets (Fig.
1E, G). The longer spacing between volleys in C. chi seems consistent with
the interdigitated volleys typical of dueting insects of that species, while C.
oculata individuals produce volleys so rapidly that exchanging volley for
volley is probably not feasible. In C. oculata, calls produced by an individ-
ual during a duet appear to be physically indistinguishable from those pro-
duced by that same individual alone. However, individuals of C. chi make
their already-large solo volley spacing significantly greater when dueting, as
if to make it even easier for their partners to synchronize with them: the
best-studied male, CHM-41, increased its between-volley interval from
0.873 + 0.251 sec (n = 16) to 1.376 + 0.507 sec (n = 7; same temperature)
when alternating volleys with the female CHF-41 (t = 3.23). It should be
stressed again, though, that acoustical alternation during heterosexual in-
teractions is rarely seen in either species.

Male-male interactions in C. oculata and C. chi are infrequent and in-
volved the exchange of acoustical signals on only one occasion (in C. chi).

198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

31
OM-—1* (43)
3 NOAH HID I ESERPOOPOD Ee
=
OM-— 1* (20)
2 a x
‘OM—5 (15)

o MIO HONE
' 28
Ww
S
en
.-¢
a
2 26
= a (44)
= cae a non

N
ul

L~)
>

115 110 105 100
FREQUENCY — ABD. STROKES/SEC

Fig. 2. Graph showing the effect of temperature on abdominal vibration frequency in Chry-
sopa oculata, for four different individuals. Solid dots are means, shaded areas are standard
deviations, and lines extend to extremes. In each case, sample size, in parentheses, is the total
number of volleys measured. The sloping line corresponds to the linear equation y = 5.87x
— 48.77, with vibration frequency plotted on the y-axis (turn graph clockwise 90 degrees).

No special patterns could thus be discerned. My impression is that males
do not interfere acoustically with the reproductive activities of other males,
since on 30 out of 34 occasions peripheral males showed no interest in
courting and copulating conspecifics in the cup-sized experimental arenas.
However, when crowded together for long periods of time, males undoubt-
edly interact aggressively, since nearly every individual of both species
exhibited antennal injury under those conditions.

DISCUSSION

Collecting data suggest that little interaction will occur between Chrysopa
oculata and C. chi under natural conditions, since each species occupies a
different microhabitat. For most of the summer, C. oculata seems to be the
only lacewing species inhabiting the low vegetation at most meadow sites
in the Northeast; only C. (Chrysoperla) carnea Stephens is likely to co-
occur with it (Smith, 1922; Bickley and MacLeod, 1956), but in practice I
have never collected those two species together. On the other hand, C.
chi’s ecological preferences overlap with those of several other lacewings,
including C. nigricornis Burmeister, C. quadripunctata Burmeister, C.
(Chrysoperla) rufilabris Burmeister, C. (2) lineaticornis, and even the co-
nifer-associated C. (Chrysoperla) harrisii Fitch and Meleoma Fitch (two

199

VOLUME 84, NUMBER |

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ay) ‘[eNplAIpul yove Joy “SUrATeYy Aq payepnoyeo sem onyeA poysedx9 ay} ‘S}S9} auenbs-1yd oy} 104 “WUROYyIUsISUT pospnf oom 1Yy9 “2D Ul puodas F0'0)
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pipjnz0 vdoskayD JO [RNPIAIPUL YORI 1OJ payeynge) ‘surseds Aa|JOA ul JURJSUOS POUILUIAL 10 ‘pasevaidep “Paseasoul Je} s]jeo jo toquinN = “7 PGP.

200 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

species) (Balduf, 1939; Bickley and MacLeod, 1956); in fact, all of those
species have been found associated with C. chi at one time or another at
the New York State site alone. Perhaps this abundance of competitors par-
tially explains why populations of C. chi always remain smaller and more
patchy than those of its closest relative. Of course, factors underlying
species composition are completely unknown in this case.

As mentioned, calls are much simpler and less consistently delivered in
C. oculata and C. chi than they are in Chrysoperla species. Of the call
types so far described, their calls most closely resemble that of C. (Chry-
soperla) carnea, which also produces trains of short, identical volleys dur-
ing courtship and mating (Henry, 1979, 1980c). In fact, volley spacing is
virtually the same in C. (C.) carnea and C. chi, and both species alternate
single volleys rather than whole trains of volleys during heterosexual duets
(Henry, 1979), so that dramatic convergence is readily apparent in oscillo-
graphs of their calls (Fig. 1F, G). However, closer inspection of volley
structure reveals that C. carnea’s volleys are significantly longer and char-
acterized by much more pronounced frequency modulation than those of
either C. chi or C. oculata (see Fig. 1B in Henry, 1980c); additionally,
neither of the latter two species depends for its mating success upon acous-
tical interactions between the sexes. At any rate, convergence in calls be-
tween two such distantly related species as C. chi and C. carnea has little
biological meaning even if calls are important premating barriers, since dif-
ferences in the habitats, food preferences, pheromone repertoires, and be-
havioral responses of the two species preclude such close-range interac-
tions.

In the basic components of their reproductive and calling behavior, C.
oculata and C. chi are not only similar to each other, but also resemble
other members of their species-group, including C. nigricornis (Toschi,
1965, and personal observation) and C. quadripunctata (unpublished data).
All species of Chrysopa s. str. that have been studied assume a character-
istic ‘head down”’ position when sexually receptive, wave their abdomens
in the manner described earlier, and produce superficially similar, simple
volleys of abdominal vibration that differ primarily in repetition rate (tem-
poral patterning). Additionally, copulation is usually swift and preceded by
the release of a strong-smelling secretion, presumably from the prothoracic
‘“‘repugnatorial glands’’ of one or both sexes (McDonnough, 1909; Sulc,
1914; Withycombe, 1924). As mentioned earlier, Chrysoperla species never
produce noticeable odors, although the prothoracic glands seem to be as
fully developed in them as in members of Chrysopa s. str. (E. G. MacLeod,
pers. comm., 1967). It is thus tempting to correlate the production of odors
during copulation with relatively low acoustical sophistication, speculating
that more complex, intense pheromones have replaced acoustical modes of
communication as species-isolating mechanisms in Chrysopa s. str. Selec-

VOLUME 84, NUMBER 1 201

tive pressures favoring radically different calling patterns in closely related
sympatric species, as seen in Chrysoperla, are thus weakened in Chrysopa
s. str., leading incidentally to the relatively weak distinctions described here
between the calls of C. oculata and C. chi. That complex jerking behavior
is the primitive, rather than derived, condition in green lacewings is an
assumption that needs more support: outgroup comparison can tell us that
some Hemerobiidae, the sister-group to Chrysopidae, also jerk their abdo-
mens (Smith, 1922), but those calls have never been studied. Similarly, it
is not necessarily valid that lacewing odors can function as reproductive
pheromones, since their repugnatorial function has always been taken for
granted and has even been confirmed by Blum et al. (1973), who isolated
the active ingredient skatole from the prothoracic exudate of C. oculata and
showed that it repelled fire ants (Solenopsis richteri Forel) and some albino
mice. Nevertheless, skatole might both repel enemies and attract conspe-
cifics in C. oculata, and slightly different substances could function as sex-
ual attractants in other members of Chrysopa s. str.; Blum et al. (1973) did
not analyze the secretions of any other species. Even if skatole is present
in other ‘“‘stinkflies,’ it may be mixed with other, rarer active ingredients
(Blum et al.’s 5% unidentified ‘‘volatiles’’) in unique, species-specific pro-
portions for sexual attraction, as are other blends of pheromones in certain
groups of Lepidoptera (Roelofs and Cardé, 1974; Silberglied, 1977). Cer-
tainly the release during lacewing courtship and mating of secretions pre-
sumed to be defensive is provocative and suggests the need for further
research.

I feel that to reject the hypothesis that calls serve a species-isolating
function in Chrysopa s. str. is premature, especially in view of the mea-
sureable and consistent differences detected between the calls of C. oculata
and C. chi. However, alternative functions for acoustical behavior in these
insects must also be considered, since calling differences do seem weak
among species of this group. For example, jerking behavior may additionally
serve to induce sexual activity in a partner, to facilitate dispersal of some
other pheromone from an exocrine gland in the male genitalia (MacLeod,
pers. comm. 1967; personal observation), or even to interfere with the mat-
ing success of other males in competition for females (see Cade, 1980, for
a review of competition among singing crickets). Short-distance attraction
of females to calling males may also be important in nature, occurring after
aggregation of conspecific individuals to a site by pheromones, local prey
abundance (Hagen et al., 1976), or plant odors (Flint et al., 1979). Several
of the above alternatives predict intensified sexual selection in males of
Chrysopa s. str., especially in view of the polygamous or even polygynous
(unpublished data) mating systems characteristic of species of that taxon
(see Halliday, 1978, for a general discussion of sexual selection). In fact, as
predicted, males of both C. oculata and C. chi do most of the calling and

202 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

display considerably more aggression toward other males than do males of
any Chrysoperla species yet studied. However, as far as I can tell, male-
male competition has not generated in males of either species studied here
the kinds of aggressive calls and chorusing interactions found in some other
polygynous singing insects and amphibians (Alexander, 1975; Wells, 1977).

The physiological basis of jerking behavior in green lacewings is not
known. Earlier work (Henry, 1980b, c) demonstrated that conspecific in-
dividuals of different size (mass) do not differ significantly from one another
in abdominal vibration frequency, suggesting that abdominal motion is con-
trolled by a neural oscillator rather than by the resonant properties of the
moving mass. That view is substantiated by the pronounced temperature
effects described in this study (Fig. 2), since resonant properties should not
change much, if at all, with temperature. In fact, the linear relationship
observed in C. oculata is consistent with the more complex exponential and
logistic curves describing various inorganic and organic chemical reactions
and biological processes (Wigglesworth, 1965), since most of those curves
approximate linear equations over a narrow range of temperatures. Here,
the temperature coefficient Q,) for vibration frequency in C. oculata is 1.86,
meaning that for temperatures from 24.5—30°C, frequency will increase by
a factor of 1.86 for each 10°C increment. If frequency differences are at all
important to lacewings in the recognition of conspecifics, for whatever fun-
damental purpose, neural control provides the best means of achieving con-
sistency among individuals of different sex and sizes under a given set of
environmental conditions.

ACKNOWLEDGMENTS

I thank Steve M. Cohen, Jane E. O’Donnell, Ellen B. Dawley, and es-
pecially Raymond J. Pupedis (University of Connecticut) for help in col-
lecting and maintaining living lacewings. Robert Dahgleish kindly permitted
us to collect on the grounds of the E. N. Huyck Preserve, Rensselaerville,
New York. The work was supported by National Science Foundation award
#DEB-79-11537, Charles S. Henry, principal investigator.

LITERATURE CITED

Alexander, R. D. 1975. Natural selection and specialized chorusing behavior in acoustical
insects, pp. 35-77. Jn Pimentel, D., ed., Insects, Science, and Society. Academic Press,
N.Y. 284 pp.

Balduf, W. V. 1939. The Bionomics of Entomophagous Insects. Part II. John S. Swift Co.,
Inc., N.Y. 384 pp.

Bickley, W. E. and E. G. MacLeod. 1956. A synopsis of the Nearctic Chrysopidae with a
key to the genera (Neuroptera). Proc. Entomol. Soc. Wash. 58(4): 177-202.

Blum, M. S., J. B. Wallace, and H. M. Fales. 1973. Skatole and Tridecene: identification and
possible role in a chrysopid secretion. Insect Biochem. 3: 353-357.

Bram, R. A. and W. E. Bickley. 1963. The green lacewings of the genus Chrysopa in Maryland
(Neuroptera: Chrysopidae). Bull. Univ. Md. Agric. Exp. Stn. A-124: 1-18.

VOLUME 84, NUMBER 1 203

Cade, W. 1980. Alternative male reproductive strategies. Fla. Entomol. 63(1): 30-45.

Flint, H. M., S. S. Salter, and S. Walters. 1979. Caryophyllene: an attractant for the green
lacewing. Environ. Entomol. 8(6): 1123-1125.

Graf, R. G. F. (ed.). 1970. Modern Dictionary of Electronics (3rd ed.). Bobbs-Merrill Co..
Minneapolis, Minn. 408 pp.

Hagen, K. S., P. Greany, E. F. Sawall, Jr., and R. L. Tassan. 1976. Tryptophan in artificial
honeydews as a source of an attractant for adult Chrysopa carnea. Environ. Entomol.
5: 458-468.

Hagen, K. S., R. L. Tassan, and E. F. Sawall, Jr. 1970. Some ecophysiological relationships
between certain Chrysopa, honeydews, and yeasts. Portici Boll. Lab. Entomol. Agr.
Filippo Silvestri 28: 113-134.

Halliday, T. R. 1978. Sexual selection and mate choice, pp. 180-213. Jn Krebs, J. R. and N.
B. Davies, eds., Behavioral Ecology. Blackwell Sci. Publs., Oxford.

Henry, C. S. 1979. Acoustical communication during courtship and mating in the green
lacewing Chrysopa carnea (Neuroptera: Chrysopidae). Ann. Entomol. Soc. Am. 72(1):
68-79.

—. 1980a. Acoustical communication in Chrysopa rufilabris (Neuroptera: Chrysopidae),
a green lacewing with two distinct calls. Proc. Entomol. Soc. Wash. 82(1): 1-8.
—. 1980b. The courtship call of Chrysopa downesi Banks (Neuroptera: Chrysopidae): its

evolutionary significance. Psyche (Camb. Mass.) 86(2—3): 291-297.

—. 1980c. The importance of low-frequency, substrate-borne sounds in lacewing com-
munication (Neuroptera: Chrysopidae). Ann. Entomol. Soc. Am. 73(6): 617-621.
McDonnough, J. 1909. Uber den Bau des Darms und seiner Anhange von Chrysopa perla L.

Arch. Naturgesch. 75: 313-360.

Propp, G. D., M. J. Tauber, and C. A. Tauber. 1969. Diapause in the neuropteran Chrysopa
oculata. J. Ins. Physiol. 15: 1749-1757.

Roelofs, W. L. and R. T. Cardé. 1974. Sex pheromones in the reproductive isolation of
Lepidoptera species, pp. 96-114. Jn Birch, M. C., ed., Pheromones. Am. Elsevier Publ.

ConNeye
Sémeria, Y. 1977. Discussion de la validité taxonomique du sous-genre Chrysoperla Stein-
mann (Planipennia, Chrysopidae). Nouv. Rev. Entomol. 7(2): 235-238.

Silberglied, R. E. 1977. Communication in the Lepidoptera, pp. 362-402. In Sebeok, T. A.,
ed., How Animals Communicate. Indiana Univ. Press, Bloomington, Ind.
Smith, R. C. 1922. The biology of the Chrysopidae. Cornell Univ. Agric. Exp. Stn. Mem. 58:

1291-1372.
Sulc, K. 1914. Uber die Stinkdriisen und Speicheldriisen der Chrysopen. Stitzungsberichte
Kgl. Bohmischen Gesellschaft der Wissenschaften 11: 1—S0.

Tauber, C. A. 1969. Taxonomy and biology of the lacewing genus Meleoma (Neuroptera:
Chrysopidae). Univ. Calif. Publ. Entomol. 58: 1-62.

Tauber, M. J. and C. A. Tauber. 1974. Dietary influence on reproduction in both sexes of
five predacious species (Neuroptera). Can. Entomol. 106: 921-925.

Tjeder, B. 1966. Neuroptera-Planipennia. The lacewings of Southern Africa. 5. Family Chry-
sopidae. S. Afr. Anim. Life 12: 228-534.

Toschi, C. A. 1965. The taxonomy, life histories, and mating behavior of the green lacewings
of Strawberry Canyon. Hilgardia 36: 391-431.

Wells, K. D. 1977. The social behaviour of anuran amphibians. Anim. Behav. 25: 666-693.

Wigglesworth, V. B. 1965. The Principles of Insect Physiology (6th ed.). Methuen and Co.,
Ltd., London. 741 pp.

Withycombe, C. L. 1924. Further notes on the biology of some British Neuroptera. Ento-
mologist 57: 145-152.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 204-207

HEMEROBIUS STIGMA STEPHENS (NEUROPTERA: HEMEROBIIDAE):
EXTERNAL MORPHOLOGY OF THE EGG

GARY L. MILLER AND PARIS L. LAMBDIN

Department of Entomology and Plant Pathology, University of Tennes-
see, Knoxville, Tennessee 37901.

Abstract.—Adult brown lacewings, Hemerobius stigma Stephens, col-
lected from white pine were maintained under laboratory conditions at
18.3°C + 1.0°C and 6L:18D. Females laid 133 (72-176) eggs. Each egg was
yellow, 730 wm (675-750) long, 335 wm (300-375) wide, and weighed 0.046
mg. The eggs possessed reticulations arranged in a diagonal pattern on a
smooth chorion. The micropylar process consisted of a series of waxlike
projections arranged in groups on the anterior pole.

Smith (1923) and Withycombe (1922) provided the first life history inves-
tigations on Hemerobius stigma Stephens (=H. stigmaterus Fitch), includ-
ing a brief description of the egg. Advances in microscopy have provided
the opportunity to examine in greater detail the external morphology of the
egg. Mazzini (1976) noted that ultrastructural morphology of the egg chorion
and micropyle has provided information useful for inferring phylogenetic
relationships for insects in several orders.

Identification of hemerobiids is presently based on morphological char-
acteristics of the adults (MacLeod and Stange, 1981). Our objective was to
describe the external morphology of the egg of H. stigma. Additional in-
vestigations on eggs of related species may assist in identifying other hem-
erobiid species.

MATERIALS AND METHODS

A laboratory colony of H. stigma was established 3 March 1981 at The
University of Tennessee from adults collected on white pine, Pinus strobus
L. The colony was maintained at 18.3°C + 1.0°C on an 8L:16D cycle.

Male-female pairs were placed in 3.5 x 9.5 cm petri dishes and allowed
to mate. The bottom of each petri dish was lined with a 9.0 cm disk of
Fisher brand coarse filter paper. A 10 dr vial filled with distilled water and
plugged with cotton was placed in each petri dish to provide water and

VOLUME 84, NUMBER 1 205

Fig. 1. a, Egg of Hemerobius stigma (116); b, Surface of anterior pole showing waxlike
projections and micropylar process (144); c, Tubercles and fissures on the micropylar surface
(1200x).

206 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

moisture. A cotton ball, ca. 1.5 cm diam., and pine needles were placed in
the petri dish to serve as a substrate for oviposition. Females were main-
tained until death to determine fecundity and ovipositional habits.

Eggs were collected within 24 h after oviposition and weighed using an
Ainsworth SCV electrobalance. Egg size was determined using an ocular
micrometer. Egg color was compared with color charts in the Munsell Book
of Color (1929), and the color description was recorded (e.g., yellowish red
in Munsell might be written as 5 YR 6/8 or a No. 5 yellow red hue, color
value No. 6 and a chroma No. 8).

External morphology was examined using an ETEC Autoscan scanning
electron microscope. Eggs were placed on aluminum studs and coated with
gold-palladium in a Denton Vacuum DV-515 vacuum evaporator. S.E.M.
photographs were taken using Polaroid 55 film, and photomicrographs were
taken using 35 mm Kodak Panatomic X film.

RESULTS

Field-collected adult females of H. stigma maintained in the laboratory
laid an average of 133 (72-176) eggs. Eggs were deposited singly or in groups
of 2-5 on filter paper, cotton balls, pine needles, and in the scaly sheath at
the base of the needles. Withycombe (1922) reported that females occasion-
ally laid up to seven eggs in groups.

Eggs were ellipsoid (Fig. 1a), 730 wm (675-750) long, 335 wm (300-375)
wide, and weighed 0.046 mg. Color of eggs 24-72 h after deposition was
yellow (SY 8/6), but changed to yellowish red (10 YR 7/6) after 96 h, followed
by more intensified darkening (10 YR 7/8) with embryonic development until
eclosion after ca. 120 h. Hinton (1981) concluded that many insect eggs
undergo coloration change during incubation due either to a change in color
of the chorion or to a color change of the embryo as seen through the
eggshell. We observed that the maturing embryo caused color changes in
the chorion.

Withycombe (1922) reported the surface of H. stigma eggs was smooth,
but broken by many small granular pits. Smith (1923) described the chorion
as dotted with undulating rows of minute or microscopic elongate white
raised, rounded spots. Scanning electron microscopy revealed that the
““spots’* consisted of irregular shaped waxlike projections somewhat evenly
spaced on a smooth chorion (Fig. 1b). These projections form diagonal
patterns around the egg. However, the chorion may lack these reticulations
where the egg is attached to the substrate. The projections near the anterior
pole fuse to encircle the micropylar process. The waxlike projections di-
rectly adjacent to the micropyle are connected by narrow bridges similar to
those on the chorion of Chrysopa carnea Stephens (Mazzini, 1976). The
projections on the surface of the chorion probably form a boundary layer

VOLUME 84, NUMBER 1 207

to retain air which establishes a humidity gradient that retards water loss
(Hinton, 1981).

Described simply as a button-like structure on most hemerobiid eggs
(Smith, 1923; Withycombe, 1923), the micropylar process is nipple-shaped
and consists of irregularly shaped, tubercle-like projections grouped into
sections separated by fissures (Fig. Ic). Narrow bridges proximally located
interconnect each tubercle with adjacent tubercles. Each tubercle possesses
an aperture usually centrally located. Additional studies may reveal that
ultrastructure of hemerobiid eggs is diagnostic at the species level.

ACKNOWLEDGMENT

Grateful appreciation is extended to A. G. Wheeler, Jr., Bureau of Plant
Industry, Pennsylvania Department of Agriculture, for his critique of this
manuscript before publication.

LITERATURE CITED

Hinton, H. E. 1981. Biology of insect eggs. Pergamon Press Ltd, New York, Vol. 1. 473 pp.

MacLeod, E. G. and L. A. Stange. 1981. The brown lacewings of Florida (Neuroptera: Hem-
erobiidae). Fla. Dep. Agric. Consum. Serv. Div. Plant Ind. Entomol. Circ. 227: 1-4.

Mazzini, M. 1976. Fine structure of the insect micropyle—III. Ultrastructure of the egg of
Chrysopa carnea Steph. (Neuroptera: Chrysopidae). Int. J. Insect Morphol. Embryol.
5: 273-278.

Munsell Book of Color. 1929. Munsell Color Co., Inc., Baltimore. 26 color charts.

Smith, R. C. 1923. The life histories and stages of some hemerobiids and allied species (Neu-
roptera). Ann. Entomol. Soc. Am. 16: 129-151.

Withycombe, C. L. 1922. Life-history of Hemerobius stigma Steph. Entomologist 55: 97-99.

. 1923. Notes on the biology of some British Neuroptera (Plannipennia). Trans. Ento-

mol. Soc. Lond. 1922: 501-594.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 208-209

NOTE

The Status of Nomia foxii Dalla Torre
(Hymenoptera: Halictidae)

T. D. A. Cockerell mixed the various species of Nomia (Curvinomia) in
his published papers (e.g., see Ribble. 1965. Univ. Kans. Sci. Bull. 45: 302-
303). Curvinomia contains six valid species and one valid subspecies. Even
though I (1965: 277-359) revised this subgenus and examined the type-spec-
imens still in existence, confusion over the proper scientific name remains
with three of the species (Hurd and Linsley. 1974. Proc. Entomol. Soc.
Wash. 76: 198-199; Hurd. 1979. pp. 1941-1942. In Krombein, K. V. et al.,
eds., Catalog of Hymenoptera in America North of Mexico. Smithsonian
Institution Press).

The three species in question and their synonyms are as follows:

1. Nomia (Curvinomia) tetrazonata Cockerell, 1910 (=moctezumae
Crawford, 1911; californiensis Michener, 1937).

2. Nomia (Curvinomia) universitatis Cockerell, 1908 (=punctata var.
(?) Fox, 1893).

3. Nomia (Curvinomia) foxii Dalla Torre, 1896, new name for punctata
Fox (=punctata Fox, 1893 [preoccupied by Smith, 1875]; mesillensis Cock-
erell, 1908).

Even though specimens of tetrazonata have been misidentified, as well
as other species as it, by Cockerell and others, there is no question that
moctezumae and californiensis are junior synonyms (see key and diagnoses
in Ribble, 1965, for identification).

In 1965 (p. 296) I stated, ‘‘Cockerell (1906a) selected Denver as the type
locality for foxii. This is an incorrect designation of the type locality, be-
cause the Denver [and South Dakota] specimen probably was an universi-
tatis, which is Fox’s ‘variety.’ Cockerell later (1908) correctly selected the
‘New Mexico species as the true foxii’ (!).”’ In selecting a lectotype, the
rules and procedures of zoological nomenclature require the lectotype to be
‘typical’ for the species and match the description (e.g., see Mayr, Linsley,
and Usinger. 1953. Methods and Principles of Systematic Zoology. Mc-
Graw-Hill, New York, pp. 241-242). I continued, ‘‘The specimen from New
Mexico most closely agrees with the description of punctata [=foxii D. T.],
while universitatis most closely agrees with Fox’s (1893. Entomol. News
4: 134-135) var. (?).”> The specimen from Denver has been lost, but on
circumstantial evidence it fits (contrary to Hurd and Linsley, 1974) univ-
ersitatis or Fox’s var. (?) which then would not be typical of foxii.

' Two errors in this paper may cause confusion: Page 198, line 20, ‘‘Ribble (1905)’’ should
be “Ribble (1965)’’; and line 38, ‘‘new Mexico species’’ should be ‘‘New Mexico species.”

VOLUME 84, NUMBER | 209

The restrictions of the type-locality by Cockerell (1906. Trans. Am. Ento-
mol. Soc. 32: 289-314; 1908. Ann. Mag. Nat. Hist. (8) 2: 323-334) were not
accompanied by the designation of a lectotype. I (1965: 298) was the first
to designate a lectotype for foxii: ‘‘Cockerell (1908) did not specifically
designate the New Mexico specimen as the lectotype, but that was clearly
his intent. I therefore here designate the female from Vega San Jose, New
Mexico, as the lectotype of N. foxii.”’ In view of the fact that the New
Mexico specimen was the only remaining syntype of foxii, this action seems
entirely prudent. An amendment to Article 74. Lectotypes.—_(a) Designation
of a specimen.—(1974. Bull. Zool. Nomencl. 31: 85) of the International
Code of Zoological Nomenclature (1964. London) stated ‘‘(ii) The first pub-
lished designation of a lectotype supersedes all previous restrictions of the
use of the name of the species. Example.—The type-locality becomes the
geographical place of origin of the lectotype, despite any previous restric-
tions of the type-locality.”’

Because of my lectotype designation, mesillensis is a junior synonym of
foxii, and the list above, therefore, shows the correct synonymies as I did
in 1965.

D. W. Ribble, 197 Poplar Avenue, Campbell, California 95008.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, pp. 209-210

NOTE

Hybridization between Gerris alacris Hussey and
Gerris comatus Drake and Hottes (Hemiptera:
Heteroptera: Gerridae) in Nature

Males of two middle-sized northeastern species of Gerris Fabricius, G.
comatus Drake and Hottes and G. alacris Hussey, are easily identified.
Tufts of golden hairs within the two impressions of the first genital segment
ventrally are unique among males of G. comatus and a prominent meso-
sternal scent gland (omphalium) characterizes G. alacris males (Calabrese.
1974. Mem. Conn. Entomol. Soc., pp. 227-266; Drake and Harris. 1934.
Ann. Carnegie Mus. 23: 179-240). While collecting specimens from adjacent
populations of G. alacris and G. comatus at a lagoon at Presque Isle State
Park, Pennsylvania in June 1980, I took a certain hybrid between the
species. The venter of the specimen, a male, is shown (Fig. 1). Both the
long hairs on the first genital segment ventrally and the prominent omphal-
ium are present.

210 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

&

Fig. |. Male hybrid of Gerris comatus x G. alacris, venter of abdomen.

Diane M. Calabrese, Department of Biology, Trinity College, Washing-
ton, D.C. 20017; present address: Department of Biology and The Wildlife
Sanctuary, Dickinson College, Carlisle, Pennsylvania 17013.

PROC. ENTOMOL. SOC. WASH.
84(1), 1982, p. 211

NOTE

New Records of Mites (Acari: Digamasellidae; Erythraeidae) Phoretic on
Biting Midges (Diptera: Ceratopogonidae)

Records of larval mites associated with adult biting midges are fairly well
known (Whitsel and Schoeppner. 1967. Proc. Entomol. Soc. Wash. 69: 284—
286; Grogan and Navai. 1975. Proc. Entomol. Soc. Wash. 77: 214-215). At
least two associations have yielded new species of Trombidiform mites
(Vercammen-Grandjean. 1957. Ann. Mag. Nat. Hist. 10: 283-286; Vercam-
men-Grandjean and Cochrane. 1974. J. Kans. Entomol.Soc. 47: 66-79).
Recent evidence by Grogan and Navai (1975) and Grogan (1977. Proc. Ento-
mol. Soc. Wash. 79: 24) indicated that adult mites in the families Phytosei-
idae and Macrochelidae (Parasitiformes: Gamasida) may be phoretic on
adult biting midges. Two new records of phoresy are presented in this paper.

An adult female Bezzia bivittata (Coquillett) taken by W. J. Hanson in
west Hodges Canyon, Cache County, Utah by Malaise trap 25-28 July 1978
had a mite attached to its abdomen. The mite was removed and mounted
in phenol-balsam and was identified as an adult Leptus sp. (Erythraeidae).
This is the first record of this mite family on ceratopogonids and the first
record of adult mites of the order Acariformes, suborder Actinedida on
these midges. Krantz (1978, A Manual of Acarology, 509 pp.) indicates that
larval erythraeid mites are parasitic on a variety of insects and other ar-
thropods. Krantz (1978) also states that adult and nymphal erythraeids are
predaceous on small insects or other mites. In this particular instance it is
most parsimonious to assume that this association represents a case of pho-
resy due to the fact that the mite was much smaller than the fly.

An adult male Dasyhelea oppressa Thomsen taken by Malaise trap 15-
21 May 1979 by W. L. Grogan, Jr., in Salisbury, Wicomico County, Mary-
land had a mite attached to its abdomen that was removed and mounted in
phenol-balsam. This mite was identified as an adult Digamasellus sp. (Di-
gamasellidae) of the order Parasitiformes, suborder Gamasida. This is the
first record of this mite family associated with ceratopogonids. According
to Krantz (1978) phoresy is common for digamasellid mites on geotrupine
dung beetles.

The above two mites are deposited in the acarology collection of the
National Museum of Natural History. Special thanks are extended to Robert
L. Smiley, Systematic Entomology Laboratory, USDA, Beltsville, Mary-
land, for identification of the mites and for suggestions on the manuscript.

William L. Grogan, Jr., Department of Biological Sciences, Salisbury
State College, Salisbury, Maryland 21801.

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CONTENTS
(Continued from front cover)
MILLER, G. L. and P. L. LAMBDIN—Hemerobius stigma Stephens (Neuroptera:
Hemerobiidae): External morphology of the egg

NORTON, R. A.—Arborichthonius n. gen., an unusual enarthronote soil mite (Acarina:
Oribatei) from Ontario

ORTH, R. E.—Five new species of Pherbellia Robineau-Desvoidy, subgenus Oxytaenia
Sack, from North America (Diptera: Sciomyzidae)

PINTO, J. D.—The phenology of the plant bugs (Hemiptera: Miridae) associated with
Ceanothus crassifolius in a chaparral community of southern California

SELANDER, R. B. and N. J. AGAFITEI—First-instar larvae of the Uniforma Group
of the genus Epicauta (Coleoptera, Meloidae)

SMITH, D. R.—Symphyta (Hymenoptera) of Sri Lanka

STEYSKAL, G. C.—A second species of Ceratitis (Diptera: Tephritidae) adventive
in the New World

STIMMEL, J. F.—Seasonal history of the white peach scale, Pseudaulacaspis pentagona
(Targ.-Tozz.) (Homoptera: Diaspididae), in northeastern Pennsylvania

THORPE, K. W.—Six Trichogramma (Hymenoptera: Trichogrammatidae) species asso-
ciated with a Maryland cornfield, with description of a new species

WHEELER, A. G., JR.—Coccobaphes sanguinarius and Lygocoris vitticollis (Hemip-
tera: Miridae): Seasonal history and description of fifth-instar, with notes on
other mirids associated with maple

WHITEHEAD, D. R.—Foods of Caulophilus spp., particularly the broadnosed grain
weevil, C. oryzae (Gyllenhal), based on interception records (Coleoptera: Curculion-
idae: Cossoninae)

NOTES:

CALABRESE, D. M.—Hybridization between Gerris alacris Hussey and Gerris
comatus Drake and Hottes (Hemiptera: Heteroptera: Gerridae) in nature

GROGAN, W. L., JR.—New records of mites (Acari: Digamasellidae; Erythraeidae)
phoretic on biting midges (Diptera: Ceratopogonidae)

KINGSOLVER, J. M.—Authorship of the family name Erotylidae (Coleoptera)
RIBBLE, D. W.—The status of Nomia foxii Dalla Torre (Hymenoptera: Halictidae) ...

SIS iy) A (ISSN 0013-8797)

hana TART

of the

‘ 84 APRIL 1982 NO. 2

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

CONTENTS

ADAMSKI, D. and T. M. PETERS—Axillary structure of the Tortricidae (Lepidop-
FES An ET gel PN ay ey ie ee OMT Eye heye pete, Slaters ohh Tcvets, wT TLeere, Oat & cis) Bis ane ool et eee

BLANCHARD, A. and J. G. FRANCLEMONT—Marilopteryx carancahua, a new
genus and new species from east Texas (Lepidoptera: Noctuidae: Hadeninae) ...

DELONG, D. M.—New species of Xestocephalinae (Homoptera: Cicadellidae) from
NMAC ear ATst cl AE CRUD ARC ER EZ lace % sofa. d<) $12 Septic a6 oct wse cereldle ate way» Seuret aye eal

DUARTE, J. and D. M. CALABRESE—Is the binomen Lygaeus kalmii Stal (Hemip-
tera: Heteroptera: Lygaeidae) applied to sibling species? .............+..0s0e00s

_ EADS, R. B., G. C. SMITH, and G. O. MAUPIN—A CO, platform trap for taking
adult Permacentor aqnaersont (Acari TXOGIdaC) 5.4... 6 seen nnd ms essere eeeenne
GORDON, R. D.—An old world species of Scymnus (Pullus) established in Pennsyl-
vane and -Newerork (Coleoptera; Coccinellidac) ca... 05 <0. s vox cus wena emis
GORDON, R. D.—Two new species of Nephaspis Casey (Coleoptera: Coccinellidae)
OM UAE TICE COIOMIS — 2.5.5.<s,5<6c esis iclaie bins WON ES Visine A eres acniatd eo wimalereetale

HENRY, T. J.—New synonymies and a new combination in the North American

TUBA Aries EMR HAN POT SLA Meee arac-Fo 2)e Rial dal hie’ ack Wtecls aba tocTMaNG cape = ors Natete Sere Oe ete EW ere mL 337
HENRY, T. J. and J. M. CAPRILES—The four ‘‘ocelli’’ of the isometopine genus
Isometocoris Carvalho and Sailer (Hemiptera: Miridae) ...........6..eee senses 245

HOEBEKE, E. R. and A. G. WHEELER, JR.—Rhopalus (Brachycarenus) tigrinus,
recently established in North America, with a key to the genera and species of

y Rhopalidae in eastern North America (Hemiptera: Heteroptera) ................ 213
KELLEY, R. W. and J. C. MORSE—A key to the females of the genus Oxyethira
(Trichoptera: Hydroptilidae) from the southern United States ................45. 256

(Continued on back cover)

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PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 213-224

RHOPALUS (BRACHYCARENUS) TIGRINUS, RECENTLY
ESTABLISHED IN NORTH AMERICA, WITH A KEY TO THE
GENERA AND SPECIES OF RHOPALIDAE IN EASTERN
NORTH AMERICA (HEMIPTERA: HETEROPTERA)

E. RICHARD HOEBEKE AND A. G. WHEELER, JR.

(ERH) Department of Entomology, Cornell University, Ithaca, New York
14853; (AGW) Bureau of Plant Industry, Pennsylvania Department of Ag-
riculture, Harrisburg, Pennsylvania 17110.

Abstract.—Rhopalus (Brachycarenus) tigrinus (Schilling) was collected
in New Jersey in 1977, the first record of this European species for North
America; it also has been found in Pennsylvania and on Long Island, New
York. This rhopalid is briefly described and its dorsal habitus, diagnostic
external features, and male genitalic characters are illustrated. Known host
plants of the family Cruciferae are recorded, and North American locality
records are listed and mapped. A key to the genera and species of Rhopal-
idae in eastern North America is presented, including for the first time R.
tigrinus.

A European scentless plant bug, Rhopalus (Brachycarenus) tigrinus
(Schilling), was recently reported new to North America (Hoebeke, 1977).
This detection resulted from the USDA-APHIS ‘‘High Hazard Pest Survey”
program and was based on specimens submitted (to ERH) for identification.
The first North American collection of this species consisted of specimens
taken in southern New Jersey in 1977. We initiated a survey of portions of
south-central and southern New Jersey to determine the distribution and
relative abundance of this newly detected species, and to gather information
on its biology, seasonal history, and host plant preferences.

In this paper we describe and illustrate the important adult characters of
R. tigrinus, record its known host plants, list and map locality records in
North America, and provide a key to the genera and species of Rhopalidae
in eastern North America, including Rhopalus tigrinus.

The adults of Rhopalus tigrinus (Fig. 1) closely resemble those of some
native species of rhopalids occurring in eastern North America (especially
species of Arhyssus, Niesthrea, and Liorhyssus), but may readily be distin-

214 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

-

Fig. 1. Rhopalus (Brachycarenus) tigrinus, dorsal habitus.

guished by the following diagnosis: Head more than 12x as broad as long;
bucculae covering *% the length of Ist rostral segment in profile; rostrum not
usually extending beyond mesosternum; metapleuron with posterior margin
strongly angulate with prominent postero-lateral projection (Fig. 4); male
pygophore and paramere as in Figs. 2, 3; dorsal coloration yellowish or
grayish with black markings on the head, anterior margin of the pronotum,
anterior corners of the scutellum, and spots on the veins of the hemelytra;
abdomen beneath hemelytra black except for yellow markings at apex; con-
nexiva generally yellow, sometimes with black spots; 6.5—7.0 mm in length.

Rhophalus tigrinus is considered a relatively common species in the Pale-

VOLUME 84, NUMBER 2 215

Figs. 2-4. Rhopalus (Brachycarenus) tigrinus. 2, Male pygophore. 3, Paramere. 4, Meta-
pleural region and base of hindleg, showing extreme prolongation of the dorsal half of meta-
pleuron (arrow). (Figs. 2 and 3 redrawn from Gollner-Scheiding, 1978b.)

216 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

arctic Region, found throughout much of Europe (except the British Isles)
and the Mediterranean Region including Algeria and Morocco, Madeira and
Canary Islands, and Asia Minor. In its native habitat little or nothing is
known about its biology and habits. The egg and third-instar nymph of R.
tigrinus have been figured and included in a key to these life stages by
Puchkov and Puchkova (1956) and Puchkova (1957). According to Lindberg
(1953), the species is found chiefly on low plants and weeds in waste places
adjoining cultivated areas. In European USSR, tigrinus is noted by Kerzh-
ner (1967) to be a general feeder with an apparent preference for members
of the mustard family, the Cruciferae. Stichel (1960) has reported figrinus
feeding on weedy crucifers such as hoary alyssum, Berteroa incana (L.)
DC; flixweed, Descurainia sophia (L.) Webb (cited as Sisymbrium sophia);
and Alyssum incanum L. Stichel also noted tigrinus on Trifolium (Legu-
minosae) and Cirsium arvense (L.) Scop. (Compositae). In a study of He-
miptera associated with crucifers in Poland, Studzinski and Malachowska
(1973) collected adults of R. tigrinus on various species of Cruciferae.

Rhopalus tigrinus at present is known in eastern North America only from
a few localities (see Fig. 25). The original collection consisted of two adult
males taken from wheat on a farm at North Vineland, New Jersey (Cum-
berland Co.), 16 August 1977, by R. Morris. In 1978, eight specimens were
collected from weeds, 5 September, and three specimens from a pasture, |
September, on farms also at North Vineland. Ten specimens were taken
from grasses in a railroad yard at Camden, New Jersey (Camden Co.),
Jefferson Avenue at 4th Street, 4 August 1978, by B. C. Emens. We have
taken specimens at the same North Vineland and Camden locations in 1979
and 1980. AGW has found populations of adults and nymphs feeding and
developing on crucifers in two railroad yards in Pennsylvania, one at west
Philadelphia (along Pennsylvania Railroad near 30th Street Station) in 1980
and the other near Harrisburg (Enola Railroad Yards) in 1980 and 1981. We
have found a population along a railroad track south of Yaphank, Long
Island, on 30 May 1981. Furthermore, AGW and T. J. Henry (Systematic
Entomology Laboratory, USDA) collected additional specimens by sweep-
ing vegetation near Medford, Long Island (Suffolk Co.), on 29 August 1981.
The fact that specimens of tigrinus have been taken on plants in three
railroad yards in New Jersey and Pennsylvania and along the railroad on
Long Island strongly indicates that this species may be dispersed passively
via the railroad. Further study and field observations will be needed to
support this speculation. Many other examples of this type of passive dis-
persal may be cited (e.g., Catorhintha mendica Stal (Balduf, 1957); Coc-
cinella undecimpunctata L. (Wheeler and Hoebeke, 1981); Subcoccinella
vigintiquatuorpunctata L. (Wheeler and Henry, 1981)).

Our field and laboratory observations supplement the list of host plants
recorded in the literature and support the supposition that R. tigrinus ex-
hibits a preference for species of Cruciferae. We have observed or have

VOLUME 84, NUMBER 2 217

———
Gi. cs
Figs. 5-10. Structural characters. 5, Dorsal view of head, Harmostes reflexulus. 6, Dorsal
view of head, H. fraterculus. 7, Dorsal view of apex of female abdomen, Arhyssus lateralis.
8, Metapleural region, Stictopleurus punctiventris. 9, Metapleural region, Arhyssus nigrister-

num. 10, Dorsal view of apex of female abdomen, A. nigristernum. (Figs. 5, 6,8, and 9 redrawn
from Hoffman, 1975.)

taken eggs, nymphal stages, or adults from the following cruciferous
species: Capsella bursa-pastoris (L.), shepherd’s purse; Lepidium virgini-
cum L., peppergrass; Thlaspi perfoliatum L., a pennycress; and Arabidopsis
thaliana (L.), mouse-ear cress. We will provide a summary of the biology
and seasonal history, and descriptions and illustrations of the egg and
nymphal instars in a forthcoming paper.

The following key is provided to aid in the recognition and identification
of all genera of Rhopalidae now known to occur in North America (modified
principally after Chopra, 1967; Harris, 1943; and Slater and Baranowski,
1978), and all species that occur in eastern North America’ (adapted from

1 Eastern North America is defined here as in Blatchley (1926): “‘the United States east of
the Mississippi River and Canada east of the 90th Meridian.”’

218 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

11 12

14

Figs. 11-16. Male genitalia. 11, Pygophore, Jadera haematoloma. 12, Pygophore, Sticto-
pleurus punctiventris. 13, Paramere, S. punctiventris. 14, Pygophore, Jadera antica. 15, Py-
gophore, Stictopleurus knighti. 16, Paramere, S. knighti. (Figs. 11 and 14 redrawn from Goll-
ner-Scheiding, 1979; Figs. 12, 13, 15, and 16 redrawn from Gollner-Scheiding, 1975.)

Hoffman, 1975; Harris, 1944; Chopra, 1968, 1973; Blatchley, 1926; and Gdll-
ner-Scheiding, 1975, 1976, 1978a, 1978b, 1979, 1980).

KEY TO GENERA AND SPECIES OF RHOPALIDAE IN
EASTERN NORTH AMERICA

1. Lateral pronotal margins distinctly notched behind anterior margin;
generally conspicuously colored; 11 mm or more in length ........ Zi
— Lateral pronotal margins straight or slightly sinuate, without a dis-
tinct notch behind anterior margin; generally inconspicuously col-
ored; usually a little more than 9 mm
2. Bucculae long, reaching base of head; Jadera Stal (2 eastern
] 0) 9) laa care ae PPA etree a GS A De 3

VOLUME 84, NUMBER 2 219

17

Sie.
ey:

Figs. 17-24. Male genitalia. 17, Pygophore, Niesthrea sidae. 18, Paramere, N. sidae.
Pygophore, N. louisianica. 20, Paramere, N. louisianica. 21, Pygophore, Arhyssus ae
22, Paramere, A. lateralis. 23, Pygophore, A. nigristernum. 24, Paramere, A. nigristernum.
(Figs. 17-20 redrawn from Chopra, 1973; Figs. 21-24 redrawn from Chopra, 1968.)

— Bucculae short, not extending beyond middle of head; Boisea Kir-
Kalen SPoe sercccat: txaidier dhe Bastia aomtewm nis Boisea trivittata (Say)
Widespread throughout North America.

3. Species predominantly brownish black; male pygophore and par-
amere as in Fig. 11; large species, 9.6-13.3 mm in length .......

«etch ae Biles ROR IAC a Jadera haematoloma (Herrich-Schaeffer)
In the U.S. known from the southeastern states west to Louisiana,
Texas, and California, also from the central states including lowa,
Illinois, Kansas, Colorado, Oklahoma, and Missouri; also in Mex-
ico, Middle America, and the West Indies to Colombia and Vene-
zuela, and in Hawaii.

? Historically, the species trivittata, best known as the boxelder bug, has been placed in the
genus Leptocoris. Boisea Kirkaldy, originally proposed as a subgenus of Leptocoris and in-
cluding trivittata, is accorded generic rank in a recent revision by Gollner-Scheiding (1980) and
is SO recognized here.

220

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Species predominantly brownish red or only reddish; male pygo-
phore and paramere as in Fig. 14; smaller species, 7.4-10.3 mm in
PESITESENT oc Beso BP aha d'c wisehs oft no: career Jadera antica (Walker)
Florida and the West Indies and south to Central America.

. Hindfemur incrassate, bearing a series of prominent spines ........ 2)

Hindfemur not incrassate, lacking prominent spines .............. 8

. Abdomen not dilated; hemelytra covering abdominal connexiva;

Harmostes Burmeister (3'Spp:) |. «+. 2462-524 Je 2 oe eee 6
Abdomen dilated laterally; hemelytra not covering abdominal con-
nexiva; connexiva widely exposed; Aufeius Stal (1 sp.) ........

SE A Cee eee rae ne I A 3 Aufeius impressicollis Stal
Ohio west to South Dakota, Colorado, and California and south-
west to Arizona and Mexico.

. Lateral pronotal margins irregularly, but distinctly, finely toothed

RO 5.5 ols Bearing sies ale Maen ke ae Harmostes serratus (Fabricius)
Texas, Florida, Arizona, California, Mexico, and Cuba.
[eateral pronotal margins entire {2% = 32.25. -- 2s eee ee 7

. Basal antennal segment large, thickened, and extending far beyond

apex of tylus (Fig. 5); costal margin of hemelytra usually unicolor-

USE PEs ac Aik Se uae Cee Ce ee Harmostes reflexulus (Say)
Generally distributed throughout U.S. and southern Canada.

Basal antennal segment relatively small, scarcely exceeding apex

of tylus (Fig. 6); costal margin of hemelytra spotted with fus-
CUS uit cu. Sitters fens fone bar Rca Ghia Harmostes fraterculus (Say)
New Jersey west to southern Indiana, Oklahoma, and California
south to Georgia and Arizona.

. Metapleuron not, or only indistinctly divided into episternum and

epimeron; metathoracic scent gland openings indistinct or absent;
pronotal cicatrices generally ending in a closed loop; Stictopleurus
Stal(2 Spp.) . 5. 0% oe od SE ee eee eee 9
Metapleuron distinctly divided into episternum and epimeron;
metathoracic scent gland openings conspicuous; pronotal cicatrices
notending ‘ina:closedi1oape ees As ie ss en 10

. Male pygophore as in Fig. 12; paramere with a pronounced globose

enlargement basally, strongly tapering distally, recurved so that
posterior face is concave (Fig. 13); female usually grayish yellow
withtdark markings...) 225.04 Stictopleurus punctiventris (Dallas)
Holarctic, across Canada and northern U.S. and southward
along mountains.

Male pygophore as in Fig. 15; paramere without subbasal enlarge-
ment, flat and tapering (Fig. 16) female distinctly reddish .......

ES ae avea eal shi auc doce ee ee Stictopleurus knighti Harris
Michigan and Minnesota west to Wyoming.

. Pronotum with a distinct, but narrow collar anteriorly; pronotum

VOLUME 84, NUMBER 2 221

\ CLINTON

| FRANKLIN }
ST. LAWRENCE a=

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.
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—t_—§_—_J waYnE

MC KEAN POTTER | TIOGA

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A —J a z
SULLIVAN | yt? Fs
z
\ as
\ LycominNG ht, rg
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\ PH & ) :
CLEARFIELD 3 Fe } ron <p a A
J> CENTRE OOD aan Sr —~<EARBON

SNYDER (OmBER- > ae Y
NOSCHUYLRILE “g, *
“4

YS 4 \ &
z eae NO \- f
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CARROLL, *,
xe /% & os

c ~~~ %
J rreoenicn> 9 %0,%0, 2 \ °
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Fig. 25. Distribution of Rhopalus (Brachycarenus) tigrinus in eastern North America. Map
shows N.Y., Pa., N.J., Md., and Del.

between collar and cicatrices forming a distinct ridge which is pol-
ished and impunctate, or at most with a few punctures; Liorhyssus
StalkGeispey. eee, fe gee ENG VERS Liorhyssus hyalinus (Fabricius)
Cosmopolitan; widespread throughout North America.
— Pronotum without collar anteriorly; pronotum anterior to cicatrices
not smooth or polished, always with numerous coarse punctures... 11
11. Head quite short, more than 12x as broad as long; rostrum short,
not or barely extending to metasternum; postero-lateral margin of

299

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

metapleuron strongly angulate, with a prominent and pointed pro-
jection laterally (Fig. 4); male pygophore and paramere as in Figs.
2, 3; Rhopalus (Brachycarenus) Schilling (1 sp.), ... ...: 2 4.2508

th RT eT eee er te 5 ke a2 Rhopalus tigrinus (Schilling)
A Palearctic species, reported new to North America; known only
from New Jersey, Pennsylvania, and New York (Long Island).
Head longer, less than 112 as broad as long; rostrum extending
beyond metasternum; postero-lateral margin of metapleuron usu-
ally straisht.or slightly’sinuate (Figs.(85 9). <0... eee ee 12

. Hindtibia possessing a number of black rings; rostrum extending

to or beyond 3rd abdominal sternum; Niesthrea Spinola (2 spp.) .. 13
Hindtibia often speckled or spotted with black or fuscous, but lack-
ing a series of black rings; rostrum not extending posteriorly be-
yond 3rd abdominal sternum; Arhyssus Stal (3 spp.) ............. 14

. Small species, usually less than 6.5 mm; male pygophore with me-

dian lobe constricted at base, broad proximally and slightly con-
cave distally (Fig. 17); paramere slender with subapical projection,
broad and somewhat flattened distally (Fig. 18) ................

Saxe Penile 8 See ones otek Race ttt rai Metre Niesthrea sidae (Fabricius)
Maryland south to Florida and west to Oklahoma, Texas, and Ar-
izona; also in South America.

Larger species, usually much greater than 6.5 mm; male pygophore
with median lobe slightly constricted medially and concave distally
(Fig. 19); paramere slender, with subapical and dorsal projection
(EE 220) ces Gi esd ecco eee Niesthrea louisianica Sailer
Long Island, New York south to Florida, west to Arizona and
California, and in Mexico; more common in southern states.

. Scutellum reaching to or beyond clavus; antenniferous tubercles

inconspicuous or obsolete ......2.. 24-24; Arhyssus hirtus (Bueno)
An eastern species, known only from New York (Long Island) and
Massachusetts.

Scutellum not reaching clavus; antenniferous tubercles readily vis-

(1 (0, SA er nee Oe Meme E! Sl Hunton Weed nell os a cos - 15

. Apex of last abdominal tergum of female broadly rounded in dorsal

outline (Fig. 7); male pygophore with median lobe slightly concave
(Fig. 21); paramere broad in middle, lateral projection with tip
broadiysrounded!(Fig::22)', =.....0henk eee Arhyssus lateralis (Say)
Widely distributed in North America and ranging into southern
Mexico.

Apex of last abdominal tergum of female subacuminate or pointed

in dorsal outline (Fig. 10); male pygophore with median lobe al-
most truncate (Fig. 23); paramere flat, broad, lateral projection with

tip narrowly rounded (Fig. 24) ..... Arhyssus nigristernum (Signoret)

VOLUME 84, NUMBER 2 223

Widely distributed in eastern North America, ranging from Quebec
and Ontario south to central Florida and west to Oklahoma and
Kansas.

ACKNOWLEDGMENTS

We thank John E. Rawlins, Cornell University, Ithaca, N.Y., for reading
and commenting on the manuscript.

LITERATURE CITED

Balduf, W. V. 1957. The spread of Catorhintha mendica Stal. Proc. Entomol. Soc. Wash. 59:
176-185.

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America, with especial
reference to the faunas of Indiana and Florida. Nature Publishing Co., Indianapolis,
1116 pp., 215 figs., XII pls.

Chopra, N. P. 1967. The higher classification of the family Rhopalidae (Hemiptera). Trans. R.

Entomol. Soc. Lond. 119: 363-399, 75 figs.

. 1968. A revision of the genus Arhyssus Stal (Rhopalidae: Hemiptera). Ann. Entomol.

Soc. Am. 61: 629-655, 146 figs.

——.. 1973. A revision of the genus Niesthrea Spinola (Rhopalidae, Hemiptera). J. Nat.
Hist. 7: 441-459, 16 figs.

Gollner-Scheiding, U. 1975. Revision der Gattung Stictopleurus Stal. Dtsch. Entomol. Z. 22:
1-60.

—. 1976. Revision der Gattung Liorhyssus Stal, 1870 (Heteroptera, Rhopalidae). Dtsch.

Entomol. Z. 23: 181-206.

. 1978a. Revision der Gattung Harmostes Burm., 1835 (Heteroptera: Rhopalidae) und

einige bemerkungen zu den Rhopalinae. Mitt. Zool. Mus. Berl. 54: 257-311.

. 1978b. Bemerkungen zu der Gattung Rhopalus Schilling einschliesslich Brachycarenus

Fieber (Heteroptera, Rhopalidae). Mitt. Zool. Mus. Berl. 54: 313-330.

. 1979. Die Gattung Jadera Stal, 1862 (Heteroptera, Rhopalidae). Dtsch. Entomol. Z.

26: 47-75.

. 1980. Revision der afrikanischen Arten sowie Bemerkungen zu weitern Arten der

Gattung Leptocoris Hahn, 1833, und Boisea Kirkaldy, 1910. Dtsch. Entomol. Z. 27:

103-148.

Harris, H. M. 1943. Concerning the Rhopalidae (Hemiptera). lowa State Coll. J. Sci. 17:
197-204.
. 1944. Concerning American Rhopalini (Hemiptera, Rhopalidae). lowa State Coll. J.
Sci. 19: 99-109.

Hoebeke, E. R. 1977. (Note). Jn Cooperative Plant Pest Report 2(40): 802.

Hoffman, R. L. 1975. Squash, broad-headed and scentless plant bugs of Virginia (Hemiptera:
Coreoidea: Coreidae, Alydidae, Rhopalidae), /n *‘The Insects of Virginia,’ No. 9, Va.
Polytech. Inst. State Univ. Res. Div. Bull. 105, 52 pp., 20 figs.

Kerzhner, I. M. 1967. Order Hemiptera (Heteroptera), pp. 851-1118. Jn Bei-Beinko, G. Ya.,
ed., Keys to the insects of the European USSR, Vol. |. (Translated from Russian. Israel
Program for Scientific Translations, Jerusalem.)

Lindberg, H. 1953. Fam. Coreidae (subfam. Corizinae), pp. 52-54. Jn Hemiptera Insularum
Canariensium. Societas Scientiarum Fennica, Commentationes Biologicae XIV.1. Cen-
tral tryckeriet, Helsingfors.

Puchkov, V. G. and L. V. Puchkova. 1956. A key to the eggs and larvae of Hemiptera-
Heteroptera injurious to crops. Tr. Vses. Entomol. Obva. 45: 218-342. (In Russian.)

224 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Puchkova, L. V. 1957. Eggs of Hemiptera-Heteroptera. III. Coreidae (Supplement), IV.
Macrocephalidae. Entomol. Obozr. 36: 44~58. (In Russian.)

Slater, J. A. and R. M. Baranowski. 1978. Family Rhopalidae, the scentless plant bugs, pp.
66-70. In How to know the true bugs (Hemiptera-Heteroptera). Wm. C. Brown Com-
pany Publishers, Dubuque, lowa.

Stichel, W. 1960. Pentatomomorpha, Coreiodea: Corizidae, vol. 4, 14 Heft, pp. 417-448. In
Illustrierte Bestimmungstabellen der Wanzen. II. (Europa Hemiptera-Heteroptera Eu-
ropae). Martin-Luther, Berlin-Hermsdorf.

Studzinski, A. and D. Malachowska. 1973. Tluskwiaki Roznoskrzydle (Heteroptera) wyste-
pujace na dziko Rosnacych roslinach Krzyzowych (Cruciferae) w Polsce w 1970 R.
[Hemipterous bugs (Heteroptera) occurring in 1970 on wild cruciferous plants (Crucif-
erae) in Poland]. Rocz. Nauk. Roln. Ser. E Ochr. Rosl. 3: 79-100. (In Polish with English
summary.)

Wheeler, A. G., Jr. and T. J. Henry. 1981. Seasonal history and habits of the European alfalfa
beetle, Subcoccinella vigintiquatuorpunctata L. (Coleoptera: Coccinellidae). Coleopt.
Bull. 35(2): 197-203.

Wheeler, A. G., Jr. and E. R. Hoebeke. 1981. A revised distribution of Coccinella undecim-
punctata L. in eastern and western North America (Coleoptera: Coccinellidae). Coleopt.
Bull. 35(2): 213-216.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 225-231

FURTHER OBSERVATIONS ON THE ETHOLOGY OF ALYSSON
CONICUS PROVANCHER (HYMENOPTERA: SPHECIDAE)

MARK F. O’BRIEN AND FRANK E. KURCZEWSKI

(MFO) Insect Division, Museum of Zoology, University of Michigan, Ann
Arbor, Michigan 48109; (FEK) Department of Environmental and Forest
Biology, S.U.N.Y. College of Environmental Science and Forestry, Syra-
cuse, New York 13210.

Abstract.—Females of Alysson conicus Provancher were observed nest-
ing in large aggregations in a sandy road and path in the Adirondacks. They
were active at ambient temperatures of from 15° to 22°C. Females stored
adult Cicadellidae (94% Empoa albicans Walsh) in their nests. From 5 to
12 leafhoppers were placed in the fully provisioned, spherical cells, 1.4 to
6.0 cm beneath the surface. Prey were transported to the nests in flight and
held by the beak and body with the wasp’s mandibles and legs, respectively.
Entrances were left open while provisioning. Prey (61%) were parasitized
by dryinid wasps in 1980 prior to capture by A. conicus.

Little is known about the bionomics of the genus Alysson in North Amer-
ica. Of the nine species in the Nearctic Region, only A. melleus Say has
been studied to any extent (Hartman, 1905; Rau and Rau, 1918; Evans,
1966; Kurczewski and Kurczewski, 1971).

Alysson conicus Provancher is one of the less commonly collected
species. It ranges throughout the northeastern U.S. from southeastern Can-
ada into Virginia (Krombein, 1979). The only known information on its
nesting behavior has been published by O’Brien and Kurczewski (1979).
The present report adds considerably more data and complements the be-
havioral information presented in 1979.

ECOLOGY

We observed A. conicus during August 6-10, 1979 and July 20-25, 1980
at the Cranberry Lake Biological Station of the College of Environmental
Science and Forestry, in St. Lawrence Co., N.Y. The wasps occupied a 52
m segment of a sandy road which parallels Sucker Brook (Fig. 1), and a
compacted, sandy-clayey path 2 m from the lake shoreline on the opposite
side of the Station. Dominant vegetation along these areas comprised sec-

226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Nesting habitat of Alysson conicus at Cranberry Lake, N.Y., showing 80 marked
nests within a 6.0 x 0.9 m area. Individual nests are denoted by white plastic stakes.

ond-growth stands of northern hardwoods, especially red maple (Acer ru-
brum), sugar maple (Acer saccharum), American beech (Fagus grandifolia),
yellow birch (Betula allegheniensis), poplar (Populus tremuloides), and
black cherry (Prunus serotina). Herbaceous plants, bracken fern (Preris
latiuscula), and grasses grew at the edges of the road and path. The road
was shaded during much of the day, and the sand in which the wasps nested
was uniformly moist. The daily ambient temperatures at the nesting sites
ranged from 15° to 22°C during periods of observation. Temperatures in 1979
were unusually cool at night, plunging once to 7°C. Several cool windy days
interspersed with rain in 1979 and periods of heavy rain in 1980 dampened
wasp activity.

Females of A. conicus were active from 0830 to 1630 h (EDT), being
most active on sunny days; however, they carried on some ‘‘typical’’ nest-
ing activities during periods of unfavorable weather. Between 0730 and 1000
h many females remained inside their nests, either in terminal cells or
lower parts of their burrows. When removed from their nests, such wasps
became active within 20 s.

Males were found in sunny spots within the nesting area or resting on
overhanging vegetation along the road between 0930 to 1400 h. Some
males were observed lapping honeydew off blackberry (Rubus sp.) leaves;
others were found on flowers of tall meadow rue (Thalictrum polygamum).

VOLUME 84, NUMBER 2 227

No copulation was observed; however, males frequently followed closely
behind provisioning females on the ground and hunting females on vegeta-
tion.

NESTING BEHAVIOR

Females were aggregative, possibly due to the limited suitable nesting
habitat. Distances between conspecific entrances ranged from 2 to 25 cm in
dense groups but over | m in others. The densest aggregation contained 80
nests within a6 x 0.9 m area (Fig. 1). Nest tumuli (276) were marked within
a 52 x 1.8 m stretch of the road in 1980. On the lakeside path, we marked
29 nest entrances within a 16.5 x 1.5 m area.

Six females were observed excavating burrows between 0830 and 1000 h,
and from 1500 to 1600 h. They formed pellets of sand with the forelegs,
transferred them backwards to the midlegs, and pushed them into the en-
trance with the hindlegs. The pygidium often assisted in pushing pellets into
the opening. The pellets were then pushed to the side of the entrance, the
female rotating in a clockwise direction. Often, after several pellets had
accumulated in the entrance, the female would move them to one side,
forming an asymmetrical tumulus. As a wasp dug deeper, she took increas-
ingly longer (10-25 s) to deposit the soil in the entrance.

Provisioning females flew into the nesting areas from the surrounding
vegetation at a sharp angle. One female, grasping a leafhopper venter-up by
its beak and body with her mandibles and legs, respectively, flew from a
red maple 1.5 m above ground level to the nest. Within 15 to 30 cm of her
entrance, she held the prey venter-up on the ground by its beak (Fig. 2).
Females walked in a zig-zag manner near their entrances, dropped their
prey just outside, entered, turned around, and pulled the leafhoppers inside
head-first. One wasp took 5 min between provisioning trips.

Provisioning wasps were skittish on the ground, and flew away at the
slightest provocation, often releasing the prey. Such prey were usually not
retrieved by the wasps. Some disoriented females flew in circles above the
area until they found their entrance, then retrieved the leafhopper and took
it into the nest.

All nests were left open during provisioning. Many nests were easily
recognized by their characteristic tumuli (Fig. 3), in the form of a simple
conical mound, an elongate-convex mound with the entrance off-center, or
a turret. Weathering was important in shaping the tumulus, the oldest ones
being either absent or inconspicuous. Dimensions of 33 tumuli varied from
4 to 18 mm high (¢ = 8.4 + 3.1 mm), and from 10 to 20 mm wide (* = 15.9
+262.3.mi).

Nest entrances (16) ranged from 1.5 to 2.5 mm in diameter (* = 2.1 + 0.2
mm). Burrows went either straight downward through symmetrical tumuli
or obliquely downward through asymmetrical ones (Fig. 4). Unfinished nests

228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 2, 3. 2, Alysson conicus female carrying prey to her nest. 3, Tumulus and nest en-
trance.

VOLUME 84, NUMBER 2 229

Fig. 4. Side views of Alysson conicus nests. Broken lines indicate burrows which could
not be traced exactly. Completed cells contained an egg (e) or larva (1). Uncompleted cells are
designated by *‘i.””

230 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(20) contained a single terminal cell, but 5 completed or nearly completed
nests had from 2 to 5 cells. Cells (34) were spherical (Fig. 4) with a mean
diameter of 4.5 + 0.7 mm (3.0-5.5 mm). Cell depth varied from 1.4 to 6.0
cm (% = 3.8 + 1.0 cm; N = 34) beneath the soil surface. The soil was in-
variably moist at cell depth.

Prey consisted of adult Cicadellidae, subfamily Typhlocylinae as follows:
218 Empoa albicans Walsh, 2 E. querci Fitch, 2 E. latifasciata Christian,
8 Empoasca atrolabes Gillette, 1 Typhlocyba persephone McAtee, and |
Ribautiana sp. The prey appeared to be either fully-paralyzed or dead, with
no visible appendage or breathing movements.

Fully-provisioned cells, i.e., those with an egg or larva, contained prey
placed head-inward and venter-upward. The mean number of prey per com-
pleted cell was 8.8 + 2 (5-12; N = 14). An egg was laid on the uppermost
leafhopper in the cell, either to the right or left of center on the sternum as
in A. melleus (see Fig. 20, Evans, 1966). Four eggs averaged 0.75 x 0.2
mm. Two cells contained prepupae within ovoidal cocoons, measuring
6.0 x 2.5 and 5.6 x 2.3 mm. The cocoons were constructed of silk, sand
grains, and leafhopper remains.

There was no evidence of parasitism or predation in 1979. In 1980, how-
ever, 61% of the prey were parasitized by dryinid wasps prior to capture by
A. conicus. Dryinidae are parasitoids of Homoptera, developing at first
internally, then extruding through the integument (Krombein, 1979). It is
doubtful that there would have been any effect upon A. conicus by the
dryinid infestation due to the faster development of the A. conicus larva.
Alysson conicus females may have captured parasitized leaf-hoppers be-
cause they were less successful than non-parasitized leafhoppers in evading
capture.

DISCUSSION

Alysson conicus exhibits the behavioral traits of the genus, as outlined by
Evans (1966). It nests in aggregations in cool, moist sandy situations, with
females hunting near their nests. Burrows are nearly vertical, with conspic-
uous, often irregular tumuli. Completed nests are 2- to 5-celled, as in A.
melleus (Evans, 1966), with the deeper cells in a nest being older.

Although we observed A. conicus females releasing prey outside their
entrances and pulling them in, other species of Alysson have been noted
carrying prey directly into their nests (Evans, 1966; Tsuneki, 1969). Perhaps
many of the entrances at Cranberry Lake were blocked with sand grains,
and the females had to clear these away prior to entering.

Alysson conicus prefers Empoa albicans as prey at Cranberry Lake, N.Y.
Empoa albicans was common on low vegetation, shrubs, and trees paral-
leling the nest sites, and the wasps hunted in these areas. Alysson melleus
takes a broader range of prey (Evans, 1966). Observations of A. conicus in

VOLUME 84, NUMBER 2 231

other areas would undoubtedly reveal differences from the Cranberry Lake
population in regards to prey selection.

It is surprising to find A. conicus preying on dryinid infested leafhoppers.
This phenomenon has not been noted for any other wasps preying on Ho-
moptera, although it would have been easily overlooked unless the dryinids
were extruding through their host’s integument.

Alysson conicus nests were shallower than those reported for A. melleus
(Evans, 1966) and were less variable in cell depth. The shallower A. conicus
nests may reflect the continual, moist sand in perpetual shade, or limitations
of the sand layer overlying the compacted organic layer of the road.

ACKNOWLEDGMENT

We thank J. P. Kramer, Systematic Entomology Laboratory, USDA,
Washington, D.C. for identifying the prey Cicadellidae.

LITERATURE CITED

Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps. Harvard Univ.
Press, Cambridge, Mass. xvi + 526 pp.

Hartman, C. 1905. Observations on the habits of some solitary wasps of Texas. Bull. Univ.
Texas 65: 1-72.

Krombein, K. V. 1979. Family Dryinidae, pp. 1240-1251; family Nyssonidae, pp. 1684-1720.
In Krombein, K. V. et al., eds., Catalog of Hymenoptera in America North of Mexico.
Vol. 2, Apocrita (Aculeata), Smithsonian Institution Press, Washington, D.C.

Kurczewski, F. E. and E. J. Kurczewski. 1971. Host records for some species of Nyssoninae.
J. Kans. Entomol. Soc. 44: 334-337.

O’Brien, M. F. and F. E. Kurczewski. 1979. Observations on the nesting behavior of Alysson
conicus Provancher (Hymenoptera: Sphecidae). Proc. Entomol. Soc. Wash. 81:
435-437.

Rau, P. and N. Rau. 1918. Wasp studies afield. Princeton Univ. Press, Princeton, N.J.
xv + 372 pp.

Tsuneki, K. 1969. Gleanings on the bionomics of the East-Asiatic non-social wasps (Hyme-
noptera). IV. Some species of Bembecini, Stizini, Gorytini, Mellinini, and Alyssonini.
Etizenia 41: 1-19.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 232-239

POECILOCRYPTICUS FORMICOPHILUS GEBIEN, A SOUTH
AMERICAN BEETLE ESTABLISHED IN THE UNITED STATES
(COLEOPTERA: TENEBRIONIDAE)'

WARREN E. STEINER, JR.

Department of Entomology, University of Maryland, College Park, Mary-
land 20742; present address: % Department of Entomology, NHB 169,
Smithsonian Institution, Washington, D.C. 20560.

Abstract.—Poecilocrypticus formicophilus Gebien, known from the At-
lantic coast of South America, has been collected recently in Alabama,
Florida, and Mississippi, and is reported as a species introduced to the
United States. The small, colorful beetle is apparently associated with the
imported fire ant. The species is illustrated and redescribed; known distri-
bution records and biological observations are given.

In Mississippi during February of 1978, I collected a single specimen of
a very small, colorful tenebrionid beetle which proved to be difficult to
identify; no similar specimens were in the National Museum of Natural
History collection. The species was apparently a member of the tribe Cryp-
ticini, but the only crypticine known to occur in the United States is Gon-
dwanocrypticus obsoletus (Say), a broadly oval, dull black species (Arnett,
1968). My specimen was examined by C. A. Triplehorn, who had seen one
other example of this beetle, taken in northern Florida in May of the same
year. He suspected (personal communication) that it represented an intro-
duction.

After searching the literature and examining types I was able to identify
it as Poecilocrypticus formicophilus Gebien (1928). The species was de-
scribed from southern Brazil and northern Argentina, and presumably has
been introduced to the Gulf States from South America. Three more spec-
imens were taken by me in Alabama in April of 1980. Evidence indicates
that the beetle may be associated with the imported fire ant.

The monotypic genus was described in detail by Gebien, but the general
body form and male genitalia of P. formicophilus were not figured. Since
the species apparently has become established in North America, the fol-

‘ Scientific Article No. A-3055, Contribution No. 6120 of the Maryland Agricultural Ex-
periment Station.

VOLUME 84, NUMBER 2 233

lowing illustrations and brief redescription are presented, so this distinctive
tenebrionid can be easily recognized by collectors. The key couplet below
will separate Poecilocrypticus from members of the other Western Hemi-
sphere crypticine genus, Gondwanocrypticus.

1. Body elongate oval in dorsal view, more than 2 as long as wide;
dorsum brightly colored, with pronotum orange and elytra yellow
atic! DIAC Kaede Sentient mirs, «eres ae ene aie Poecilocrypticus Gebien

— Body broadly oval in dorsal view, not more than 2 as long as wide;
dorsum usually dark and uniformly colored; if elytra maculate then
pronotum brown or black, not orange ... Gondwanocrypticus Espanol

Poecilocrypticus formicophilus Gebien

Description.—Length 2.6-2.8 mm; greatest width 1.0-1.2 mm; greatest
thickness 0.7-0.8 mm. Form elongate oval, head prognathous; dorsum
brightly colored, polished, punctate (Fig. 1).

Head dark brown to black, epistoma lighter, gular area yellowish; form
rounded, with evenly spaced setigerous punctures. Eye small, reniform.
Antenna long, reaching base of elytron if folded back along body; basal 6
segments dull yellow, nearly twice as long as wide; apical 5 segments dark
brown, thicker, nearly as wide as long. Mouthparts yellow except for dark
mandibles.

Pronotum uniformly bright reddish orange, finely margined laterally,
slightly wider than long, widest at midlength, rounded and narrowed ante-
riorly, nearly parallel sided near base; apex and base evenly truncate; dorsal
surface with scattered punctures that are smaller toward middle and each
with a short, inconspicuous seta. Scutellum orange, small, triangular, round-
ed apically, without punctures.

Elytra together as wide as pronotum, elongate, tapering to a point, with
longitudinal rows of alternating large and small punctures, each with a mi-
nute, decumbent seta; color pattern dark brown to black and yellow, with
a less distinct dark basal band, a prominent, wide, median quadrate patch
of black extending nearly to margin, and a black apical patch across suture
and extending up suture to about midlength of elytra, leaving a prominent
C-shaped area of yellow on each elytron.

Ventral surfaces punctate, yellowish orange in color except apical 2 vis-
ible abdominal sterna black; punctures very fine and dense on abdominal
sterna, with relatively long, fine setae. Legs yellow, setose; tibiae spiny and
pubescent, with 2 large, unequal apical spurs; tarsal formula 5-5-4; claws
small, simple.

Male genitalia (Figs. 2, 3) with tegmen well sclerotized; basal piece dor-
soventrally flattened, arched dorsally, widest at apical 4, rounded and
slightly asymmetrical at base, membranous ventrally. Fused parameres

234

Fig. 1.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Poecilocrypticus formicophilus, dorsal habitus.

VOLUME 84, NUMBER 2

Figs. 2, 3.

Fie te

as
cS

Ce
co,

KE
GREER

~AS oSe
aA

ates
rVSONS

Scat
MITER

235

Poecilocrypticus formicophilus, male genitalia. 2, Dorsal view. 3, Lateral view.

forming a conical apex, with scattered small punctures dorsally; basal pro-
cesses (struts) short. Median lobe membranous; internal sac laterally lined
with 2 longitudinal, dense patches of fine spicules; lining of sperm duct
granular. Female genitalia unmodified; styli short, 1 segmented, setose api-

cally.

Variation.—The elytral color pattern in P. formicophilus is quite variable,
as was noted by Gebien (1928). He figured three elytra which differed in the
extent and shape of the black areas. The form described and illustrated
above is that of the holotype, and represents an intermediate coloration in
the series of specimens available. The basal dark band varies in shape and
extent, and in one of the paratypes from Argentina the elytra are entirely

236 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

yellow at the base, and the median and apical black patches are reduced in
size. The head is reddish in this specimen also. Other specimens from the
same locality and the U.S. have a darkened stripe along the suture that
connects the basal dark band and apical patch; this stripe varies from a fine
sutural line (as in the Alabama specimens) to a wide band that blends into
the black median patch, separating the C-shaped yellow area into two quad-
rate patches (as in the specimen from Mississippi). The antennae may be
uniformly pale colored.

All four specimens in the type-series are probably males; the female spec-
imens from Florida and Alabama are slightly larger than the others, with a
more robust and rounded pronotum. In addition, the antennae in females
are shorter and less robust in proportion to body size than in males. Unlike
many other tenebrionids, however, there are no noticeable sexual differ-
ences in the front tarsi.

Figures 2 and 3 of the male genitalia were drawn from one of the paratypes
of P. formicophilus ; these structures in males from North America are iden-
tical to those of the paratype from Argentina.

Specimens examined and known distribution.—In addition to the type-
series from South America and the 5 specimens from the Gulf States, 29
other specimens found in collections of unidentified beetles were examined,
for a total of 38 specimens with the following data:

South America: ARGENTINA: Buenos Aires: Mendoza, 3 6 labeled
‘‘cotype’’; Santiago del Estero: “‘Chaco,’’ Sept. 1903 (E. R. Wagner), |
3; 1914 (E. R. Wagner), 2 6,3 2. BOLIVIA: Santa Cruz: 20 km N. Mon-
tero, 31 Dec. 1970(R. T. Allen), ‘treading,’ 8 d,6 29;4km W. Portachuelo,
17 Dec. 1970 (R. T. Allen), ‘‘cane debris,’’ 1 ¢. BRAZIL: Pernambuco:
Pery-Pery, 5 June 1892 (Gounelle), 1 2; Rio Grande do Norte: Ceara Mirim,
6-7 July 1969 (P. & P. Spangler), | 2; Rio Grande do Sul: Sao Leopoldo
(Heyer), | 6 labeled “‘type.’” PARAGUAY: Paraguari: Sapucay, Mar. (W.
T. Foster), | ¢. URUGUAY: Montevideo, 30 Aug. 1962 (Silviera-Guido),
*‘saevissima richteri nest,’ 1 ¢; 21 Dec. 1922 (F. Felippone), 1 ¢.

North America: USA: ALABAMA: Russell Co., 6 km S. Crawford, 9
April 1980 (W. E. Steiner), 1 ¢d, 1 2; 10 April 1980 (W. E. Steiner), 1 ¢.
FLORIDA: Leon Co., Tall Timbers Res. Sta., 29 May 1978 (M. Altieri),
‘‘In pitfall trap; in corn field,’’ 1 2. MISSISSIPPI: Hancock Co., Bay St.
Louis (Diamondhead), 19 Feb. 1978 (W. E. Steiner), ‘‘Pine woods; in humus
under opossum dung,” | 6.

An additional 3 specimens are labeled as intercepted at U.S. ports in
shipments from Argentina: *‘Ex Argentina, 6 May 1933, N.Y., in grapes,”’
| 5d; ‘Ex Argentina, New York, N.Y., 20 May 1940,” 1 6; ‘‘Ex Argentina,
29 May 1937, on grapes, New Orleans,’’ 1 @.

The type-specimens are deposited in the Frey Museum, Munich (Tutzing),
West Germany; other specimens are in the National Museum of Natural

VOLUME 84, NUMBER 2 237

History, Washington, D.C., the Museum National d’Histoire Naturelle,
Paris, the University of Arkansas Collection, Fayetteville, the Florida State
Collection of Arthropods, Gainesville, and in my private collection.

Remarks.—Although the species is apparently widespread, it probably is
not often collected because of its small size. It also appears to be flightless,
so it would not be taken in light traps. I suspect that the species is much
more abundant than indicated by the collection data at present; the obser-
vations on habits and habitat discussed below suggest that this beetle is
probably a common soil surface insect in open, disturbed areas and would
be expected to occur frequently in turf samples and pitfall traps.

The beetle may not be recognized as a tenebrionid by the general collec-
tor; the bright coloration, body form, and active running behavior make it
resemble a mycetophagid. Poecilocrypticus formicophilus may also be con-
fused with another tenebrionid, Alphitophagus bifasciatus (Say), a wide-
spread beetle of occasional economic importance (Triplehorn, 1965). The
markings and coloration are not as striking in the latter species, but the light
and dark banded elytra, general body form and size make it resemble P.
formicophilus. In A. bifasciatus the elytral apices and abdominal sterna are
light colored rather than black, the dorsal pubescence is more prominent,
and (in males) the clypeus is ornately sculptured.

BIOLOGY

The type-specimen from southern Brazil is pinned with two ants on cards,
labeled as *‘Prenolepis fulva Mayr.’ According to Gebien (1928), the beetles
were found under moist bark among soil partitions of an abandoned part of
an ant nest. The hindgut of the dissected paratype contained rough, dark
colored, granular material, probably soil particles.

The specimen from Montevideo, Uruguay is labeled ‘‘saevissima richteri
nest,’’ in reference to the fire ant, Solenopsis richteri Forel. A survey of
animals associated with fire ants was done by Silviera-Guido (1972), who
collected the above specimen; 28 species of Coleptera were listed but not
identified. The ant above and the closely related S. invicta Buren have been
introduced to the Gulf States from South America (Buren, 1972); Poecilo-
crypticus could have been introduced with one or both of these species of
imported fire ants. The localities where P. formicophilus has been taken
surround Mobile, Alabama, where Solenopsis spp. were supposedly first
introduced.

Two other myrmecophilous beetles were presumably introduced from
South America in this manner. Myrmecosaurus ferrugineus Bruch (Staphy-
linidae), a species described from Argentina, has been taken in fire ant nests
in Alabama, Florida, and Louisiana (Frank, 1977). Myrmecaphodius exca-
vaticollis (Blanchard) (Scarabaeidae), also from Argentina, is common
wherever fire ants occur in the Gulf States (Woodruff, 1973). These and

238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

other beetles were listed as inquilines in fire ant nests in the U.S. (Collins
and Markin, 1971) but no tenebrionids have previously been associated with
imported fire ants.

Poecilocrypticus formicophilus has not been found in close association
with Solenopsis spp. in the U.S., but fire ant mounds were very common
at the localities where I collected the beetle in Mississippi and Alabama;
some ant nests that I examined yielded Myrmecosaurus and Myrmeca-
phodius. At the Alabama site, specimens of P. formicophilus were seen
running rapidly in open, sparse turf in bright sunlight; specimens of the
related tenebrionid Gondwanocrypticus were also taken there in this situ-
ation. My single specimen of P. formicophilus from Mississippi was taken
(with some leiodids) in leaf litter beneath old remains of opossum droppings,
in a narrow strip of disturbed, secondary growth pine forest. Fire ant
mounds were located 20-30 m from each of these sites.

The beetle seems to be feeding on decaying organic plant debris. Gut
contents of the above specimens was made up of fibrous plant tissue, plant
hairs, fungus spores, and other particulate matter.

The collection data at present suggest that P. formicophilus is associated
with imported fire ants, but probably is an occasional ant nest scavenger
rather than a specialized myrmecophile, being more agile and free-living
than some other more specialized ant guests. Its immature stages are un-
known, however, and may be found to be more closely associated with ant
nests. The establishment of this beetle in North America probably will be
of no economic importance, but further study is needed to better define the
niche of this interesting species.

ACKNOWLEDGMENTS

With the generous help of Gerhard Scherer, Zoologische Sammlung des
Bayerischen Staates, Munich, I was able to examine the types of P. for-
micophilus in the Gebien collection. Theodore J. Spilman, Systematic Ento-
mology Laboratory, USDA, Washington, D.C., provided references on the
Crypticini and reviewed the manuscript. Charles A. Triplehorn, Ohio State
University, Columbus, called my attention to other specimens in collec-
tions, and also reviewed the study. Robert E. Woodruff, Florida State Col-
lection of Arthropods, Gainesville, lent the Florida specimen for my ex-
amination. Help with literature and identifications concerning fire ants was
given by David R. Smith, Systematic Entomology Laboratory, USDA,
Washington, D.C. Joseph J. Anderson, University of Maryland, College
Park, assisted in collecting in Alabama. For the kind hospitality given during
my trips to the Gulf States, I thank Standley and Phyllis Haight, Bay St.
Louis, Mississippi, and Mary Love Smith, Seale, Alabama.

VOLUME 84, NUMBER 2 239

LITERATURE CITED

Arnett, R. H., Jr. 1968. The Beetles of the United States. Am. Entomol. Inst., Ann Arbor,
Mich. 1112 pp.

Buren, W. F. 1972. Revisionary studies on the taxonomy of the imported fire ants. J. Ga.
Entomol. Soc. 7(1): 1-26.

Collins, H. L. and G. P. Markin. 1971. Inquilines and other arthropods collected from nests
of the imported fire ant, Solenopsis saevissima richteri. Ann. Entomol. Soc. Am. 64(6):
1376-1380.

Frank, J. H. 1977. Myrmecosaurus ferrugineus, an Argentinian beetle from fire ant nests in
the United States. Fla. Entomol. 60: 31-36.

Gebien, H. 1928. Uber einige Gruppen Amerikanischer Tenebrioniden (Col. Heter.). Stettin.
Entomol. Ztg. 89(1): 97-234.

Silviera-Guido, A., J. Carbonell, and C. Crisci. 1972. Animals associated with the Solenopsis
(fire ants) complex, with special reference to Labauchena daguerri. Proc. Tall Timbers
Conf. Ecol. Control Habitat Manage. 4: 41-52.

Triplehorn, C. A. 1965. Revision of Diaperini of America north of Mexico with notes on
extralimital species (Coleoptera: Tenebrionidae). Proc. U.S. Natl. Mus. 117: 349-458.

Woodruff, R. E. 1973. The scarab beetles of Florida (Coleoptera: Scarabaeidae). Part I. The
Laparosticti. Arthropods Fla. Neighboring Land Areas 8: 1-220.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 240-244

NOTES ON THE WINTER STONEFLY GENUS ALLOCAPNIA
(PLECOPTERA: CAPNITDAE)

Boris C. KONDRATIEFF AND RALPH F. KIRCHNER

(BCK) Department of Entomology, Virginia Polytechnic Institute and
State University, Blacksburg, Virginia 24061; (RFK) 5960 East Pea Ridge,
Ridgeview Apt. 1, Huntington, West Virginia 25705.

Abstract.—Allocapnia zekia Ross is shown to be only a variant of A.
wrayi Ross and is synonymized with that species. The female of A. fumosa
Ross is described for the first time from specimens collected in Virginia,
and the male of A. frisoni Ross and Ricker is reillustrated to aid other
workers in the identification of this species. Variations in habitat and male
morphology are noted for A. stannardi Ross.

All material listed and not specifically marked otherwise is in the collec-
tions of the Virginia Polytechnic Institute and State University (BCK) and
R. F. Kirchner (RFK).

Allocapnia wrayi Ross
Bigss lie

Allocapnia wrayi Ross, 1964: 170. Holotype and Allotype, Richmond, Vir-
ginia.

Allocapnia zekia Ross, 1964: 171. Holotype, Zekiah Swamp, La Plata,
Charles County, Maryland, examined. New synonymy.

Allocapnia wrayi is a common winter stonefly east of the Appalachian
Mountains ranging from Maryland to Georgia. Allocapnia zekia was de-
scribed from a single male (Ross, 1964). Ross and Ricker (1971) already
indicated that A. zekia may be only a local variant of A. wrayi. Recent
collections of Allocapnia from the type-locality of A. zekia and surrounding
areas in Maryland and throughout eastern Virginia confirm this. Characters
used to separate A. zekia from A. wrayi (‘‘seventh tergite with a complete
sclerotized basal strap and a large apical hump, and the lower limb of the
epiproct shorter and relatively deeper’’) were found to completely inter-
grade with typical wrayi populations. Several A. wrayi populations exam-
ined (Maryland and Virginia) contained individuals which varied from typ-
ical wrayi to ‘‘zekia’’ forms. Figures 1-2 illustrate some of these variations.

VOLUME 84, NUMBER 2 241

Material examined—MARYLAND: Charles Co., Zekiah Swamp Run,
Rt. 382, 28 Feb 1976, R. M. Patterson, 4 ¢, (USNM); Zekiah Swamp Run
at Rt. 5, 29 Nov 1975, R. M. Patterson #3, 15 6, 1 2, (USNM); Zekiah
Swamp Run at Rt. 6, 29 Nov 1975, R. M. Patterson #1, 15 6,5 2,(USNM):
Zekiah Swamp Run, Rt. 382, Waldorf, 14 Feb 1981, M. Firth, | ¢; Zekiah
Swamp Run, Rt. 6, Dentsville, 14 Feb 1981, BCK, 2 3; Zekiah Swamp Run,
Rt. 6, 14 Feb 1981, M. Firth, 3 ¢; Clark Run, Rt. 558, 14 Feb 1976, R. M.
Patterson, | d6, (USNM); Clark Run, 5 mi. SE of La Plata, Bel Alton-
Newtown Rd., Rt. 558, 14 Feb 1981, M. Firth, 2 ¢; Clark Run, Rt. 6, La
Plata, 14 Feb 1981, BCK, 1 3; Clark Run, Springhill-Newtown Rd., | mi.
SE of La Plata, 14 Feb 1981, BCK, | 36; Bryantown Bridge, Rt. 232, 28 Feb
1976, R. M. Patterson, 10 6, 1 2, (USNM); Dentsville Bridge, Rt. 6, 14
Feb 1976, R. M. Patterson, | ¢6 (USNM); Gilbert’s Creek, Rt. 231, 1 mi. W
of Hughesville, 14 Feb 1981, BCK, | 3; Kerrick Swamp at Rt. 488, 29 Nov
1975, R. M. Patterson, 1 6, (USNM); small stream 1.5 mi. S off Rt. 382 on
Rt 1252, 14 Feb. 1981 7BCK, | od; stream at Rts 232, 1 mi. Nob ee 15. 14
Feb 1981, M. Firth, 2 d; Calvert Co., Penny’s Run, Scientist’s Cliff at Gate
B, 15 Feb 1981, BCK, 5 ¢d,2 2. VIRGINIA: Albemarle Co., North Rivanna
River, Co. Rt. 606, off St. Rt. 29 March 1979, P. Firth, 6 6; Bedford Co.,
Big Otter River, St. Rt. 122, 7 Feb 1981, J. Despins, 1 ¢; Halifax Co., North
Fork of Aarons Creek, Co. Rt. 732 bridge, 17 March 1978, BCK, 6 6;
Aarons Creek, Co. Rt. 604, 17 March 1978, BCK, 4 ¢, 3 2; Hanover Co.,
Falling Creek, Co. Rt. 667 bridge, 25 Feb 1978, BCK, 1 6, 3 2; Stagg Creek,
Co. Rt. 696 bridge, BCK, 25 Feb 1978, 2 ¢6; Loudoun Co., small spring-fed
stream into Goose Creek, Co. Rt. 733, 15 Feb 1981, BCK, 1 6; Lunenburg
Co., Middle Meherrin River, St. Rt. 49, 17 March 1978, J. R. Voshell, |
36, 1 2; Mecklenburg Co., Butcher’s Creek, St. Rt. 92, 17 March 1978,
BECK. 2...

Allocapnia fumosa Ross
Fig. 3

Allocapnia fumosa Ross, 1964: 174; Ross and Ricker, 1971: 26.

Allocapnia fumosa has been known previously from a few localities in
the mountains of North Carolina and Tennessee. The female has remained
undescribed. Populations of this distinctive species were found in several
streams in the Mount Rogers National Recreation Area, Virginia (the highest
point in Virginia, 1746 m). Most adults were captured running about on
small trees and shrubs surrounding small spring-fed seeps flowing into first
order streams. The description and morphological terms follow those of
Ross and Ricker (1971).

Female.—Length of body 7-8 mm; wings reaching to 6th tergum. Eighth
sternum more heavily sclerotized and separate from 7th, with medial area
heavily sclerotized into a truncate process (Fig. 3).

242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-5. Terminalia of Allocapnia spp. 1, 2, A. wrayi, male, lateral view; 1, Mecklenburg
Co., Va.; 2, Calvert Co., Md. 3, A. fumosa, female, ventral view; Grayson Co., Va. 4, 5, A.
frisoni, male, Greenbrier Co., W. Va.; 4, dorsal view; 5, lateral view.

Material examined.—VIRGINIA: Grayson Co., spring seeps into Lewis
Fork, .05 mi. off Co. Rt. 603, 16 Feb 1981, BCK and RFK, 13 6,2 @ (one
pair in copulo); same, 3 April 1981, | 2; small spring-fed stream into Fox
Creek, off Co. Rt. 603, 16 Feb 1981, BCK, 1 36; Fox Creek, 2 mi. W of

VOLUME 84, NUMBER 2 243

Troutdale, Co. Rt. 603, 23 Dec 1980, RFK, 1 ¢; Lewis Fork, Co. Rt. 603,
7 Feb 1981, BCK, 1 3; same, but 27 Dec 1978, RFK, 1 6; Smyth Co., small
spring-fed stream at entrance of Grindstone Campground, Co. Rt. 603, 7
Feb 1981, BCK, 1 d, 1 2; Washington Co., Straight Branch of Big Laurel
Creek, 4 mi. E of Damascus, St. Rt. 58, 25 Dec 1977, RFK, 1 ¢.

Allocapnia frisoni Ross and Ricker
Figs. 4, 5

Allocapnia frisoni Ross and Ricker, 1964: 90. Holotype and Allotype, Ev-
ansville, West Virginia; Ross and Ricker, 1971: 30.

The description of A. frisoni is adequate. However, the illustration in
Ross and Ricker (1971, Fig. 25b) is not representative of the rugosity of the
dorsal process on the eighth tergum. Apparently the extent and shape of the
dorsal rugosities were not completely illustrated. This has caused some
problems with the identification of this species. We present additional illus-
trations (Figs. 4, 5) to assist other workers in identifying A. frisoni. Para-
types were examined from Tompkins Co., New York.

Material examined.—VIRGINIA: Rockingham Co., Kephart Run, St. Rt.
33, W of Harrisonburg, 24 Dec 1977, P. K. Powell, 4 ¢; Rocky Run, St.
Rt. 33, W of Harrisonburg, 24 Dec 1977, P. K. Powell, 3 ¢. WEST VIR-
GINIA: Logan Co., Frogtown Hollow of Copperas Mine Fork, Holden, 27
Jan 1975S, RFK 43 ¢, 26 2; same but 11 Dec 1975,8 6,6 2; same but 1
March 1976, 37 6, 20 2; Fayette Co., Big Hollow of Paint Creek, 2 mi. NW
of Kingston, WV-Turnpike, 22 Dec 1979, RFK, 4 d, 4 2; same but 24 Jan
1980, 5 3o, 1 2; same but 1 Feb 1981, 61 6, 52 9; Greenbmer Co., Rocky
Run, Forest Rd. 139, Blue Bend Rec. Area, 29 Dec 1980, BCK and RFK,
5 6; Meadow Creek, Co. Rt. 14, near Rucker Gap, 29 Dec 1980, BCK and
RFK, 10 6.

Allocapnia stannardi Ross

Allocapnia stannardi Ross, 1964: 174. Holotype and Allotype, Sevier
Co., Tennessee; Ross and Ricker, 1971: 42.

Allocapnia stannardi Ross was collected at two locations in Virginia. The
habitat of both localities vary from the description of Ross and Ricker
(1971). Three males and one female were collected from small spring-seeps
in Patrick County, Virginia, (BCK) at an altitude of only 580 m. Nearly 200
males and females were collected in Smyth County, Virginia, (BCK and
RFK), also from spring seeps and from a nearby small head water stream
at an altitude of 1060 m. This species is found in a broader range of habitats
and at lower elevations than mentioned by Ross and Ricker (1971). Several
males from the second collection site had small processes on the sixth ter-
gum and others had the process of the seventh tergum slanted distad rather
than erect.

244 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Both A. fumosa and A. stannardi represent new range extensions and
new state records for Virginia (Kondratieff and Voshell, 1979).

ACKNOWLEDGMENTS

We thank John D. Unzicker of the Illinois Natural History Survey, Cham-
paign, for loan of the holotype of A. zekia and Oliver S. Flint, Jr., National
Museum of Natural History, Washington, D.C., (USNM) for loan of para-
types of A. frisoni and additional specimens of A. wrayi. Penelope F. Kon-
dratieff is gratefully acknowledged for the illustrations. We also thank Mar-
tin J. Firth, Port Republic, Maryland, for helping with the Maryland
collections.

LITERATURE CITED

Kondratieff, B. C. and J. R. Voshell, Jr. 1979. A checklist of the stoneflies (Plecoptera) of
Virginia. Entomol. News 90: 241-246.

Ross, H. H. 1964. New species of winter stoneflies of the genus Allocapnia (Plecoptera,
Capniidae). Entomol. News 75: 169-177.

Ross, H. H. and W. E. Ricker. 1964. New species of winter stoneflies, genus Allocapnia

(Plecoptera: Capniidae). Trans. Ill. Acad. Sci. 57: 88-93.

. 1971. The classification, evolution, and dispersal of the winter stonefly genus Allo-

capnia. Ill. Biol. Monogr. 45: 1-166.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 245-249

THE FOUR “‘OCELLI’’ OF THE ISOMETOPINE GENUS
ISOMETOCORIS CARVALHO AND SAILER
(HEMIPTERA: MIRIDAE)

THOMAS J. HENRY AND J. MALDONADO CAPRILES

(TJH) Systematic Entomology Laboratory, IIBIII, Agricultural Research
Service, USDA, % National Museum of Natural History, Washington, D.C.
20560; (JMC) Department of Anatomy, Ponce School of Medicine, G.P.O.
Box 7004, Ponce, Puerto Rico 00731.

Abstract.—lsometocoris blantoni Carvalho and Sailer (Hemiptera: Miri-
dae), described as having four ocelli and previously placed in the subfamily
Isometopinae, is re-examined and transferred to the subfamily Psallopinae
on the basis of pronotal, hemelytral, and pretarsal characters. This is the
first report of Psallopinae in the Western Hemisphere. Electron microscopy
is used to show that J. blantoni does not have true ocelli.

Carvalho and Sailer (1954) described the attractive species /sometocoris
blantoni from a single male collected in Panama. On the basis of four minute
‘“‘ocelli’’ located on the vertex (Figs. 1, 2), they placed this genus in the
Isometopidae (now a subfamily of the Miridae), a family characterized by
two distinct ocelli. In reference to this peculiar condition, they commented,
‘‘Only with the discovery of additional specimens will it be possible to
determine with certainty whether this specimen is a freak or whether it is
a representative example of the species.”

We recently discovered seven additional specimens of blantoni in the
collection of the National Museum of Natural History and the personal
collection of the second author. Because blantoni was described from a
single male, our original interest was to present new distribution records
and a description of the female of this unusual species. Closer examination,
however, provided new information regarding the presence of the four ocel-
li, a condition not known to occur in any other group of insects (unless the
‘‘ocelli’’ of certain male sternorrhynchus Homoptera are considered).

The holotype of blantoni exhibits four tiny, symmetrically placed, shiny
spots on the vertex. When viewed with a stereomicroscope, these *‘ocelli”’
are, at best, difficult to see; their pattern, suggestive of more than a mere
random placement of small shiny spots, was mistaken for true ocelli.

246 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1, 2. Line drawings indicating position of “‘ocelli’’ on vertex of Isometocoris blantoni
(redrawn from Carvalho and Sailer, 1954). 1, Frontal view of head. 2, Dorsal view of head and
pronotum.

We first doubted that these structures were true ocelli when we discov-
ered that all of our specimens possessed additional similarly developed spots
between the compound eyes (Figs. 3—5) and on the pronotum (Fig. 4). Elec-
tron microscopy revealed that the 4 spots originally described as ocelli for
blantoni are not ocelli but are actually enlarged, pigmented setigerous
cavities (Figs. 3, 5) common to several parts of the insect’s dorsum. When
a seta is broken or lost, the remaining cavity (Fig. 5) appears as a small
polished spot or ‘‘ocellus’’ when viewed at magnifications of 100x or less.
Carvalho and Sailer’s (1954) drawings are reproduced here (Figs. 1, 2) to
allow comparison with our electron micrographs (Figs. 3, 4). Arrows on our
figures indicate corresponding setal bases shown in Carvalho and Sailer’s
line drawings.

The lack of ocelli in this genus requires that Isometocoris be removed
from the subfamily Isometopinae and from Henry’s (1980) key to the New
World genera. Further examination of the claws (Fig. 6), tarsi, and anterior
margin of the pronotum (Fig. 4) indicates that [sometocoris belongs in the
Old World subfamily Psallopinae.

Schuh (1976) defined Psallopinae based on the only included genus, Psal-
lops Usinger. Most previous workers had put this genus in the subfamily
Phylinae, although Carvalho (1956), in describing Psallops occulatus, point-
ed out that Psallops has all the facies of the subfamily Isometopinae (then
considered a family), but the lack of ocelli placed the genus outside this
group. He also noted that the genitalia of Psallops were entirely different
from those of the Phylinae. Eyles (1972), following Carvalho, transferred
this genus to Isometopidae. Schuh (1976) defined Psallops as having a fine
upturned anterior pronotal margin, a simple male vesica without spicules,
subapical claw teeth, one or two cells in the membrane of the forewings,
2-segmented tarsi, and nine metafemoral trichobothria.

VOLUME 84, NUMBER 2 247

Figs. 3-6. Micrographs of Isometocoris blantoni. 3, Frontal view of head (x250); arrows
indicate position of false ocelli. 4, Head and pronotum showing numerous ocelli-like setal bases
(<x 250); arrows indicate position of false ocelli. 5, High magnification (2500) of the setal base
indicated by arrow in upper right corner of Figs. 3 and 4. 6, Claw showing subapical teeth
(2500).

Isometocoris exhibits the finely upturned and narrow anterior margin of
the pronotum (Fig. 4), simple male genitalia, a subapical tooth on each claw
(Fig. 6), a single closed cell in the membrane of each forewing, and 2-seg-
mented tarsi. This combination of characters dictates that /sometocoris be

248 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

transferred to Psallopinae; thus it becomes the first member of this subfamily
to be recognized in the Western Hemisphere.

Isometocoris is separated from Psallops by the broader vertex that is
equal to the dorsal width of an eye and by the strongly concave basal margin
of the pronotum. In Psallops, the eyes cover much of the dorsal surface of
the head, the vertex is less than % the dorsal width of an eye, and the base
of the pronotum is nearly straight.

Isometocoris blantoni Carvalho and Sailer

Diagnosis.—Female, length 1.74-1.76 mm, width 0.90-0.94 mm. Head:
Width 0.46-0.48 mm, vertex 0.22 mm. Rostrum: Length 0.76 mm, reaching
beyond base of ovipositor. Antenna: Segment I, length 0.08 mm; I, 0.38
mm; III, 0.18 mm; IV, broken. Pronotum: Length 0.20-0.22 mm; basal
width 0.74-0.76 mm.

General coloration of dorsum pallid, set with numerous brown setigerous
pores; head pallid, becoming somewhat darker toward frons, set with nu-
merous brown setigerous pores, including 4 symmetrically positioned ones
on vertex (Figs. 3, 4); pronotum pallid with a few dark blotches on disc and
at lateral angles; hemelytra pallid with a large orange blotch near middle of
clavus and corium, on apical % of corium and on apical %4 of embolium;
inner angle of cuneus fuscous; hemelytra with numerous brown setigerous
spots that become more reddish on cunei and apical ¥% of coria; membrane
opaque white, tinged with brown; membrane with a single closed cell and
distal spurious vein. Venter fuscous, meso- and metapleura tinged with pink
or red. Ostiolar evaporative area brownish except for a pale or whitish
winglike plate arising from apex of ostiolar channel. Legs brown, meta-
femora becoming fuscous. Setae of body simple, long pallid and semierect,
nearly all arising from darkened bases; eyes thickly pubescent.

Diagnosis.—Male, length 1.72-1.74 mm, width 0.86 mm. Head: Width
0.52 mm, vertex 0.20 mm. Rostrum: Length 0.80 mm, reaching near 5th
abdominal segment. Antenna: Segment I, length 0.08 mm; II, 0.44 mm; III,
0.16 mm; IV, broken. Pronotum: Length 0.20 mm; basal width 0.70-0.72
mm.

Males are very similar to females in markings and coloration, but the head
and pronotum are more strongly infuscated and the eyes are proportionately
larger and more deeply faceted.

Specimens examined.—Holotype ¢, Pedroesa, Republic of Panama, Oct.
28, 1953, F. S. Blanton coll., USNM type no. 61950; 2 ¢, Pt. Aguadulce,
R. P., June 21, 1952, F. S. Blanton coll. (USNM); 1 6, Mojinga Swamp,
Canal Zone, Panama, Jan. 9, 1953; F. S. Blanton coll. (USNM); 1 6, 2
?, Panama, Lt. Barro Colorado Isl., Jan.-Apr.—June, 1972, D. Engleman
coll. (Maldonado colln.); 1 2, Panama, Lt. Coco Solo, Canal Zone, July
1973, D. Engleman coll. (Maldonado colln.).

VOLUME 84, NUMBER 2 249

LITERATURE CITED

Carvalho, J. C. M. 1956. Heteroptera: Miridae. Insects Micronesia 7(1): 1-100.

Carvalho, J. C. M. and R. I. Sailer. 1954. A remarkable new genus and species of Isometopidae
from Panama (Hemiptera: Isometopidae). Entomol. News 65: 85-88.

Eyles, A. C. 1972. Supplement to list of Isometopidae (Heteroptera: Cimicoidea). N.Z. J. Sci.
15: 463-464.

Henry, T. J. 1980. Review of Lidopus Gibson and Wetmorea McAtee and Malloch, descrip-
tions of three new genera and two new species, and key to New World genera (Hemip-
tera: Miridae: Isometopinae). Proc. Entomol. Soc. Wash. 82: 178-194.

Schuh, R. T. 1976. Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of
relationships within the family. Am. Mus. Novit. 2601, 39 pp.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 250-255

AN OLD WORLD SPECIES OF SCYMNUS (PULLUS)
ESTABLISHED IN PENNSYLVANIA AND NEW YORK
(COLEOPTERA: COCCINELLIDAE)

RoBERT D. GORDON

Systematic Entomology Laboratory, IIBUI, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—A European species of Coccinellidae, Scymnus (Pullus) su-
turalis Thunberg, is reported for the first time from North America. All
collection localities are in Pennsylvania and New York. The known distri-
bution, possible origins of the introduction, and taxonomic affinities of S.
suturalis are discussed.

In a taxonomic treatment of certain genera of North American Scymnini
(Gordon, 1976), I discussed Scymnus (Pullus) coniferarum Crotch. The dis-
tribution of this species is primarily Californian with outlying records from
British Columbia, South Dakota, and Colorado (Fig. 1) (this map is repro-
duced as published in 1976). I also included in the treatment of this species
data from specimens taken from several Pennsylvania localities with the
speculation that they had been artificially distributed on nursery stock. Sub-
sequent investigation has revealed that all of the Pennsylvania specimens
are Scymnus (Pullus) suturalis Thunberg, a species not previously recorded
from North America. A single New York record (1981) was received from
A. G. Wheeler. Korschefsky (1931) gives the distribution of S. suturalis as
‘‘Europa, Caucasus, Sibirien, Tunis.’’ Scymnus suturalis and S. conifera-
rum are extremely similar in appearance, with the primary diagnostic dif-
ference being the shape of the infundibulum in the female genitalia. This
and other differences are discussed below.

Scymnus (Pullus) suturalis Thunberg
Scymnus (Pullus) suturalis Thunberg, 1975: 106.

Description as for Scymnus (Pullus) coniferarum except: Body slightly
broader, less elongate in appearance; punctures on elytron coarse, dense,
separated by the diameter of a puncture or less; basal lobe of male genitalia
broad in ventral view, abruptly narrowed in apical 4 (Fig. 2), apex in lateral

VOLUME 84, NUMBER 2

‘(961

) piuBalAsuudagd Ul SI/Danins “g pu ROLAWY YON UWaysomM Ul uinapsafiu0d SnuUUAIE JO VONNGLISIG

i |

Figs. 2-6. Scymnus suturalis. 2-5, Male genitalia. 6, Female genitalia.

view distinctly bent downward (Fig. 3), apex of sipho S-shaped (Figs. 4, 5);
female genitalia with infundibulum slender, rodlike (Fig. 6).

Scymnus coniferarum differs from Scymnus suturalis in being extremely
elongate (Fig. 7), with elytral punctures small and separated by about twice

VOLUME 84, NUMBER 2

te
nn
es)

A,

Figs. 7-12. Scymnus coniferarum. 7, Habitus view. 8-11, Male genitalia. 12, Female gen-

italia.

Fig. 13. Known distribution of Scymnus suturalis in Pennsylvania (1981).

the diameter of a puncture; the male genitalia are different as in Figs. 8-11,
and the female infundibulum is short and thickened at both ends (Fig. 12).

The known distribution of S. suturalis in Pennsylvania is depicted in Fig.
13. However, it also occurs in New York, and it is not likely that the species
is restricted to those states. All specimens were received from A. G. Wheel-
er and T. J. Henry of the Pennsylvania Department of Agriculture and were
taken mostly on pine (Pinus sylvestris, P. strobus, P. resinosa). One record
was from spruce (Picea pungens). Nearly all records were from trees used
as ornamentals such as ornamental plantings, nursery stock, or Christmas
tree plantations.

Both S. suturalis and S. coniferarum are aphid predators and apparently
feed mainly or entirely on aphids infesting conifers. Gordon (1976) recorded
S. coniferarum on lodgepole pine and Monterey pine infested with woolly
aphids. I have no specific predation records for §. suturalis, but its food
preference is apparently very similar to that of S. coniferarum.

Because the shipment of two- and three-needled pines into this country
has been prohibited by quarantine laws since 1915 (Wheeler and Henry,
1973), S. suturalis was probably established prior to that year. Before 1915
(prior to current quarantine laws), vast amounts of live nursery stock were
imported from Europe and distributed throughout Pennsylvania and else-
where (personal communication, A. G. Wheeler). Under those conditions
S. suturalis had many opportunities to become established.

VOLUME 84, NUMBER 2 255

Collection localities for S. suturalis. —PENNSYLVANIA: (20 specimens
examined) (Allegheny Co.) Cheswick, Pittsburgh; (Berks Co.) Shillington;
(Bucks Co.) Coopersburg; (Butler Co.) Herman; (Centre Co.) State College;
(Chester Co.) Longwood Gardens; (Clarion Co.) Rimersburg; (Columbia
Co.) Berwick Cemetery; (Crawford Co.) Conneaut Lake; (Cumberland Co.)
Camp Hill, Hagestown; (Dauphin Co.) Hershey; (Erie Co.) Fair View; (In-
diana Co.) Indiana, Shelocta; (Mercer Co.) Sheakleyville; (Mongomery Co.)
Norristown; (Northampton Co.) Easton; (Washington Co.) Marianna, Squir-
rel Hill Nursery. NEW YORK: (2 specimens examined) (Tompkins Co.)
Ithaca.

ACKNOWLEDGMENTS

I thank A. G. Wheeler, Jr., Pennsylvania Department of Agriculture,
Harrisburg, and T. J. Henry, now with the Systematic Entomology Labo-
ratory, USDA, Washington, D.C., for help in locating specimens and dis-
cussing the probable reasons for the establishment of suturalis in Pennsyl-
vania and New York.

LITERATURE CITED

Gordon, R. D. 1976. The Scymnini (Coleoptera: Coccinellidae) of the United States and Can-
ada: Key to genera and revision of Scymnus, Nephus and Diomus. Bull. Buffalo Soc.
Nat. Sci. 28: 1-362.

Korschefsky, R. 1931. Coleopterorum catalogus. Coccinellidae I, pars 118. 224 pp.

Thunberg, C. P. 1795. Dissertatio entomologica sistens Insecta Suecica. Pp. 105-113. Upsaliae.

Wheeler, A. G., Jr. and T. J. Henry. 1973. Camptozygum aequale (Villers), a pine-feeding
mirid new to North America (Hemiptera: Miridae). Proc. Entomol. Soc. Wash. 75:
240-246.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 256-269

A KEY TO THE FEMALES OF THE GENUS
OXYETHIRA (TRICHOPTERA: HYDROPTILIDAE)
FROM THE SOUTHERN UNITED STATES!

RoBERT W. KELLEY AND JOHN C. MORSE

Department of Entomology, Fisheries and Wildlife, Clemson University,
Clemson, South Carolina, 29631; Graduate Research Assistant (RWK) and
Associate Professor (JCM).

Abstract.—Associations of females with the males for most species of the
genus Oxyethira known from the southern United States has made possible
the construction of a key to females of that region. Illustrations for 24 of
the 27 species are provided along with a discussion of the general mor-
phology of the genital segments of females of the genus. Three new syn-
onyms are proposed: O. barnstoni Harper is a synonym of O. mirabilis
Morton, O. allosi Blickle is a synonym of O. dualis Morton, and O. cirrifera
Flint is a synonym of O. arizona Ross.

Microcaddisflies (Hydroptilidae) are among the most poorly known Tri-
choptera, primarily due to their small size. Blickle (1979) provided a key to
the males of North American species of the family, but females of few
species have been associated with corresponding males (see Table 1). Pub-
lished illustrations for most of these females are insufficiently detailed to
allow comparisons adequate for species discrminations. Furthermore, the
majority of species have remained unassociated.

This paper provides illustrations and a key for 24 of the 27 species of
Oxyethira that are known from across the southern half of the continental
United States. A list of these species and their known geographical ranges
are provided in Table 1. No females were available of three species; O.
coercens Morton, O. leonensis Kelley, and O. setosa Denning. Oxyethira
coercens is known primarily from the northeastern United States and the
latter two from the Florida panhandle and southern Georgia.

Characters of both external and internal morphology of the genital seg-
ments are used in the key, such that specimens must be cleared in KOH in

' Technical contribution no. 1916 of the South Carolina Agricultural Experimental Station,
Clemson University.

VOLUME 84, NUMBER 2 257

Table 1. Species of Oxyethira from the southern United States.

Known Distribution

Piedmont Previous

; Moun- or Coastal Descriptions
Species States tain Plateau Plain of Female
O. abacatia Denning lalnn (Gils Sa Ce X X X None
O. aculea Ross Ariz., N. Mex., X X None
Okla.
O. arizona Ross Ariz. X Flint, 1968a
O. azteca (Mosely) exe Xx Flint, 1968b
O. coercens Morton Okla., Va. X None
O. dualis Morton Ark., Ariz., N. x Ross, 1944
Mex., Tenn.,
exe as
O. dunbartonensis Galas. X X Kelley, 1981
Kelley
O. elerobi (Blickle) RlatansnG: X Kelley, 1981
O. florida Denning Fla. X Botosaneanu, 1979
O. forcipata Mosely GaryNiGs,. S:€e, X X None
Va.
O. glasa (Ross) Fla., Ga., La., X None
Okla., S.C.
O. grisea Betten Si@wlenn...Via- X X None
O. janella Denning late Gal lear. X X X Flint, 1968a
Miss., S.C.
O. leonensis Kelley Fla. X None
O. lumosa Ross Flas Ga s:C€: X X None
O. maya Denning Fla., Ga. X None
O. michiganensis Mosely Ga., S.C., Va. X None
O. novasota Ross Plas. Gas. lay, X None (‘‘novasota”’
Miss., S.C., of Ross, 1944, is
exe actually verna)
O. pallida (Banks) ING, [EIB (Gels. x x ax Ross, 1944
La., Miss.,
Okla., S.C.
O. rivicola Blickle and Renn eaVa- X None
Morse
O. rossi Blickle and Tenn. X None
Morse
O. serrata Ross Tenn. X Ross, 1944
O. setosa Denning Fla., Ga. X None
O. sininsigne Kelley Blas. leas. 9: X Kelley, 1981
O. ulmeri (Mosely) Tex. x None
O. verna Ross Blan iba S.C.5 X Denning, 1947;
Tex: Ross, 1944 (as
““novasota’’)
O. zeronia Ross Blay,.Ga., Ia... X X X None
IN; Ge s.G;,
Tenn., Va.

a —— —————

258 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Species groups of Oxyethira in North and Central America (adapted from Mar-
shall, 1979).

nnn nnn ee eae aE Eye a a yn SSE

Azteca Group
azteca (Mosely)
janella Denning
quelinda Botosaneanu
puertoricensis Flint
Bidentata Group
abacatia Denning
aeola Ross
anabola Blickle
mirabilis Morton
barnstoni Harper, new synonym

Distinctella Group
araya Ross
serrata Ross
setosa Denning
Dualis Group
dualis Morton
allosi Blickle, new synonym
sininsigne Kelley
Falcata Group!
rossi Blickle and Morse

Forcipata Group

forcipata Mosely
michiganensis Mosely
obtatus Denning

' Primarily Palearctic.

Grisea Group

allagashensis Blickle
coercens Morton
dunbartonensis Kelley
grisea Betten

lumosa Ross

novasota Ross

rivicola Blickle and Morse
sida Blickle and Morse

Pallida Group

alaluz Botosaneanu
arizona Ross

cirrifera Flint, new synonym
campesina Botosaneanu
maya Denning
pallida (Banks)
verna Ross

Ulmeri Group
aculea Ross
florida Denning
simulatrix Flint
ulmeri (Mosely)

Zeronia Group

glasa (Ross)
jamaicensis Flint
leonensis Kelley
zeronia Ross

Incertae sedis
elerobi (Blickle)

order to see the distinguishing characters of many of these species. Details
of the clearing procedure were provided by Ross (1944). Morphological
nomenclature is based upon the work of Nielsen (1980). Species groups
recognized herein (Table 2) have been modified from those of Marshall
(1979). Voucher specimens are in the collections mentioned in Acknowl-
edgments and in the Clemson University Insect Museum.

Three synonyms were discovered in the course of this research. Exami-
nation of specimens of O. mirabilis Morton from the British Museum (Nat-
ural History) showed them to be identical with O. barnstoni Harper. Type-
specimens of both O. arizona Ross and O. cirrifera Flint were studied and
found to be synonyms. Blickle (1980) did not mention O. dualis Morton in

VOLUME 84, NUMBER 2 259

Fig. |. Generalized views of genital segments of Oxyethira females. A, Left lateral view.
B, Left dorsolateral oblique view of internal sclerites. C, Left ventrolateral oblique view
of internal sclerites. a = apicomesal spur of segment VI; b = segment VII; c = tergum VIII;
d = apodeme of tergum VIII; e = sternum VIII; f = tergum IX; g = tergum X: h = cercus;
i = spermathecal sclerite; } = spermathecal process; k = foramen of spermathecal process; | =
horizontal lamella; m = collaterial duct; n = venter of oviduct; o = posterior ring sclerite; p =
spermathecal duct.

his diagnosis of O. allosi Blickle. A review of the illustrations of each
species reveals their synonymy.

GENERAL MORPHOLOGY

Nielsen (1980) reviewed the internal and external anatomy of the genital
segments of female Trichoptera, including the microcaddisfly genera Agray-
lea, Hydroptila and Orthotrichia. Although similar in general features to
these genera, several modifications can be noted in Oxyethira (Fig. 1). In
females of most species of the genus, segment VIII is incomplete with a
short tergum. Laterally the segment narrows into apodemes which proceed
anteriorly 4% to % the length of segment VII. Ventrally, segment VIII is
represented by a short sternum which is sclerotized to various degrees in
different species. In some species, sclerites are present on sternum VIII and
still others have sternum VIII retracted within segment VII. Segment IX is

260 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

reduced to a short, lightly sclerotized tergum. Tergum X is more heavily
sclerotized with a pair of cerci distally.

Several sclerites are associated with the internal reproductive system
(Figs. 1A—C). The spermathecal sclerite is similar to that of other Trichop-
tera with a spermathecal process originating anterioventrally. A keyhole-
shaped foramen can be seen ventrally in the spermathecal process through
which the spermathecal duct enters dorsally. The sclerotized dorsum of the
oviduct, which Nielsen called the horizontal lamella, is prominent in most
Oxyethira females. In many species the posteriolateral corners of this scler-
ite continue posterioventrally to encircle the oviduct and fuse ventrally,
forming the floor of the oviduct. Immediately posterior to the horizontal
lamella the collaterial duct enters the dorsum of the oviduct. Further pos-
teriorly another sclerite nearly encircles the oviduct, although it is reduced
to a membranous structure in many species. Nielsen did not mention a
homolog for this sclerite in other hydroptilids; thus, we refer to it here as
the “‘posterior ring sclerite.”’ Species of the azteca group bear sclerotized
processes supporting the oviduct ventrally which do not appear to be pres-
ent in other species groups (Figs. 2A, C).

KEY To THE FEMALE OXYETHIRA FROM THE SOUTHERN UNITED STATES

1. Pair of sclerotized parallel rods proceeding anterioventrally from

spermathecal sclerite, nearly reaching sternum VII (Figs. 2A, C) ..
wages syd s Sremsgabiey = ered wareatk a ogee PAPC eRe eee janella Denning

— Pair of rods shorter or lacking beyond spermathecal sclerite (Figs.

BAns ©) isctiachet. sencurmach os SSeurts Ae obeabie ac.b colic Serena 2
2. Sternum VIII with 1| or 2 distinct sclerites ventrally (Figs. 3A, C;

BAC) aos tise 2 eye sie ed iedd ow 006.5 a9 COR eee te 3
— Sternum VIII without ventral sclerites, retracted within segment

VII, or entire sternum moderately sclerotized (Figs. 22A, C) ...... 6

3. Sternum VIII with ventral paired sclerites which are congruent
anteriorly, diverging posteriorly (Figs. 3A, C); apico-mesal process

lackine-fromrstemum Vii... 2k eee ee elerobi (Blickle)
— Sternum VIII with single sclerite (Figs. 4C; 6C); apico-mesal pro-
cess: present*on sternum! VI fee: )2o58/9) i Gea ee ee 4

4. Sternite VIII broad, as wide as, or wider than, long (Figs. 4C; 5C)

— Sternite VIII small, longer than wide, trapezoidal (Figs. 6A, C) ...
Salo A RPE A. ea rae ee dunbartonensis Kelley

5. Apodemal rods of segment VIII proceeding anteriorly as far as, or
farther than, spermathecal sclerite (Figs. 4A, B) ..... novasota Ross

— Apodemal rods of segment VIII not proceeding as far anteriorly as
spermathecalsclerite (Figs: SAVB)" 22 eee grisea Betten

VOLUME 84, NUMBER 2 261

+

Figs. 2-5. Terminal abdominal segments of Oxyethira spp., females. 2, O. janella. 3, O.

elerobi. 4, O. novasota. 5, O. grisea. A, Lateral. B, Dorsal. C, Ventral.

262

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
. Segment VII infolded laterally (Figs. 7A, C; 8A, C; 9A, C) ........ |
Segment VII not infolded laterally (Figs. JOA, C) ................ 9
. Tergum IX elongate (Fig. 7A); distal end of tergum VIII nipple-
shaped jin dorsal-view (Fig. 7B) 2g eeeee eee arizona Ross
Tergum IX shorter (Figs. 8A; 9A); distal end of tergum VIII broadly
rounded or truncate in dorsal view (Figs. 8B; 9B) ................ 8

. Spermathecal sclerite broadly rounded anteriorly (Fig. 8C); terga

VIII and IX flattened dorsally in lateral view (Fig. 8A) .........
Pie ae dee ean 68 a SENS ee OS «TE SL eae ae oe maya Denning
Spermathecal sclerite more acutely rounded anteriorly (Fig. 9C);
tergum VIII with knoblike process dorso-distally and tergum IX
proceeding ventrally from tergum VIII in lateral view (Fig. 9A) ...
Rept PEAR oe ans sol aha th as: Ace Se pallida (Banks)

. Tergum VIII large, longer than wide; tergum IX reduced; tergum

Mheavily sclerotized: (Figs 10A.sB)atase Steet eee glasa (Ross)
Tergum VIII small, wider than long; tergum X usually lightly scler-
etized) (Fics: 11B; 13B).3.6 Sear nts. bo Ie et eee 10
Posterior margin of sternum XII sinuate, with tongue-shaped pro-
cess in ventral view (Fig. 11C) .......... rivicola Blickle and Morse
Posterior margin of sternum VII not both sinuate and tongue-
shaped. (Fis7*14C) 220 hek a ice eee a en eee 11

. Sclerotized process proceeding ventrally from spermathecal scler-

ite, appearing distally forked in lateral and ventral views (Figs.

LP AMC) rare ie, oh RE. ee ee anes azteca (Mosely)
No such sclerite proceeding ventrally from spermathecal sclerite
(Ig PASAY 55.2055. iS. Bee SE. Se ee nr 2

. Posterior end of oviduct floor with paired sclerites (Figs. 13 A,C);

paired, lightly sclerotized tergites on tergum X (Fig. !3B) .....
alist tee BMlek Ns BECUASE ee A ae eine rossi Blickle and Morse

Oviduct floor without paired sclerites; sclerotized tergites usually

absent from tergum) X:(Bie wiS@) ia... 1s. Joe oe 13

. Terga VIII, IX and X and sternum VIII largely retracted within

segment VII (Figs. 14A—C); apodemes of segment VIII nearly par-
allel in dorsal view and much nearer each other than to lateral walls

Ofsegment, Vil (Rig. (4B) ee ee ee michiganensis Mosely
Terminal segments exserted in normal position, apodemes of VIII
much farther apart (Figs 17A):..05 5 ee ee 14

. Spermathecal sclerite visible as distinct oval structure in ventral

view and set anteriorly to horizontal lamella (Figs. 15A, C) .....

PS EI tan Bieh ives Ops, w idx WOR, cig ce et Re eee sininsigne Kelley
Spermathecal sclerite not anterior to horizontal lamella and may or
may not /be:oval (Fig. 14C). 4.24 b.4. 2e9e eee 15

VOLUME 84, NUMBER 2 263

Figs. 6-9. Terminal abdominal segments of Oxyethira spp., females. 6, O. dunbartonensis.
7, O. arizona. 8, O. maya. 9, O. pallida. A, Lateral. B, Dorsal. C, Ventral.

264 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

13B

Figs. 10-13. Terminal abdominal segments of Oxyethira spp., females. 10, O. glasa. 11,

O. rivicola. 12, O. azteca. 13, O. rossi. A. Lateral. B, Dorsal. C, Ventral.

VOLUME 84, NUMBER 2

265

Figs. 14-17. Terminal abdominal segments of Oxyethira spp., females. 14, O. michiganen-
sis. 15, O. sininsigne. 16, O. dualis. 17, O. abacatia. A, Lateral. B, Dorsal. C, Ventral.

266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 18-21. Terminal abdominal segments of Oxyethira spp., females. 18, O. florida. 19,
O. aculea. 20, O. zeronia. 21, O. verna. A, Lateral. B, Dorsal, C. Ventral.

——— EOE

VOLUME 84, NUMBER 2 267

Figs. 22-25. Terminal abdominal segments of Oxyethira spp., females. 22, O. ulmeri. 23,
O. serrata. 24, O. forcipata. 25, O. lumosa. A, Lateral. B, Dorsal. C, Ventral.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

. Spermathecal sclerite extending anteriorly at least 74 the length of

segment VII; tergum VIII broadly rounded dorsally in lateral view
(Pigs AIGA IPA) 2... Fass «+o. cafes 2 lee oleae ees ete earn 16
Spermathecal sclerite extending anteriorly less than 12 the length
of segment VIII; tergum VIII with bulbous process dorsally (Figs.
244: 25A).or acutely pointed (Figs. 20AR 21°): .8 Sa oa eee 17

. Spermathecal sclerite triangular in ventral view (Fig. 16C); apo-

demes of segment VIII shorter than segment dorsally (Figs. 16A,
2 Se Ai a ener en aac CP Mas. SAK TAES dualis Morton
Spermathecal sclerite more nearly diamond-shaped in ventral view
(Fig. 17C); apodemes of segment VIII long, much longer than seg-

ment dorsallva¢Higs.. .17A; 'C) o2.2.. .. - Seas oe abacatia Denning
. Floor of oviduct with sclerotized area attenuated posteriorly (Figs.

its | OM LC C) eh ne acre Sree. Aor. 5 18

Sclerotized area on floor of oviduct broader posteriorly (Figs. 20C;

QUO) oi NS oe AN Bae oh Moe ols Cae HOR AGS a 20

. Sternum VIII extending anteriorly into segment VII to encircle

oviduct and fuse dorsally (Figs. 18A; 19A); attenuate sclerite con-
tinuing posteriorly as spinelike process (Figs. 18C; 19C); tergum
VIII rounded or pointed posterio-dorsally (Figs. 18A; 19A) ....... 19
Sternum VIII not extending into segment VII (Figs. 22 A,C); at-
tenuate sclerite not continuing posteriorly as spinelike process; ter-
gum VIII with truncate process in lateral view (Fig. 22A) .......

bg 1 ae en i. NP Seaete e ulmeri (Mosely)

. Tergum X moderately excised distally in dorsal view (Figs. 18B,

©) 0 Fics EB oo 8 Fo dow Bids wguhiike Oa ee florida Denning
Tergum X only slightly excised distally in dorsal view (Figs. 19B,
Cr pS Pie Pais Be os. 5c. 6a As ee oe eee aculea Ross

. Apico-mesal process absent from venter VI or very minute; dorso-

distal end of tergum VIII angulate in lateral view, dropping sharply

posteriorly (Figs:.20AG 2 WA), Gi ae ons oes ee eee 4)
Apico-mesal process present on venter VI (Fig. 1A); dorso-distal
end. of tergum VILL bulbous se5.3.....\. 2.3 6 Seed. 0 eee 22

. Tergum X with two sclerites dorsally (Fig. 20B); floor of sperma-

theca sclerotized only posteriorly (Figs. 20A, C); apex of tergum
VIII angled at approximately 90° in lateral view (Fig. 21A) ......
Sb is Oe ee Ae Ae, Sik Sn Sy ch, so as ata zeronia Ross
Tergum X only lightly sclerotized in a single plate (Fig. 21B); floor
of spermatheca completely sclerotized, oval in ventral view (Figs.
21A, C); apex of tergum VIII protruding posterio-dorsally, acutely
angled m lateral view (Fig::21/A) 2. ssquuh, 4). Us etn verna Ross

. Tergum X widening distally in dorsal view (Figs. 23B, C); terga

Vi bs and -X ‘distinct (Figs... 23A/B)) ee eee serrata Ross

VOLUME 84, NUMBER 2 269

— Tergum X narrowing distally in dorsal view; terga VIII, [IX and X
nomelcagly. distmct (Figs. 244A. B: 254. :B)scc. 5205 eet l. ooo eepes

23. Distal end of tergum VIII not proceeding posteriorly as far as an-
terior end of tergum X in dorsal view (Figs. 24A, B) ...........

i IEICE EAC thie STAT dines SOE hs See forcipata Mosely
— Distal end of tergum VIII proceeding posteriorly as far as anterior
end: ofgtereum~exn( Bigs: 25A By o28 idee 26h. de cles lumosa Ross

ACKNOWLEDGMENTS

We express our appreciation to O. S. Flint, Jr., of the National Museum
of Natural History, Washington, D.C., David C. Etnier of the University
of Tennessee, Knoxville, D. G. Denning of Moraga, California, and Peter
C. Barnard of the British Museum (Natural History), London, for the loan
of specimens that helped make this research possible. A portion of this work
was funded as part of a baseline study for the U.S. Department of Energy's
Savannah River National Environmental Research Park, Aiken, South Car-
olina.

LITERATURE CITED

Blickle, R. L. 1979. Hydroptilidae (Trichoptera) of America north of Mexico. Bull. N.H.

Agric. Exp. Stn. 509: 1-97.

. 1980. A new Oxyethira (Hydroptilidae, Trichoptera) of the aeola Ross group; with a

key to separate the five males of the group. Pan-Pac. Entomol. 56(2): 101-104.

Botosaneanu, L. 1979. The caddis flies (Trichoptera) of Cuba and of Isla de Pines. Stud. Fauna
Curacao Other Caribb. Isl. 59(185): 1-62.

Denning, D. G. 1947. Hydroptilidae (Trichoptera) from southern United States. Can. Entomol.
79: 12-20.

Flint, O. S. 1968a. The caddisflies of Jamaica (Trichoptera). Bull. Inst. Jam. Sci. Ser. 19: 1-68.

. 1968b. Bredin-Archbold Smithsonian Biological Survey of Dominica, 9. The Trichop-

tera (caddisflies) of the Lesser Antilles. Proc. U.S. Natl. Mus. 125(3665): 1-86.

Kelley, R. W. 1981. New species of Oxyethira (Trichoptera: Hydroptilidae) from the south-
eastern United States. J. Ga. Entomol. Soc. 16(3): 368-375.

Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera). Bull. Br.
Mus. (Nat. Hist.) Entomol. 39(3): 135-239.

Nielsen, A. 1980. A comparative study of the genital segments and the genital chamber of
female Trichoptera. Biol. Skr. Dan. Vid. Selsk. 23(1): 1-200.

Ross, H. H. 1944. The caddisflies, or Trichoptera, of Illinois. Ill. Nat. Hist. Surv. Bull. 23:
1-236.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 270-276

MARILOPTERYX CARANCARUA, A NEW GENUS
AND NEW SPECIES FROM EAST TEXAS
(LEPIDOPTERA: NOCTUIDAE: HADENINAE)

ANDRE BLANCHARD AND JOHN G. FRANCLEMONT

(AB) 3023 Underwood Street, Houston, Texas 77025; (JGF) Department
of Entomology, Comstock Hall, Cornell University, Ithaca, New York
14853.

Abstract.—A new genus and new species of hadenine Noctuidae, Mari-
lopteryx carancahua, is described from East Texas. The genus includes one
other species occurring in the United States, lutina (Smith), new combi-
nation, from Florida, and a few tropical American species, all of which,
except /amptera (Druce), new combination, are apparently undescribed.

The new species described here is one of the very striking moths that
have been collected in recent years in Texas; its description is to be attrib-
uted to both authors. The new genus has been in manuscript by the junior
author for a long time, and its description is to be attributed to him.

Marilopteryx Franclemont, New Genus

Type-species.—A grotis lamptera Druce, 1890, new combination.

Diagnosis.—A genus in the large American complex that includes Erio-
pyga Guenée, 1852, Chabuata Walker, 1857 [1858], Orthodes Guenée, 1852,
and relatives. Hampson, 1905 (vol. 5, p. 313) placed /amptera in Eriopyga
to which there is some superficial similarity, but he gave it a section by
itself. The genitalia of both sexes, especially those of the male, are strikingly
different from those of Eriopyga. The males of Marilopteryx have the gen-
italia of a more or less normal configuration although with some special
modifications; those of the males of Eriopyga have the valves much less
sclerotized, extended laterally, and deeply cleft. The females of Marilop-
teryx have the sternum of the seventh abdominal segment highly modified
and heavily sclerotized; those of Eriopyga have the sternum of the seventh
abdominal segment little or not modified and no more heavily sclerotized
than those of the other segments.

Description.—Head with vestiture deep, front rounded, ridge of scales
above antennae, male antenna pubescent, with fine, short, closely set setae

VOLUME 84, NUMBER 2 271

of more or less equal length, female antenna with shorter pubescence, eye
with hairs over entire surface, palpus with Ist segment short, 2nd segment
moderate, upcurved to about middle of eye, 3rd segment short, porrect,
haustellum well developed, strong; thorax with vestiture of hairlike scales
and narrow, forked scales, a prominent median, longitudinal ridge from
behind patagia, venation as figured by Hampson (1905, text fig. 70, p. 313),
forewing with R, from beyond middle of discal cell, R, from near apex of
accessory cell, R; and R, short stalked from apex of accessory cell, R;
connate with stalk of R; + Ry, M, from upper angle of discal cell, hindwing
with Rs and M, connate from upper angle of discal cell, M. obsolescent, M,
and Cu, connate from lower angle of discal cell, discocellular veins more or
less erect, legs of male with large tufts on femora and tibiae, especially the
latter, tarsi with 3 rows of spines beneath, abdomen with vestiture of long
hairlike scales overlying flat scales, male with evident lateral tufts on seg-
ments | through 6, dorsal tufts on segments | through 3, conspicuous caudal
tufts, last sternum with large, spreading tuft from pocket, Ist sternum mod-
ified, usually with lateral hair-pencil on each side, female without tufts ex-
cept those on dorsum, last sternum, 7th segment, highly modified, heavily
sclerotized.

Male genitalia—The species agree in basic structure with carancahua
but with various individual modifications of tegumen, uncus, and aedoeagus.
(The type of the genus, /Jamptera, is most similar to carancahua.)

Female genitalia.—The species agree with carancahua, differing in mod-
ifications of 7th abdominal sternum, sclerotization of ductus bursae, shape
of bursa, and shape of appendix bursae.

Remarks.—The species of the genus are mostly dull reddish or reddish-
brown moths with some grayish overtones; the new species described here
has most of the examples grayish with a pink tint, but there are some red-
dish-brown ones, thus the name Marilopteryx (ember + wing).

In addition to the type-species and the new species the genus contains
Leucania lutina Smith, 1902, new combination (L. velutina Smith, 1900,
preoccupied), placed in Tricholita by McDunnough (1938: 73) and three or
four undescribed species from the American tropics.

Marilopteryx carancahua Blanchard and Franclemont, New Species
Figs. 1-6

Diagnosis.—This species is most closely related to /amptera and an ap-
parently undescribed species from Panama. It differs from /amptera in many
obvious characters of the male genitalia. The expansion of the penicillus of
the tegumen is highly rugose in carancahua; it is not rugose in lamptera.
The upper lobe of the vinculum of /amptera is widely expanded, but that
of carancahua is not. The cuculli of the valves of carancahua are two times
the width of those of the valves of /Jamptera; the expanded lobe of the right

272 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

valve of /amptera is ligula-like in shape, not racquet shaped as is that of
carancahua, the lobe of the left valve of /Jamptera is two times as wide as
that of carancahua, and lamptera has an additional small, rounded lobe at
the angulation of the cucullus with the longitudinal axis of each valve, but
in carancahua in addition to the absence of the small lobe, the angulation
of the cucullus is absent, and the cucullus is on the longitudinal axis of each
valve. The vesica of the aedoeagus of /amptera has a single cornutus, not
two cornuti as in carancahua. The male genitalia of the undescribed species
from Panama have the rugose lobes of the penicilli smaller, the lobes of the
valves much different in shape, and the uncus with two long projections.
The only other species, /utina, occurring in the United States differs from
carancahua in having the male genitalia with very small cuculli, very small
lobes on the valves, and a narrow uncus. The females of carancahua, lutina,
and lamptera have the seventh abdominal sternum of the same basic con-
firmation; that of carancahua is more or less quadrate with the apical ex-
cavation broad; that of /utina is somewhat oval, flattened at the base, and
with the excavation narrow; and that of /amptera has the lobes prolonged
and the excavation broad. The ductus bursae of /utina and that of lamptera
are longer and narrower than that of carancahua; the appendix bursae of
lutina is much longer than those of carancahua and lamptera; lutina has a
separate, longitudinal, sclerotized plate near the origin of the appendix bur-
sae. The female of the Panamanian species is not known to us. Superficially
carancahua is unique in the possession of the conspicuous black spot in the
subterminal line and the somewhat sinuous, slightly irregular antemedial
and post medial lines which in all the other species, except the Panamanian,
are straight from costa to inner margin.

Description.—Head, patagia, disc of thorax, and tegulae concolorous with
ground color of forewing, palpus with Ist and 2nd segments reddish black,
upper margin of 2nd segment and all of 3rd segment light tan, all segments
with some reddish scales, behind, below, and in front of eye black, pale
above; forewing varying from ashy, olive gray with just a faint pinkish tinge
to dark vinaceous red, color uniform on any one specimen, all steps in
variation present from extremes, basal, antemedial, and postmedial lines
double, filling between darker lines ground color or slightly paler, basal line
traceable from costa to cubital vein, outer of 2 lines often faint, antemedial
line crenulate, forming 3 loops, black dots on vague outer line on Cu and
2nd A, postmedial line slightly excurved to M, then incurved to inner mar-
gin, vague outer line followed by black points on veins M, through 2nd A,
often with an indication of one on R;, median shade often obsolescent,
dusky, angled from costa to inner side of reniform then straight to inner
margin, subterminal line irregularly wavy, paler than ground, often with
considerable white scaling, conspicuous black spot on inner side of subter-
minal line between veins M, and Mg, often with smaller black spots between

VOLUME 84, NUMBER 2 273

Figs. 1, 2. Marilopteryx carancahua. 1, Holotype, Deutchburg, Jackson Co., Texas, 6

March 1975, A. & M. E. Blanchard; collection USNM. 2, Male genitalia with aedoeagus
removed, aedoeagus with vesica everted at right; genitalia slide AB 3618.

274 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 3-6. Marilopteryx carancahua. 3, Male abdomen, 8th abdominal segment, sternum
(right) and tergum (left); genitalia slide AB 3618. 4, Female genitalia, 7th abdominal segment
in place; genitalia slide AB 3279. 5, Female genitalia, 7th abdominal segment removed; genitalia
slide AB 3620. 6, Female, 7th abdominal segment, modified sternum; genitalia slide AB 3620.

VOLUME 84, NUMBER 2 075

veins M, and M; and between veins R; and M,, sometimes other small black
spots toward costa, terminal line a series of small black dots in the inter-
spaces between the veins connected by a very fine, dentate, black line,
fringe concolorous with ground, reniform and orbicular large, with incon-
spicuous, pale annuli, reniform barely darker than ground; orbicular no
darker than ground, often not discernible; hindwing much less variable than
forewing, vinaceous gray, heavily infuscate toward outer margin, veins con-
trasting, blackish, discal dot moderate, fringe pinkish; dorsum of abdomen
tending to agree in color with ground of forewing, with some darker scales
and hairlike scales, terminal and lateral tufts of male with a decided reddish
tint. Beneath: forewing dark gray, paler toward extreme base, a conspicuous
postmedial line and discal spot, veins pale scaled beyond postmedial line;
hindwing sordid, pinkish gray, darker beyond conspicuous, dark, punctate
postmedial line, discal spot punctiform, dark, fringe of both wings much as
above, of hindwing less contrasting.

Expanse.—Male, 36.5-40.5 mm, female, 35.4-41.4 mm; forewing length
20.5—23 mm.

Male genitalia.—As figured; Fig. 3 illustrates sclerotized areas of tergum
and sternum of 8th abdominal segment, sternum, on right, shows pocket
(dark) from which large tuft arises.

Female genitalia—As figured; Fig. 4 illustrates genitalia with 7th sternum
in place; Fig. 5 illustrates genitalia with sternum removed, sclerotization of
ductus bursae visible, ligulate plate, excavated at apex, is on back (upper)
wall of ductus; Fig. 6 illustrates the modified 7th sternum, a more or less
quadrate ventral plate with wide apical excavation and 2 apically pointed,
more dorsad, lateral plates.

Holotype.—Male (Fig. 1). Deutchburg [near Carancahua Creek], Jackson
County, Texas, 6 March 1975, A. & M. E. Blanchard. In the National
Museum of Natural History, Washington, D.C.

Paratypes—45 do, 10 2; 54: Deutchburg, Jackson County, Texas, 14
March 1975 (1), 18 March 1974 (8), 3 March 1975 (18), 6 March 1975 (26),
15 March 1977 (1); 1: Eagle Lake, Colorado County, Texas, 27 April 1978;
A. & M. E. Blanchard. Paratypes are in the following collections: American
Museum of Natural History, New York (2), British Museum (Natural His-
tory), London (2), Canadian National Collection, Ottawa (2), National Mu-
seum of Natural History (27), A. Blanchard (12), E. C. Knudson (2), J. G.
Franclemont (8).

ACKNOWLEDGMENTS

We wish to thank R. W. Poole for reading the manuscript and E. C.
Knudson for numerous kindnesses.
The photographs are by the senior author.

276 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

LITERATURE CITED

Druce, H. 1890. Descriptions of new species of Lepidoptera Heterocera from Central and
South America. Proc. Zool. Soc. (London), 1890: 493-520, pl 42 & 43.

Hampson, G. F. 1905. Catalogue of the Lepidoptera Phalaenae in the British Museum. Volume
5; xvi + 634 pp. Trustees of the British Museum (Natural History), London.

McDunnough, J. H. 1938. Check List of the Lepidoptera of Canada and the United States of
America, Part 1, Macrolepidoptera. Mem. South Calif. Acad. Sci. Vol. 1, 174 pp.

Smith, J. B. 1900. A hundred new moths of the family Noctuidae. Proc. U.S. Natl. Mus. 22:

413-495.

. 1902. Contributions toward a monograph of the Lepidopterous family Noctuidae of

Boreal North America. A revision of the moths referred to the genus Leucania, with

descriptions of new species. Proc. U.S. Natl. Mus. 25: 159-209.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 277-280

OVOVIVIPARITY IN THE BLATTELLID COCKROACH,
SYMPLOCE BIMACULATA (GERSTAECKER)
(DICTYOPTERA: BLATTARIA: BLATTELLIDAE)

Louis M. RotH

U.S. Army Natick Research and Development Laboratories, Natick,
Massachusetts 01760; send correspondence to 81 Brush Hill Road, Sher-
born, Massachusetts 01770.

Abstract.—The only cockroach genera considered to be ovoviviparous
belong to the Blaberidae. However, the African blattellid, Symploce_ bi-
maculata (Gerstaecker), is ovoviviparous. Apparently ovoviviparity arose
independently in both the Blattellidae and Blaberidae. Symploce capitata
(Saussure) is a new synonym of S. pallens (Stephens).

According to McKittrick (1964), cockroaches have evolved along two
phyletic lines, the superfamilies Blaberoidea and Blattoidea. Species in the
former group evolved ovoviviparity and viviparity, whereas the Blattoidea
remained oviparous. McKittrick’s hypothesis is supported by considerable
biological evidence (Roth, 1970).

During the course of evolution of ovoviviparity in the Blattaria, the 00-
theca changed from a hard, rigid egg case that was dropped shortly after its
formation, to a soft, flexible, reduced structure which was retracted into a
uterus or brood sac. Such changes have taken place in the Blaberoidea
(Polyphagidae, Blattellidae, and Blaberidae), which evolved internal incu-
bation, but not in the Blattoidea (Cryptocercidae and Blattidae) (Roth,
1968a, 1971).

Because species of Blattella carry their odthecae externally for about the
entire embryogenetic period, Roth and Willis (1958) suggested that the Blar-
tella type of oviposition behavior is an intermediate stage between oviparous
and ovoviviparous forms. Two other blattellid genera, Chorisia and Ony-
cholobus have similar odthecae and oviposition behavior (Roth, 1968a,
1971). After studying the ovarioles of various species of Blattaria, I (Roth,
1968b, 1970) suggested two possible pathways within the Blattellidae that
led to ovoviviparity in the Blaberidae. At that time, the available evidence
indicated that only the Blaberidae are ovoviviparous or viviparous [only
Diploptera punctata (Eschscholtz) is known to be viviparous, but presum-
ably the other species of Diploptera are also}.

278 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1 y Imm A 0.1mm

04mm

Figs. 1-5. Symploce bimaculata. 1, Abdomen (dorsal) of a pregnant female showing the
end of the dotheca (arrow) protruding slightly beyond the supraanal plate. 2, Embryo removed
from an ootheca that was carried internally in the brood sac. 3, Odtheca removed from the
uterus. 4, Part of the keel region of an o6theca showing the absence of respiratory tubules and
calcium oxalate crystals. 5, Spongy bodies in the keel region of the odtheca. Figures 2, 4, and
5 are chitin preparations.

Recently while working on a taxonomic revision of the blattellid genera
Blattella and Symploce, | found several female specimens of the African
Symploce bimaculata (Gerstaecker) that had odthecae internally in a uterus
(Fig. 1). The egg cases occupy practically the entire abdominal cavity, as
they do in pregnant blaberids. Shelford (1910) stated that the odtheca of S.

VOLUME 84, NUMBER 2 279

BLABEROIDEA
Viviparity

Diplopterinae

had

Haun Ovoviviparity | 2

Ovoviviparity 9 subfamilies 2 ~
Blattellinae

BLATTOIDEA

Oviparity
Blattidae
Cryptocercidae

Oviparity
Blattellinae and
4 other subfamilies

Blattellidae

Oviparity
Polyphagidae

Fig. 6. Phylogenetic tree of the Blattaria (abbreviated and modified from McKittrick, 1964),
showing the evolution of ovoviviparity and viviparity.

bimaculata is similar to that of Blattella germanica (L.), but he did not
indicate that it is carried internally.

Two of the odthecae of bimaculata contained 38 and 40 eggs, which, in
the latter, contained developing embryos (Fig. 2). Superficially the odtheca
resembles that of species of Blattella but is more transparent throughout
(Fig. 3; however, the portion near the end of the female’s body is slightly
darker than the remainder of the egg case), and its keel is further reduced
and lacks respiratory tubules (Fig. 4). The spongy bodies in the interior of
the seam (Fig. 5) are still present and are similar to those of Blattella ger-
manica (Lawson, 1951). Calcium oxalate crystals are completely absent
from the walls of the egg case. The odthecae of various species of Blattella
may have a few calcium oxalate crystals, or none at all (Roth, 1968a). That
Symploce bimaculata incubates its eggs internally shows that ovoviviparity
actually has evolved independently in at least two families of the Blaberoi-
dea, the Blattellidae and Blaberidae (Fig. 6).

Symploce pallens (Stephens) [=Symploce capitata (Saussure), new
synonymy] is oviparous and deposits its odtheca shortly after its formation.
The hard, rigid o6thecae of Symploce ruficollis (F.) (reported as Symploce

280 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

bilabiata Rehn and Hebard), and Symploce jamaicana Rehn, both of which
contain large amounts of calcium oxalate crystals, indicate that these species
also are oviparous (Roth, 1968a). Several species of African Symploce have
oothecae and oviposition behavior similar to species of Blattella, but these
actually belong to the latter genus (unpublished data). That Symploce bi-
maculata is ovoviviparous suggests that it is not a Symploce, and based on
male characters. I believe, should be removed from that genus, which I
have done elsewhere (Roth, in press).

There are about 209 genera and about 1740 species of Blattellidae, and
approximately 155 genera, and about 1020 species of Blaberidae, The ovi-
position behavior of relatively few genera and species in these two families
is known. Probably other genera of blattellids will be found that are ovo-
viviparous. I (Roth, 1970: 79) stated that ‘It would be safe to say that any
species which carries its egg case internally is a blaberid .. . .”’ I should
have known better than to be so dogmatic, and I would not be too surprised
if oviparous blaberids are eventually found.

ACKNOWLEDGMENT

I thank Inge Persson of the Swedish Museum of Natural History, Stock-
holm, for the series of specimens of Symploce bimaculata.

LITERATURE CITED

Lawson, F. A. 1951. Structural features of the odthecae of certain species of cockroaches
(Orthoptera, Blattidae). Ann. Entomol. Soc. Am. 44: 269-285.

McKittrick, A. F. 1964. Evolutionary studies of cockroaches. Cornell Univ. Agric. Exp. Stn.
Mem., No. 389, 197 pp.

Roth, L. M. 1968a. Odthecae of the Blattaria. Ann. Entomol. Soc. Am. 61: 83-111.

. 1968b. Ovarioles of the Blattaria. Ann. Entomol. Soc. Am. 61: 132-140.

. 1970. Evolution and taxonomic significance of reproduction in Blattaria. Annu. Rev.

Entomol. 15: 75-96.

. 1971. Additions to the odthecae, uricose glands, ovarioles, and tergal glands of Blat-

taria. Ann. Entomol. Soc. Am. 64: 127-141.

. In press. Stayella, a new genus of ovoviviparous blattellid cockroaches from Africa

(Dictyoptera: Blattaria: Blattellidae). Entomol. Scand.

Roth, L. M. and E. R. Willis. 1958. An analysis of oviparity and viviparity in the Blattaria.
Trans. Am. Entomol. Soc. 83: 221-238.

Shelford, R. 1910. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition
nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deutsch-Os-
tafrikas 1905-1906, Dr. Yngve Sjdstedt. 17. Orthoptera. 2. Blattodea. Konigl. Schwed.
Akad. Wissen. (1907), 3 (17.2): 13-48.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 281-296

A REVISION OF EUSCHISTUS DALLAS SUBGENUS
LYCIPTA STAL (HEMIPTERA: PENTATOMIDAE)

L. H. RoLsTon

Department of Entomology, Louisiana Agricultural Experiment Station,
Center for Agricultural Sciences and Rural Development, Louisiana State
University, Baton Rouge, Louisiana 70803.

Abstract.—The subgenus Lycipta Stal of Euschistus Dallas is redefined
and five species, one of them new, are added to it. Euschistus (Lycipta)
machadus, n. sp., is described, E. (L.) luridus (Dallas) is transferred from
Agroecus, and E. aceratos Berg is placed in the synonymy of this species.
A key is given for species of the subgenus.

The subgenus Lycipta of the genus Euschistus contains ten South Amer-
ican species from the southern temperate zone and bordering tropics. One
species, E. cornutus Dallas, seems to be the most abundant South American
member of the genus.

Stal (1862a) proposed Lycipta as a genus to contain three species which
earlier (1860) he had placed in Euschistus; but in his work on the Hemiptera
of Mexico, which also appeared in 1862, he demoted Lycipta to subgeneric
rank by describing E. (Lycipta) spurculus. A decade later Stal (1872) cor-
rected his initial placement of E. spurculus by relocating this species in the
nominate subgenus. At the same time he retrieved one of the species, now
Agroecus scabricornis (Herrich-Schaffer), from those that he originally
placed in Lycipta and included this species among the “‘species incerti ge-
neris.’’ Also, he added three more species to the subgenus Lycipta. From
then until now Lycipta has included only the five South American species
assigned to it by Stal: E. triangulator (Herrich-Schaffer), E. illotus Stal, E.
cornutus (Dallas), E. cribrarius Stal, and E. picticornis (Stal). Of the ten
species under consideration, only the above five were known to Stal.

Five additional species are here added to the subgenus: E. circumfusus
Berg, E. imitator Berg, E. luridus (Dallas), E. machadus, n. sp., and E.
sharpi Bergroth. Euschistus luridus is transferred from Agroecus and is a
senior synonym of Euschistus aceratos Berg.

282 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Euschistus subgenus Lycipta Stal

Lycipta Stal, 1862a: 58.
Euschistus (Lycipta): Stal, 1862b: 100; Stal, 1872: 23; Kirkaldy, 1909:
XXIX.

Type-species.—Cimex triangulator Herrich-Schaffer, 1842: 95, fig. 667
(by subsequent designation, Kirkaldy, 1909: X XIX).

Diagnosis.—Lateral walls of genital cup consisting in part of inflatable
membrane located just beneath rim of cup and turning ventrad near anterior
wall of cup (Fig. 11). Penisfilum coiled (Fig. 12). Thecal processes incrassate
basally (Fig. 13). Spermatheca with secondary dilation at base of proximal
flange; duct bent rather abruptly at proximal end of primary dilation, often
with funnel shaped pigmented area enclosing base of dilation and bend of
duct (Fig. 15). Apex of head incised between each jugum and tylus (Fig. 7).

Comments.—The partially membranous lateral walls of the genital cup,
which appear as a white cushion when inflated, are apparently unique within
Euschistus. The secondary dilation of the spermatheca basad of the proxi-
mal flange may also be unique to Lycipta, although the spermatheca of many
Euschistus species has not been examined. However, the incisions in the
apex of the head, one between each jugum and the tylus, also occur in at
least some specimens of E. quickua Piran and E. rufimanus Stal.

The spermatheca of eight of the ten species of Lycipta were examined,
but no females of E. cribrarius or E. sharpi were available for dissection.

The subgenus is remarkable in that it contains three species pairs, each
pair with virtually identical genitalia but with non-genitalic differences that
are apparently consistent. While such genitalic similarity among species
arouses suspicion that the forms may represent disjunct variation of a single
species, a comparable situation exists within the servus group of Euschistus
species in North America, and in the latter instance genetic isolation, al-
though incomplete, has been demonstrated (Sailer, 1954). It would be de-
sirable to subject the members of these three species pairs to tests for ge-
netic isolation. Until such testing is done the status of the form most recently
named in each pair is likely to remain in doubt. The species involved are:
luridus and cribrarius; circumfusus and imitator; and triangulator and pic-
ticornis.

KEY TO SPECIES OF SUBGENUS LYCIPTA

1. | Venation in membrane of hemelytra simple or branched with only
1 or 2 closed cells at most; male with conspicuous tuft of setae at
posterolateral corners of genital cup in addition to scattered setae
(Five 2) eines. 02. dee lung ee ee ee 2
_ Venation in membrane of hemelytra reticulate or if not scutellum

VOLUME 84, NUMBER 2 283

bearing small black macule on each side of apex; setal tufts in-

CUMS PICUONS) OF ADSM be Fr ri a in vis tg giao te ee Oe a ee 6
2(1). Three irregular black vittae on abdominal venter, | median and

1 on each side; veins in membrane of hemelytra with many short

Bi anene SiN OM)) 2 Ye oh sce ak oh eee ee illotus Stal
- Abdominal venter lacking vittae; veins in membrane of hemelytra

Spaescivabtamchem! cP. Fide. « wids's seed enalc als CARRE gon ee 3
3(2). Margins of connexiva alternated, pale with black macules at trans-

WE GRC EGUIMLINE See sont Shea s See. sus SRT oe Oe machadus, new species

Connexiva withoutmacies) .<s.. 26's 4s sesh ae a ac) eee ee 4

4(3). Humeral angles spinose, turned obliquely forward (Fig. 16)
Sh > SIGE RRO A cee en I rere aie Goer es? imitator Berg

- iumeralsangles: OOtUSE: 375. f.o...4:. se hee ae ee ee 5
5(4). Denticles on anterolateral margin of pronotum red or yellow; con-
nexiva entirely dark or with yellow margin ...... circumfusus Berg
~ Denticles black; connexiva entirely pale .......... sharpi Bergroth
6(1). Apex of scutellum covered with large pale spot ................. ff
— Apex of scutellum not strongly differentiated by color ........... 8
7(6). Last 2 antennal segments predominately reddish; apex of scutel-
lum usually somewhat reflexed ..... triangulator (Herrich-Schaffer)
- Last 2 antennal segments predominately blackish; apex of scu-
Tellmanomnehexediud. wiaty.tn. kusedaiahed nae <r picticornis Stal

8(6). Humeri cornute, apically obtuse, projecting anterolaterad about
as far forward as base of head (Fig. 33); apex of scutellum with

Smalliplack mark omeach:side .: ..ks.caeasercoes we cornutus Dallas
- Humeri not cornute; apex of scutellum uniformly colored ........ 9
9(8). Humeri produced laterad of base of coria by width of eye or less,

OOUUSEN sant tonto. wow ethane) heels welaieainiaeee luridus (Dallas)
~ Humeri produced laterad of base of coria by about 1.5 width of

evertacuie yiiyriqtot. se Seek vals Cael. pice ee cribrarius Stal

Euschistus (Lycipta) illotus Stal
Figs. 1-6

Euschistus illotus Stal, 1860: 19.
Lycipta illotus: Stal, 1862a: 58 (listed).
Euschistus (Lycipta) illotus: Stal, 1872: 24 (keyed, descriptive note).

Veins in hemelytral membranes with many short branches but forming
few or no closed cells (Fig. 1). Three irregular vittae present on abdominal
venter, | median (sometimes tenuous) and 2 lateral, formed by black
patches. Gonocoxae 2 not evident (Fig. 6). Emargination in posterior py-
gophoral margin broad and moderately deep from caudal view (Fig. 4),

284 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-6. Euschistus (L.) illotus. 1, Membrane of right hemelytron. 2, Genital cup, proc-
tiger and parameres removed; tuft of setae (st); tumescence (tu). 3, Paramere. 4, Posterior
pygophoral margin, caudal view. 5, Same, ventral view. 6, Genital plates, caudoventral view.
Dimensional lines equal 0.5 mm.

sinuous from both ventral and dorsal views (Figs. 2, 5). Cup of paramere
densely setose, apex minutely ridged (Fig. 3).
Distribution.—Argentina (Misiones); Brazil (Espirito Santo, Mato Gros-
so, Sao Paulo, Santa Catarina, Rio de Janeiro); Paraguay; Bolivia.
Comment.—The holotype was examined.

Euschistus (Lycipta) machadus Rolston, New SPECIES
Figs. 7-15

Light yellowish brown above, becoming reddish yellow on humeri; punc-
tures rather small and evenly distributed, black in clusters on each side of
calloused spot at posterior margin of each cicatrice, along anterolateral pro-
notal margins, especially on anterior humeral margins, and at posterior mar-
gin of ocelli, concolorous on humeral apex, otherwise dark brown to fus-
cous; each side of scutellar apex with black spot. Ventral surfaces as dark
as or a little darker than dorsum; punctation fuscous to black, very dense
on abdomen. Length of body 10.1-11.5 mm.

Width and length of head subequal, 2.1-2.4 mm wide at eyes, lateral

VOLUME 84, NUMBER 2 285

i/

Figs. 7-15. Euschistus (L.) machadus. 7, Head and pronotum. 8, Genital plates; caudo-
ventral view; basal plates (bp); second gonocoxae (gx 2). 9, Posterior margin of pygophore,
caudal view. 10, Same, ventral view. 11, Genital cup, parameres and proctiger removed; in-
ferior ridge (ir); tuft of setae (st); tumescence (tu). 12, Aedeagus, lateral view; conjunctival
appendage (a); penisfilum (p); median penal lobes (mpl). 13, Aedeagus, dorsal view; thecal
processes (tp). 14, Paramere. 15, Spermatheca; proximal flange (pf); dilation of spermathecal
duct (d).

286 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

margins subparallel for middle 4% of distance from eyes to apex; juga and
tylus subequal in length, separated at apex by shallow incision between each
(Fig. 7). Basal antennal segment pale with black spots usually coalescing
into streak on dorsal and ventral surfaces, remaining segments black except
basal '/, of last and usually basal '/, of 4th pale; basal segment reaching apex
of head; length of segments 0.6—0.7; 0.9-1.1; 1.0-1.1; 1.2-1.5; 1.4-1.7 mm.
Bucculae subtruncate at base of head, surpassing slightly basal segment of
rostrum.

Pronotum 2.3—2.7 mm long at meson, 2.7—3.0 times as wide at humeri.
Anterolateral margins concavely arcuate, denticulate onto humeri; denticles
obtuse, well separated from each other, yellowish brown. Humeral angles
produced moderately laterad, slightly cephalad, a little elevated; apex nar-
rowly rounded. Disk slightly rugose; cluster of black punctures laterad of
calloused spot at posterior border of each cicatrice larger than mesal cluster,
subcircular, sometimes subfoveate; mesial cluster linear, rarely lacking.

Scutellar width and length subequal, 3.6—4.2 mm across base; fovea in
basal angles moderately large and deep, black; apex slightly depressed be-
tween marginal black spot on each side, sometimes weakly emarginate.
Membrane of hemelytra hyaline; venation light brown, usually with a few
closed cells. Connexivum moderately exposed, concolorous with corium
excepting black marginal spot on each side of segmental sutures and be-
tween these a smaller black spot on protruding apical angle of sternite.

Evaporative areas small for genus, extending about 0.4 distance from
ostiole to lateral margin of metapleuron; sparingly black punctate. Large
black spots on femora and tibiae, many coalescing on femora; tarsi fuscous
to black usually excepting superior face of basal segment. Apical angles of
abdominal sternites produced into small obtuse tubercle, black at apex;
basal angles with larger black spot. Spiracles concolorous with or darker
than supporting sternum.

Broad emargination in posterior margin of pygophore irregularly concave
from both caudal and ventral views, shallowly so from dorsal view (Figs.
9, 10, 11). Inferior ridge entirely exposed from caudal view. Large tubercle
on each side on posterior border of genital cup bearing dense tuft of setae;
large pale tumescence on lateral walls of genital cup paralleling rim for most
its length, bending abruptly ventrad anteriorly. Parameres terminating in
rather thin simple hook (Fig. 14). Conjunctiva with 2 appendages on each
side; apex of appendage nearest median penal lobes subacute, sclerotized;
other appendage hyaline (Fig. 12). Penisfilum coiled, making about 2 com-
plete loops. Median penal lobes projecting dorsad at apex. Lateral lobes of
theca moderately developed, median lobe obscure.

Holotype.—¢, labeled Brazil, Sao Paulo, Serra Bocaina, S. J. Barreiro,
1650 m. (b) Oct.-Nov. 1969, Alvarenga and Seabra. Deposited in American
Museum of Natural History, New York.

VOLUME 84, NUMBER 2 287

Figs. 16-21. Euschistus (L.) imitator. 16, Humeral angle. 17, Posterior pygophoral margin,
caudal view. 18, Same, ventral view. 19, Genital cup, parameres and proctiger removed; tuft
of setae (st); tumescence (tu). 20, Paramere. 21, Genital plates, caudoventral view; basal plates
(bp); 9th paratergites (pt 9); 2nd gonocoxae (gx 2).

Paratypes.—10 ¢, 23 2. Same data as holotype (¢6, 2 LHR; 6, 2 UNLP;
36, 2 USNM; ¢, 2? NR; 6, 2 BMNH; 6, 8 2 AMNH); same data as
holotype except Nov. 1968, M. Alvarenga (¢ TAMU; 6, 2 LHR; 6 ¢
AMNH) same data as holotype except Jan. 1969, M. Alvarenga (2 AMNH);
Italiaya, 1100 m, E. do Rio, 30-XI-42, W. Zikan (6 RNH); “‘Itatiaia,”’ 1960
m, N. B. Fagundes, 1-933 (2 RNH); “‘Itatiaia,”’ N. B. Fagundes, 1-933
(6 RNH); Bananal, Bocaina, S. Paulo, D. Mendes, 1-937 (2 RNH).

Deposition of paratypes——American Museum of Natural History, New
York (AMNH); British Museum (Natural History), London (BMNH); au-
thor’s collection (LHR); Naturhistoriska Riksmuseet, Stockholm, Sweden
(NR); Rijksmuseum van Naturlijke Historie, Leiden, Netherlands (RNH);
Texas A&M University, College Station (TAMU); Universidad Nacional de

288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

22

Figs. 22, 23. Euschistus (L.) circumfusus. 22, Humerus and anterolateral margin of the
pronotum. 23, Paramere.

LaPlata, Argentina (UNLP); National Museum of Natural History, Wash-
ington, D.C. (USNM).

Distribution.—Southern Brazil in states of Mato Grosso, Sao Paulo, and
Rio Grande do Sul, and presumably in states of Santa Catarina and Parana.

Euschistus (Lycipta) imitator Berg
Figs. 16-21

Euschistus imitator Berg, 1878: 306; Berg, 1879: 45-46 (reprint); Piran, 1948:
11 (record); Piran, 1968: 19, fig. 1G (record, pygophore).

Euschistus sellowi Berg, 1883: 208-209; Berg, 1884: 24-25 (reprint). New
Synonymy.

Humeral angles moderately produced laterad, acute, directed anterolat-
erad (Fig. 16). Connexiva broadly margined laterally with brownish yellow
to castaneous. Abdominal venter brownish yellow or rufous; dark puncta-
tion usually becoming sparse mesially. Veins in hemelytral membranes sim-
ple or branched, sometimes with | or 2 closed cells basally. Posterior margin
of basal plates mesially and mesial margin of 9th paratergites much thick-
ened, leaving 2nd gonocoxae sunken (Fig. 21). Emargination in posterior
pygophoral margin broad, concave, moderately deep from both caudal and
ventral view (Figs. 17, 18), shallow and transverse from dorsal view (Fig.
19). A large tuft of setae located on prominence at posterolateral corners of
genital cup. Parameres as in Fig. 20.

Distribution.—Argentina (Buenos Aires, Entre Rios, Santa Fé); Brazil
(Rio Grande do Sul, Santa Catarina); Paraguay; Uruguay.

VOLUME 84, NUMBER 2 289

| 24

=

Figs. 24-26. Euschistus (L.) sharpi. 24, Humerus and anterolateral pronotal margin. 25,
_ Genital cup; paramere (p); tumescence (tu). 26, Paramere.

Types.—The following specimen is designated lectotype of Euschistus sel-

— lowi: male, labeled (a) ‘‘Euschistus selowianus Berg Typen!,”’ (b) *‘Typus,”’

(c) ‘“*Zoolog. Museum Berlin,’’ (d) ‘‘1010,’’ (e) ‘“Montevideo Sello.”’ Three

female paralectotypes are all labeled (a) ‘‘Typus’’ (b) ‘‘Montevideo Sellow

No. 1010” (c) ‘‘Zoolog. Museum, Berlin.’’ All 4 specimens are in the Mu-

— seum fur Naturkunde der Humboldt—Universitat zu Berlin. These speci-
mens and the holotype of Euschistus imitator Berg were examined.

Euschistus (Lycipta) circumfusus Berg
Figs. 22, 23
Euschistus circumfusus Berg, 1883: 208; Berg, 1884: 24 (reprint); Buckup,
1961: 10 (record).

Dorsum fuscous with lateral margins of pronotum and basal margins of
coria yellow or red. Connexiva entirely dark or yellow margined. Abdominal
venter dark brown, densely and uniformly black punctate. Humeri scarcely
produced laterad, obtusely angulate (Fig. 22). Venation of hemelytral mem-
brane simple or furcate, sometimes with | or 2 closed cells basally. Genital
plates and pygophore as in L. imitator. Parameres as in Fig. 23.

Distribution.—Argentina (Entre Rios); Brazil (Santa Catarina, Rio Grande
do Sul); Paraguay; Uruguay.

Types.—The following specimen is designated lectotype: Male, labeled

290 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(a) ‘‘Euschistus circumfusus typen! Berg’’ (b) ‘““Typus”’ (c) **Zoolog. Mu-
seum Berlin’’ (d) ‘‘795’’ (e) ‘‘Montevideo, Sell.’’ The 3 paralectotypes, 2
2.13, are all labeled (a) “‘Typus”’ (b) *‘Montevideo, Sellow no. 795” (c)
‘‘Zoolog. Museum Berlin.”’ All 4 specimens are in the Museum fur Natur-
kunde der Humboldt—Universitat zu Berlin and were examined.
Comments.—The genitalia of this species and of E. imitator seem insep-
arable excepting perhaps minor differences in the parameres. In the rela-
tively few specimens of E. circumfusus and E. imitator that have been
available, differences in the humeral form and color have been consistent.

Euschistus (Lycipta) sharpi Bergroth
Figs. 24-26

Euschistus sharpi Bergroth, 1891: 223.

Dorsum including anterolateral pronotal margins fuscous to black, only
posterolateral margins of humeri, basal portion of coria along margin, and
entire connexiva pale. Abdominal venter brownish yellow with small punc-
tures rather weakly and uniformly dark colored. Humeri weakly produced
laterad, obtusely angled (Fig. 24). Venation in membrane of hemelytra sim-
ple or furcate. Posterior margin of pygophore sinuously convex from dorsal
view. A tuft of setae present at posterolateral corners of genital cup (Fig.
25). Parameres as in Fig. 26.

Distribution.—Brazil (Minas Gerais, Sao Paulo).

Type.—The type is apparently missing from the Fallou collection.

Comment.—This species closely resembles E. circumfusus but differs in
having the anterolateral pronotal margins black, in the punctation and col-
oration of the abdominal venter, in the uniformly pale connexiva, and in the
convexity of the posterior pygophoral margin when viewed dorsally.

I have seen no females of this species.

Euschistus (Lycipta) triangulator (Herrich-Schaffer)
Figs. 27-32
Cimex triangulator Herrich-Schaffer, 1842: 95-96, fig. 667.
Euschistus triangulator: Stal, 1860: 19 (listed).
Lycipta triangulator: Stal, 1862a: 58 (listed).
Euschistus (Lycipta) triangulator: Stal, 1872: 23 (keyed).

Veins of hemelytral membrane reticulate. Apex of scutellum ivory, usu-
ally somewhat reflexed. At least last 2 segments of antenna rufous, occa-
sionally pale at base. Humeri strongly projecting anterolaterad, acute api-
cally (Fig. 27). Posterior margin of basal plates emarginate at base of 9th
parategites, mesially thickened; mesial margin of one basal plate overlapping
other basally (Fig. 31). Posterior margin of pygophore trisinuate from dorsal

VOLUME 84, NUMBER 2 291

&

—

4

Figs. 27-32. Euschistus (L.) triangulator. 27, Humerus and anterolateral pronotal margin.
28, Posterior pygophoral margin, caudal view. 29, Same ventral view. 30, Genital cup, para-
meres and pygophore removed; tuft of setae (st); tumescence (tu). 31, Genital plates, caudo-
ventral view; basal plates (bp); 9th paratergites (pt 9). 32, Paramere.

and ventral views (Figs. 29, 30), broadly concave from caudal view (Fig.
28); setae tuft in posterolateral corners of genital cup inconspicuous amongst
other setae in cup.

Distribution.—Brazil (Santa Catarina, Sao Paulo, Rio de Janeiro).

Comment.—This species and the following one, E. picticornis, apparently
differ consistently only in the color of the antennae.

The type was not located. My concept of this species is based on the
literature.

Euschistus (Lycipta) picticornis Stal

Euschistus (Lycipta) picticornis Stal, 1872: 23.
Euschistus picticornis: Berg, 1879: 280-281 (nymph described); Berg, 1880:

292 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 33-38. Euschistus (L.) cornutus. 33, Humerus and anterolateral pronotal margin. 34,
Genital plates, caudoventral view; basal plates (bp); 9th paratergites (pt 9). 35, Posterior margin
of pygophore, caudal view. 36, Same, ventral view. 37, Genital cup, proctiger and parameres
removed; tumescence (tu). 38, Paramere.

8—9 (reprint); Berg, 1883: 206-208 (description); Berg, 1884: 22-24 (re-
print); Piran, 1948: 11 (record); Buckup, 1961: 11 (record); Grazia, 1977:
165-166.

Apparently differing consistently from E. triangulator only in rufous color
of antenna, especially last 2 segments. Apex of scutellum not reflexed. Dor-
sum usually a little darker in color than in E. triangulator.

Distribution.—Argentina (Buenos Aires, Parana de las Palmas, Sierra de
Cordoba); Brazil (Rio Grande do Sul, Santa Catarina, Sao Paulo).

Comments.—Berg (1883, 1884) listed a number of characters in which E.

VOLUME 84, NUMBER 2 293

Figs. 39-42. Euschistus (L.) luridus. 39-40, Variation in humeri. 41, Genital plates, cau-
doventral view. 42, Paramere.

picticornis and E. triangulator purportedly differ. None of these characters,
unfortunately, consistently differentiates between these species.
The holotype of E. picticornis was examined.

Euschistus (Lycipta) cornutus (Dallas)
Figs. 33-38

Euschistus cornutus Dallas, 1851: 201; Berg, 1878: 303-305 (description):
Berg, 1879: 42-44 (reprint); Bergroth, 1891: 223 (descriptive note); Jensen-
Haarup, 1922: 11, figs. 2a, 2b (habitus); Buckup, 1961: 11 (record); Piran,
1962: 6, 9, fig. 6 (pygophore).

Euschistus (Lycipta) cornutus: Stal, 1872: 23 (keyed).

Veins of hemelytral membrane usually reticulated. Apex of scutellum with
a small dark marginal macule on each side. Humeri cornute, projecting
anterolaterad, narrowly rounded at apex (Fig. 33). Dorsum yellowish brown,
humeri rufous. Costal margin of hemelytra toward base with small denticles.
Posterior margin of basal plates emarginate at base of 9th paratergites, thick-
ened mesially; mesial margin of one basal plate overlapping other basally;
2nd gonocoxae sulcate along meson (Fig. 34). Posterior margin of pygophore
broadly emarginate to moderate depth from caudal view (Fig. 35), less deep-
ly emarginate and with bottom of emargination slightly convex from ventral
view (Fig. 36), sinuous from dorsal view (Fig. 37). Setal tufts absent. Par-
amere as in Fig. 38.

Distribution.—Argentina (Misiones); Brazil (Rio Grande do Sul, Santa
Catarina, Minas Gerais); Paraguay.

294 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

<< tu

: Sea
;

49

Figs. 43-47. Euschistus (L.) cribrarius. 43, Humerus. 44, Posterior margin of pygophore,
caudal view. 45, Same, ventral view. 46, Genital cup; paramere (p); tumescence (tu). 47,
Paramere.

Comments.—This common species resembles E. machadus in coloration,
but the cornute humeri of E. cornutus separate these two species at a glance.
The type was examined.

Euschistus (Lycipta) luridus (Dallas), NEw COMBINATION
Figs. 39-42

Agroecus luridus Dallas, 1851: 200; Jensen-Haarup, 1937: 171 (keyed);
Buckup, 1961: 8, 11, Pl. 1 fig. 1, Pl. 2 fig. 1 (keyed, synonymy, descrip-
tion).

Euschistus aceratos Berg, 1894: 17. New synonymy.

Veins in membrane of hemelytra reticulate. Apex of scutellum not differ-
entiated by color or punctation. Humeri obtuse, produced laterad of base
of coria by width of eye or less (Figs. 39, 40). Genital plates (Fig. 41) similar
to those of E. cornutus (Fig. 34). Posterior margin of pygophore like that
of E. cribrarius (Figs. 44-46). Setal tufts absent. Pigmentation of parameres
disappearing basad of cup (Fig. 42).

Distribution.—Argentina (Misiones); Brazil (Rio Grande do Sul, Santa
Catarina, Minas Gerais); Bolivia.

VOLUME 84, NUMBER 2 295

Comment.—This species and E. cribrarius seem to differ only in the form
of the humeri and in minor differences in the parameres.
The types of Agroecus luridus and Euschistus aceratos were examined.

Euschistus (Lycipta) cribrarius Stal
Figs. 43-47

Euschistus (Lycipta) cribrarius Stal, 1872: 24 (keyed).

Differing from E. luridus in form of humeri and pigmentation of para-
meres. Humeri produced laterad of base of coria by about 1.5x width of
eye, apically acute and slightly retrorse (Fig. 43). Posterior margin of py-
gophore broadly and sinuously concave from caudal view (Fig. 44) sinuous
from ventral view (Fig. 45), convex with median projection from dorsal view
(Fig. 46). Paramere pigmented to basal disk (Fig. 47).

Distribution.—Brazil (Rio de Janeiro); Bolivia.

Comments.—The only substantial and apparently consistent difference
that I see between E. cribrarius and E. luridus is the form of the humeri,
which in many species of the genus, but by no means all of them, is noto-
riously variable. I have seen few males and no females of E. cribrarius, all
uniform with respect to their humeri, and about a score of E. luridus, the
humeri of none approaching the form of E. cribrarius.

The holotype was examined.

ACKNOWLEDGMENTS

Type-specimens for this study were graciously loaned by W. R. Dolling,
British Museum (Natural History), London; U. Gollner-Scheiding, Museum
fur Naturkunde der Humboldt—Universitat zu Berlin, East Germany; Per
Inge Persson, Naturhistoriska Riksmuseet, Stockholm, Sweden; and Luis
de Santis, Universidad Nacional de La Plata, Facultad de Ciencias Natu-
rales y Museo, Argentina.

Richard C. Froeschner, National Museum of Natural History, Washing-
ton, D.C., offered valuable suggestions for improving the manuscript.

LITERATURE CITED

Berg, C. 1878-80. Hemiptera Argentina enumeravit speciesque novas descripsit. An. Soc.
Cient. Argent. 5(6): 297-314 (1878); 9(1): 5-22 (1880). Reprinted 1879. Buenos Aires and
Hamburg. 316 pp. (Berg’s papers under this title in the Anales de la Sociedad Cientifica
Argentina were published as a book in December, 1879. The papers in this series that
appeared in the Anales in 1880 (vol. 9, no. 1) were also included, and presumably first
published, in the book.)

—. 1883. Addenda et emendanda ad Hemiptera Argentina. An. Soc. Cient. Argent. 15:
193-217. (Reprinted 1884. Buenos Aires and Hamburg. 213 pp.)

—. 1894. Descripciones de algunos Hemipteros Heteropteros nuevos 0 poco conocidos.
An. Mus. Montev. 1: 13-17.

296 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Bergroth, E. 1891. Contributions a l'étude des pentatomides. Rev. Entomol. 10: 200-235.

Buckup, L. 1961. Os pentatomideos do estado do Rio Grande do Sul (Brasil) (Hemiptera-
Heteroptera-Pentatomidae). Iheringia Ser. Zool. 16: 5-23.

Dallas, W. S. 1851. List of the specimens of hemipterous insects in the collection of the British
Museum. Part 1. London.

Grazia, J. 1977. Revisao dos pentatomineos citados no ‘‘Quarto Catalogo dos Insetos Que
Viven nas Plantos do Brasil’’ (Hemiptera-Pentatomidae-Pentatomini). Dusenia 10(3):
161-174.

Herrich-Schaffer, G. A. W. 1842. Die Wanzenartigen Insecten. Vol. 6, 118 pp., 34 pls.

Jensen-Haarup, A. C. 1922. Hemipterological notes and descriptions II. Entomol. Medd. Co-
penhagen 14: 1-16.

—_—. 1937. Einige neue Pentatomidenarten aus der Sammlung des Zoologischen Museums
in Hamburg (Hem. Het.) Entomol. Rundschau 54: 169-171, 321-324.

Kirkaldy, G. W. 1909. Catalogue of the Hemiptera (Heteroptera) Vol. 1. Cimicidae. XL +
392 pp.

Piran, A. A. 1948. Contribucion al conocemiento de la dispersion geographica de los Hem-

ipteros neotropicales. Acta Zool. Lilloana 5: 5-17.

. 1968. Hemiptera neotropica. XI. Rev. Soc. Entomol. Argent. 30: 17-25.

Sailer, R. I. 1954. Interspecific hybridization among insects with a report on cross breeding
experiments with stink bugs. J. Econ. Entomol. 47: 377-383.

Stal, C. 1860-62a. Bidrag till Rio Janeiro-traktens Hemipter-fauna. K. Svenska Vet.-Ak.

Handl. 2(7): 1-84 (1860); 3(6): 1-75 (1862).

. 1862b. Hemiptera Mexicana enumeravit speciesque novas descripsit. Stettin. Entomol.

Ztg. 23: 81-118.

—. 1872. Enumeratio Hemipterorum. Enumeratio Cimicinorum Americae. K. Svenska
Vet.-Ak. Handl. 10(4): 3-65.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 297-300

CLANONEURUM AMERICANUM (DIPTERA: EPHYDRIDAE), A
LEAFMINER OF THE LITTORAL CHENOPOD
SUAEDA LINEARIS

A. G. WHEELER, JR.

Bureau of Plant Industry, Pennsylvania Department of Agriculture, Har-
risburg, Pennsylvania 17110.

Abstract.—The first larval feeding habits are recorded for Clanoneurum
americanum Cresson, a psilopine ephydrid restricted to saline habitats. In
the Florida Keys larvae were found mining the leaves of sea blite, Suaeda
linearis (Ell.) Mog. (Chenopodiaceae). Brief notes on biology and behavior
and a scanning electron micrograph of the peculiar adult habitus are pre-
sented.

Clanoneurum americanum Cresson, a psilopine ephydrid or shore fly of
saline environments, exhibits a distinctive adult habitus, particularly the
wing venation. Cresson (1940) described this species from San Diego, Cal-
ifornia, noting that it was closely related to the Palearctic C. cimiciforme
(Haliday) and that all records of the genus in North America should refer
to this new species. Previous records included Maine (Mount Desert), Mas-
sachusetts (Woods Hole), New Jersey (Morgan, in salt marsh), and New
York (Gardiners Island) (Weiss, 1924). Cresson (1942) summarized the dis-
tribution of this ‘‘comparatively scarce species’’ as the Atlantic and Gulf
States, in addition to Arizona, California, and Utah. Wirth (1968) added
Mexico and the West Indies (Bahamas) to the known range of C. ameri-
canum, the only known North American species of the genus.

Larval feeding habits are recorded for few psilopine species (Foote and
Eastin, 1974), and no information is available on the habits of C. america-
num (Simpson, 1976). Old World members of the genus are known to mine
leaves of Chenopodiaceae (Hering, 1957; Wirth, 1965), with C. cimiciforme
recorded from sugar beets, Beta vulgaris L., by Balachowsky and Mesnil
(1936) and Hennig (1943) (cited as C. menozzii, a junior synonym of C.
cimiciforme); and from a glasswort, Salicornia europaea L. (cited as S.
herbacea L.), by Meijere (1947). Meijere’s paper included a brief description
and an illustration of the puparium and an illustration of the cephalopha-
ryngeal skeleton. Because the habits of C. americanum are unknown, it

298 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

. a ¥
Figs. 1, 2. Clanoneurum americanum. 1, Adult habitus, scanning electron micrograph

(50x). 2, Proboscis extended. feeding on brewer’s yeast.

VOLUME 84, NUMBER 2 299

seems worthwhile to record this species as a leafminer of a littoral chenopod
and present notes on its life history and habits.

BIOLOGICAL NOTES

I made observations on C. americanum during a trip to the Florida Keys
in April 1981. The habitat on Upper Key Largo consisted of coral rock land
dominated by various halophytes. Large colonies of saltwort, Batis mar-
itima L. (Bataceae), were growing among smaller patches of sea purslane,
Sesuvium portulacastrum L. (Portulacaceae), and perennial saltwort, Sali-
cornia virginica L. (Chenopodiaceae). Clanoneurum americanum, how-
ever, was observed only on another chenopod, Suaeda linearis (Ell.) Mog.,
or sea blite. This herbaceous halophyte has numerous fleshy, linear leaves
that form terminal spikes. It occurs in salt marshes from Maine south to
Florida and west to Texas, and also is known from the West Indies (Long
and Lakela, 1976).

In the field, adults of C. americanum appeared more homopteran than
dipteran. The infumate wings held almost rooflike over the abdomen (Fig.
1) and the short hopping flight were suggestive of a small black fulgoroid,
especially the macropterous form of the issid planthopper, Bruchomorpha
oculata Newman. Adults also superficially resembled a small dark weevil
(Fig. 2) as they walked over leaves, proboscices extended, apparently for
probing the surface.

Field-collected adults were placed in small plastic boxes containing sprigs
of S. linearis. Pairing occurred within minutes, apparently without a pre-
mating display or elaborate mating behavior; some pairs remained in copula
for several hours. The white eggs were deposited singly on host foliage and,
upon hatching, larvae bored into tissue beneath or adjacent to the egg shell
and tunneled through the linear leaf. Leaves containing mature larvae ap-
peared hollowed out or traversed by a wavy, light-colored thread. In the
laboratory, puparia were found within the mines or on the bottom of the
rearing containers just beneath mined leaves.

Adult emergence from foliage began on May 5-6; eggs had been laid on
the same foliage on April 15. Under laboratory conditions, the approximate
developmental time was 3 weeks. Old mines and eggs were present in the
field on April 13; it is likely that C. americanum is multivoltine, producing
successive generations in Florida. A careful study of this specialized shore
fly would add to the knowledge of feeding habits reported for the large and
diverse subfamily Psilopinae.

ACKNOWLEDGMENTS

I thank my Harrisburg colleagues K. Valley, for identifying the ephydrid
and reading the manuscript, and J. F. Stimmel, for the photograph. E. R.
Hoebeke (Department of Entomology, Cornell University, Ithaca, New

300 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

York) and K. W. Simpson (Div. Lab. and Research, State of New York
Department of Health, Albany) also made valuable comments on the manu-
script.

LITERATURE CITED

Balachowsky, A. and L. Mesnil. 1936. Les insectes nuisibles aux plantes cultivées. Vol. 2.
Presses des Etablissements Busson, Paris. Pp. 1141-1921.

Cresson, E. T., Jr. 1940. Descriptions of new genera and species of the dipterous family

Ephydridae. Paper XII. Acad. Nat. Sci. Phila. Notulae Nat. 38: 1-10.

. 1942. Synopses of North American Ephydridae (Diptera). Trans. Am. Entomol. Soc.

68: 101-128.

Foote, B. A. and W. C. Eastin. 1974. Biology and immature stages of Discocerina obscurella
(Diptera: Ephydridae). Proc. Entomol. Soc. Wash. 76: 401-408.

Hennig, W. 1943. Ubersicht tiber die bisher bekannten metamorphosesstadien der Ephydriden
Neubeschriebungen nach dem Material der Deutschen Limnologischen Sunda-Expedi-
tion (Diptera: Ephydridae). Arb. Morphol. Taxon. Entomol. 10: 105-138.

Hering, E. M. 1957. Bestimmungstabellen Blattminen von Europa einschliesslich des Mittel-
meerbeckens und der Kanarischen Inseln. Band I. Pfanzengattungen A—L. Dr. W. Junk,
‘s-Gravenhage. 648 pp.

Long, R. W. and O. Lakela. 1976. A flora of tropical Florida; a manual of the seed plants and
ferns of southern peninsular Florida. Banyan Books, Miami, Fla. 962 pp.

Meijere, J. C. H. de. 1947. Over eenige Dipterenlarven, waaronder een galmug, die mijngangen
maakt, en twee Dipteren, die gallen op paddenstoelen veroorzaken. Tijdschr. Entomol.
88: 49-62 (1945).

Simpson, K. W. 1976. Shore flies and brine flies (Diptera: Ephydridae), ch. 17, pp. 465-495.
In Cheng, L., ed., Marine insects. North-Holland Publishing Co., Amsterdam.

Weiss, H. B. 1924. Diptera collected on a New Jersey salt marsh. Can. Entomol. 56: 255-257.
Wirth, W. W. 1965. Family Ephydridae, pp. 734-759. In Stone, A. et al., eds., A catalog of
the Diptera of America north of Mexico. U. S. Dep. Agric. Agric. Handb. 276.
———. 1968. Family Ephydridae, pp. 1-43. In Papavero, N., ed., A catalogue of the Diptera
of the Americas south of the United States. Dep. Zool. Sec. Agric., Sao Paulo.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 301-303

IS THE BINOMEN LYGAEUS KALMII STAL
(HEMIPTERA: HETEROPTERA: LYGAEIDAE)
APPLIED TO SIBLING SPECIES?

JOSE DUARTE AND DIANE M. CALABRESE

(JD) Department of Biology, University of Texas at El Paso, El Paso,
Texas 79968; (DMC) Department of Biology, Trinity College, Washington,
D.C. 20017; present address, Department of Biology and The Wildlife Sanc-
tuary, Dickinson College, Carlisle, Pennsylvania 17013.

Abstract.—Populations of Lygaeus kalmii Stal from west Texas show a
diploid number of 18 and a homogametic sex determination mechanism in
the male. A diploid number of 14 and a heterogametic male have been
reported for the species in the literature.

We report unexpected results obtained when one of us (JD) began a cy-
togenetic study of Lygaeus kalmii Stal from west Texas in order to learn
cytogenetic technique. A diploid number of 14 (XY) has already been re-
ported for the species (Ueshima and Ashlock, 1980; Ueshima, 1979) and a
diagram of its chromosome complement given (Ueshima and Ashlock,
1980).

MATERIALS AND METHODS

Lygaeus kalmii adults and immatures were collected from Solanum sp.
in El Paso, Tex. at different times over a three month period from June to
August 1980. Some bugs were lab-reared, and some were used immediately
in preparations.

Testes of fifth-instar stage nymphs were dissected in insect Ringer’s so-
lution and fixed (15 ml glacial acetic, 45 ml ethanol, 5 ml acetone). Testes
were stained with lacto-proprio orcein and squashed. Numerous squashes
were made. Preparations showing metaphase I (meiotic) chromosomes were
photographed, although other stages were observed. Figures were photo-
graphed at about 6400 under oil on a Zeiss compound microscope with a
Wetzler large format camera attachment.

RESULTS

All results for west Texas populations are consistent. The chromosome
number appears to be 2N = 18 (N = 8A + W) based on spermatogenic
metaphase I cells (Fig. 1). The male appears to be homogametic (Fig. 1).

302 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. |. Lygaeus kalmii, metaphase I chromosomes photographed at about 6400x.

DISCUSSION

The difference in chromosome number between L. kalmii collected from
west Texas and specimens collected from California and reported by Ue-
shima and Ashlock (1980) suggest the possibilities of clinal variation and
sibling species. One of us (DMC) contacted N. Ueshima for his opinion of
the discrepancy in results between the two populations. He suggested (per-
sonal communication) that a series of fusions may have led to the differences
between the populations. Such an explanation would be consistent with the
large size of two of the chromosomes (Fig. 1), if Fig. 1 were a mitotic phase
as Ueshima believes (personal communication). However, after viewing se-
ries of maturation stages in spermatogenic cells of L. kalmii we are certain
that we have metaphase I chromosomes (e.g. Fig. 1). Tetrads are visible
and kinetochores are terminal (typical meiotic kinetochore behavior in the
Heteroptera; Ueshima, 1979) and therefore, our conclusion is well-support-
ed. No figures were photographed by Ueshima or Ashlock for L. kalmii
(personal communication, Ueshima and Ashlock).

Within the Heteroptera fragmentation and fusion of autosomes have prob-
ably been important in producing changes in chromosome numbers (Ue-
shima, 1979; Schrader and Hughes-Schrader, 1956; Schrader, 1947). Thus,
it is not unreasonable to assume that we are looking at a dynamic process
in evolution within the populations recognized under the binomen L. kalmii.

VOLUME 84, NUMBER 2 303

The question of whether we have clinal variation in chromosome number
or sibling species will be determined by a combination of hybridization
studies (including cytogenetic study of eastern United States populations
(DMC) and life history and host plant studies (JD)).

ACKNOWLEDGMENTS

We thank Peter Ashlock, University of Kansas, Lawrence, for verifying
our identification of voucher specimens and N. Ueshima, Matsusaka Col-
lege, Matsusaka-shi, Mie, Japan, for examining our photographs of sper-
matocyte metaphase I chromosomes. P. Tallerico shared invaluable pho-
tographic expertise with us.

LITERATURE CITED

Schrader, F. 1947. The role of the kinetochore in the chromosomal evolution of the Heter-
optera and Homoptera. Evolution 1: 134~142.

Schrader, F. and S. Hughes-Schrader. 1956. Polyploidy and fragmentation in the chromosomal
evolution of various species of Thynata (Hemiptera). Chromosoma (Berl.) 7: 469-496.

Ueshima, N. 1979. Animal Cytogenetics. Vol. 3. Insecta 6. Hemiptera II, Heteroptera. Ge-
briider Borntraeger, Stuttgart. 117 pp.

Ueshima, N. and P. D. Ashlock. 1980. Cytotaxonomy of the Lygaeidae (Hemiptera-Heter-
optera). Univ. Kans. Sci. Bull. 51: 717-801.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 304-314

DESCRIPTION OF THE LARVA AND FEMALE OF
PYCNOPSYCHE FLAVATA (BANKS) WITH COMPARATIVE NOTES
ON THE ECOLOGY OF P. FLAVATA AND P. GENTILIS
(MCLACHLAN) (TRICHOPTERA: LIMNEPHILIDAE)

JOHN A. WOJTOWICZ

Department of Zoology, University of Tennessee, Knoxville, Tennessee
37916.

Abstract.—The larva and female of Pycnopsyche flavata (Banks) are de-
scribed and figured for the first time. Ecological notes are provided for
sympatric populations of P. flavata and P. gentilis (McLachlan). The highly
variable morphology of P. gentilis larvae throughout the range of the species
is discussed.

Members of the genus Pycnopsyche are of widespread and common oc-
currence throughout eastern North America. Larvae have been described
for only six of the presently recognized 16 species in the genus (Flint, 1960).
Reared material and/or metamorphotypes are available for P. subfasciata
(Say) (Vorhies, 1909; Flint, 1960; G. B. Wiggins, personal communication)
and P. sonso (Milne) (Flint, 1960); however, no specific characters for their
identification have yet been elucidated.

The known distribution of P. flavata (Banks) is restricted to the mountains
of western North Carolina, southwestern Virginia (C. R. Parker, personal
communication), eastern Tennessee, northwestern South Carolina (J. C.
Morse, personal communication), and northern Georgia. Collections of P.
flavata \arvae in Tennessee and North Carolina indicate a fairly specialized
habitat preference.

Herein is described the larva of P. flavata with notes on its ecology
obtained from year-round collections at a site in Tennessee. Because of the
close morphological and behavioral similarity and frequent sympatric oc-
currence of P. flavata and P. gentilis (McLachlan), information on their life
history in the southern Appalachians and on the variability of certain char-
acters of P. gentilis larvae from five localities in eastern North America is

also included. The female of P. flavata is described and figured for the first
time.

VOLUME 84, NUMBER 2 305

Fig. 1. Larvae of Pycnopsyche flavata and P. gentilis. A, Head of P. flavata. B, Head of
southern P. gentilis. C, Head of northern P. gentilis. D, Dorsal view of first abdominal segment
of P. gentilis. E, Dorsal view of first abdominal segment of P. flavata. DSH = dorsal spacing
hump. Camera lucida drawings.

MATERIALS AND METHODS

Collections of P. flavata larvae were first made in December 1977 in the
Tellico Wildlife Management Area of the Cherokee National Forest, Monroe
Co., Tennessee. Light trap samples of adults were obtained in July 1978,
and pupae and larvae were brought back for rearing. Specimens were reared
in aluminum window-screen envelopes partially submerged in a spring seep.
Ten collections of larvae and/or pupae were made on an approximately
monthly basis between July 1978 and June 1979. Additional larvae and pupae
were obtained for rearing in June 1979. In all, 18 pupae and larvae were

306 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

reared for association purposes. An additional reared specimen was also
discovered in some unidentified material borrowed from the Illinois Natural
History Survey.

The description of the larva of P. flavata is based on 11 fifth-instar larvae
from the Tellico site. Numerous other larvae from several localities were
examined for much of the additional information in this paper. Larval instar
determinations for P. flavata and P. gentilis were made using the method
of approximate head widths (Dyar, 1890; Mackay, 1978). The notes on vari-
ation in P. gentilis were obtained from a study of 9 to 10 larvae from each
of five localities including Essex Co., N.Y., Westmoreland Co., Pa., Bath
Co., Va., Monroe Co., Tenn., and Morgan Co., Tenn. The larval description
follows the format of Flint (1960) who provided a comparative account of
Pycnopsyche larvae. The description of the female is based on 8 adults from
Tennessee and North Carolina.

DESCRIPTION OF FINAL INSTAR LARVA OF PYCNOPSYCHE FLAVATA

Length, 19.8-24.8 mm. Maximum width, 3-4 mm. Head: Yellowish
brown. Muscle scars with dark borders on frons and posterior portion of
genae. Front of head distinctly flattened with ridge of spines arcing from
eye to eye. All 18 major setae present. Setae 13 and 16 flat and bladelike.
Seta 16 on ridge of head appearing as member of arcuate band of spines.
Setae | and 4 hyaline and appressed. Setae 6, 10, 11, and 12 light colored
and cryptic. Seta 18 small and not obvious. Head width, 1.7—2.1 mm. Tho-
rax: Pronotum yellowish brown with a series of spines on anterior edge and
spinelike and hairlike setae dorsally. Mesonotum yellowish brown with
spinelike and hairlike setae dorsally. Muscle scars present on pro- and meso-
notum. Metanotum with Sal, Sa2, and Sa3 sclerites. Metanotal Sal sclerites
unfused. Setae present on membrane of metanotum. Legs yellowish brown.
Prothoracic leg with 2 major spinelike setae ventrally on femur. Femora of
meso- and metathoracic legs each with 2 major hairlike setae on ventral
ridge. Abdomen: First segment with lateral spacing humps and remnants of
dorsal spacing hump. Elongate sclerite present at posterior of each lateral
hump. Dorsum (Sal + Sa2 areas) of Ist segment with 42-54 setae on either
side of dorsal hump. Venter (Sal area) of Ist segment with 20-31 setae on
either side. Sa2 sclerites on venter of Ist segment with 4-9 setae each.
Eighth segment with 9-14 setae posteriorly on dorsum. Dorsal sclerite of
9th segment with 10-15 setae. Gills: Dorsal, I] 0,1; If 1,1; IV 1,1; V 1,0-1;
VI 0-1,0. Dorso-lateral, IIT 1,0; IV 0-1,0. Ventrolateral, II 0,1; II 0,0-1.
Ventral) Tl 030-1; TH 1,1: TV 1,12 Va svi Vee

Diagnosis.—Of the known larvae of Pycnopsyche, that of P. flavata bears
closest resemblance to that of P. gentilis. The larvae of both species possess
bladelike 13th and 16th head setae. In P. gentilis both seta 13 and 16 are
located on the curvature of the genae distinctly separate from other setae

VOLUME 84, NUMBER 2 307

ev |
ae at
—* 4
: ra , x
»~

»
Be
Fig. 2. Pycnopsyche flavata larva showing placement of setae 13 and 16. 76x. SEM pho-
tograph.

308 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

and spines of the head (Figs. 1B, C; 3, 4). In P. flavata seta 16 is located
in close apposition to an arcuate band of spines extending from eye to eye
(Figs. 1A, 2). In addition, the head of P. flavata is distinctly flattened an-
teriorly. The head of P. gentilis, although often with a band of spines,
displays a gradual rounding both laterally and dorsally. Finally, P. gentilis
larvae lack a dorsal spacing hump on the Ist abdominal segment while P.
flavata larvae possess at least the remnants of such a hump (Fig. 1D, E).

DESCRIPTION OF FEMALE OF PYCNOPSYCHE FLAVATA

Length of forewing, 18-21 mm. Pale orange to tan. Forewing with a slight
darkening along cord from R; to M, and with M-vein just proximal to cord,
pale. Spur formula, 1-3-4. Opening of abdominal scent gland on antero-
lateral portion of Sth sternum. Genitalia. (Fig. 5): Segment IX in lateral
view extended posteriad to sharp point dorsally, narrowed medially and
widened and rounded ventrally. Dorsal IX with central concavity when
viewed obliquely. Segment IX narrowed, rounded and tonguelike with me-
dian notch posteriorly in dorsal view. Ventral IX produced into cercus-like
lobes. Dorsal X short and indistinct, slightly visible in dorsal view. Ventral
X produced posteriorly into long, sclerotized scoop. Supragenital plate
(SG.P.) membranous and not produced in lateral view. Vulvar scale (V.S.)
produced to sharp point in lateral view, trilobed with central lobe mucronate
viewed ventrally. Patches of setae on posterior VII and lateral to V.S. on
VIII.

Diagnosis.—Pycnopsyche flavata is a member of the divergens group of
Schmid (1955). Within this group the closest relative of P. flavata is P.
gentilis. The female of P. flavata differs from that of P. gentilis mainly in
the greater development of segment X. In P. flavata ventral X extends well
beyond dorsal IX in lateral view. In P. gentilis ventral X extends at most
even with dorsal IX and is often shorter.

VARIATION IN THE LARVAE OF P. GENTILIS

Although in all known instances the larvae of P. gentilis are readily dis-
tinguishable from those of P. flavata, a sizeable amount of variation is
evident in P. gentilis from different localities. As previously reported by
Flint (1960) the southern larvae of P. gentilis (Fig. 1B) display a much
greater development of head spination than do the northern larvae (Fig. 1C),
making them strikingly similar to the larvae of P. flavata (Fig. 1A). In this
study, larvae from northern New York State were found to show extremely
sparse spicule development. Those from southern Pennsylvania and south-
ward, however, show considerably greater spicule development. Because
of the difficulty of quantifying such a character, no attempt was made to
determine if the variation is clinal. In addition, significant mean differences
were shown to occur among localities in dorsal (Sal + Sa2) and ventral

VOLUME 84, NUMBER 2

ste. e
pr a
“ee F¢

ae 13 and 16. 76

Fig. 3. Pycnopsyche gentilis larva, southern, showing placement of set
SEM photograph.

310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(Sal) setal counts of the first abdominal segment (P > F = .01). Gill for-
mulas also showed a degree of variability (mostly on the posterior seg-
ments), for the most part showing a greater development of gills than re-
ported by Flint (1960).

Recently, it was brought to the author’s attention that certain populations
of P. gentilis adults have males with aedeagal titillators strikingly different
from those of typical P. gentilis (D. A. Etnier, personal communication).
Further study has revealed variability in tibial spur counts both among and
within localities of P. gentilis, most of the males with the unique titillators
having 1-2-4 spur formulas as opposed to the typical 1-3-4. Unfortunately,
the 1-2-4 spur formulas are not restricted to the P. gentilis with the unique
titillators and further work will be necessary to clarify the significance of
this variability.

NOTES ON THE LIFE HISTORY AND ECOLOGY OF
P.. FLAVATA AND P. GENTILIS

Habitat.—The larvae of P. flavata are restricted to a rather specific hab-
itat type in the southern Appalachians. Collections in North Carolina and
Tennessee show that the larvae are most often found in the headwater areas
of streams, typically at elevations near or above 1000 m. Within this habitat
P. flavata larvae prefer spring heads where the water is slow moving and
shallow (ca. 2-5 cm), although they may be found quite commonly in larger
streams earlier in their development. Within their range, the larvae of P.
flavata are nearly always found to occur sympatrically with the larvae of P.
gentilis and occasional collections have also yielded both P. sonso and P.
luculenta (Betten) at the same locality.

Case structure.—Monthly collections during 1978 and 1979 at sites in
eastern Tennessee indicate that early instar P. gentilis and P. flavata may
construct triangular leaf cases as reported by Flint (1960) and Mackay (1972)
or may instead by-pass these and immediately begin construction of stone
cases. The later instar cases of both species are cylindrical and constructed
of small stones. The type and particular size of stones used in the case
appear to be more a function of availability than of specific preference, as
indicated by variability among collection sites, especially in P. genitilis.
Although selection for specific substrate sizes in case building may occur
within locality (Mackay, 1977) my observations suggest that this is not a
species-wide phenomenon.

Larval development.—The third-, fourth-, and fifth-instars were readily
identifiable for both species. Mean head widths of comparable instars of P.
flavata and P. gentilis differ noticeably, those of P. gentilis being somewhat
smaller. A comparison of data for P. gentilis with that of Mackay (1972)
indicates a slightly smaller head width for the southern larvae. Third-instar
P. flavata show a mean head width of .82 mm (range: .71—.87 mm; mode:

VOLUME 84, NUMBER 2

Fig. 4. Northern Pycnopsyche gentilis
SEM photograph.

311

*

larva showing placement of setae 13 and 16. 76»

312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Female genitalia of Pycnopsyche flavata. A, Lateral view. B, Dorsal view. C,
Ventral view. SG.P. = supragenital plate; V.S. = vulvar scale. Camera lucida drawings.

.74 mm; n = 66) vs. a mean of .76 mm (range: .71—.90 mm; mode: .74 mm;
n = 44) in P. gentilis. Mean head widths for fourth-instar larvae are 1.25
mm (range: 1.15—1.36 mm; mode: 1.26 mm; n = 83) for P. flavata and 1.17
mm (range: 1.05—1.29 mm; mode: 1.18 mm; n = 23) for P. gentilis. Fifth-
instar larvae of P. flavata have a mean head width of 1.84 mm (range:
1.68-2.06 mm; mode: 1.84 mm; n = 121) while those of P. gentilis have a
mean head width of 1.65 mm (range: 1.58—1.87 mm; mode: 1.71 mm; n =
30). Identification of second-instar P. flavata was tenuous at best. Second-
instar identification of P. gentilis was more dependable because of the ob-
vious lack of a dorsal spacing hump. Mean head width for second-instar P.
gentilis larvae is .47 mm (range: .45—.52mm; mode: .48 mm; n = 32). No
first-instar larvae could be positively identified for either species. Results
indicate that larvae of P. flavata may begin to reach fifth (final) instar as
early as October, with a large number of fifth-instar larvae occurring by
December and January. In P. gentilis, on the other hand, although second-
and third-instar larvae were collected in late November, no fifth-instar lar-
vae were evident until March!. This agrees with the findings of Mackay
(1972) which is surprising considering the difference in the latitude of the
populations studied.

' The fifth-instar of P. gentilis may have been reached as early as February; however,
weather conditions did not permit collection of a sample in February.

VOLUME 84, NUMBER 2 313

Diapause and pupation.—A number of larvae of both species attain final
instar as early as five or six months before pupation and adult eclosion. This
suggests some sort of resting period such as has been reported for other
species of Pycnopsyche (Wiggins, 1977; Mackay, 1972; Cummins, 1964).
Present data do not allow distinction between a period of reduced larval
activity or true physiological diapause.

The larvae of P. flavata do not appear to seal the front of their cases with
small stones much in advance of the actual time of pupation. Sealed cases
are evident only infrequently before June. Placement of an anterior sieve
plate appears to occur just prior to pupation. Larvae with sealed cases but
no anterior sieve plate often re-emerge when disturbed.

Prior to their resting period and subsequent pupation the larvae of both
P. gentilis and P. flavata may anchor their cases to either larger stones or,
occasionally, pieces of wood. In some instances a number of smaller stones
may be attached to the front of the case to serve as an anchor.

Adult emergence.—Available records for emergence of P. flavata adults
range from May 18 in Virginia (C. R. Parker, personal communication) to
October 2 in North Carolina (G. B. Wiggins, personal communication).
Southern Appalachian P. gentilis emerge later with records ranging from
August 12 to October 30 in Virginia (C. R. Parker, personal communication).
The majority of records, however, indicate that most P. flavata emerge
from early May to late August, while most P. gentilis from comparable
latitudes emerge in September and October. The virtual lack of overlap
between the adult flights of the two sympatrically occurring species probably
serves as a reproductive isolating mechanism leading to later hatching and
development of P. gentilis larvae which limits competition between com-
parable instars of both species.

ACKNOWLEDGMENTS

This paper could not have come about without the aid of a number of
people. For assistance in collecting I thank Robert L. Jones, Raymond C.
Stone, Jr., Elizabeth B. Williams, Richard A. Wojtowicz, and G. William
Wolfe. For permission to collect at the Powdermill Run Nature Reserve of
the Carnegie Museum I thank Joseph Merritt. For loan of specimens I thank
J. D. Unzicker, Illinois Natural History Survey, Champaign. For use of
personal and unpublished collecting records I thank J. D. Unzicker; Glen
B. Wiggins, Royal Ontario Museum, Toronto; Charles R. Parker, V.P.1.
& S.U., Blacksburg, Virginia; J. C. Morse, Clemson University, Clemson,
South Carolina; O. S. Flint, Jr., National Museum of Natural History,
Washington, D.C. Camera lucida and other equipment were kindly made
available by D. L. Bunting, University of Tennessee. I thank Dick Williams
and M. L. Pan for assistance in SEM work. For helpful hints on biological
illustration I thank Wynne Brown. Finally, for review of the manuscript in

314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

part or in whole, I thank D. L. Bunting, D. A. Etnier, J. C. Morse, and M.
L. Pan. This research was supported in part by a grant from Sigma Xi, The
Scientific Research Society.

LITERATURE CITED

Dyar, H. G. 1890. The number of molts of lepidopterous larvae. Psyche (Camb. Mass.) 5:
420-422.

Cummins, K. W. 1964. Factors limiting the microdistribution of larvae of the caddisflies Pycno-
psyche lepida (Hagen) and Pycnopsyche guttifer (Walker) in a Michigan stream (Tri-
choptera: Limnephilidae). Ecol. Monogr. 34: 271-295.

Flint, O. S., Jr. 1960. Taxonomy and biology of Nearctic limnephilid larvae (Trichoptera),
with special reference to species in eastern United States. Entomol. Am. 40: 1-117.

Mackay, R. J. 1972. Temporal patterns in life history and flight behaviour of Pycnopsyche
gentilis, P. luculenta, and P. scabripennis (Trichoptera: Limnephilidae). Can. Entomol.

104: 1819-1835.

. 1977. Behavior of Pycnopsyche (Trichoptera: Limnephilidae) on mineral substrates in

laboratory streams. Ecology 58: 191-195.

. 1978. Larval identification and instar association in some species of Hydropsyche and

Cheumatopsyche (Trichoptera: Hydropsychidae). Ann. Entomol. Soc. Am. 71: 499-509.

Schmid, F. 1955. Contribution a l'étude des Limnophilidae (Trichoptera). Mitt. Schweiz. Ento-
mol. Ges. 28: 1-245.

Vorhies, C. T. 1909. Studies on the Trichoptera of Wisconsin. Trans. Wis. Acad. Sci. 16:
647-738.

Wiggins, G. B. 1977. Larvae of the North American caddisfly genera (Trichoptera). Univ. of
Toronto Press, Toronto. 401 pp.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 315-324

THE NOCTUOID MOTHS OF THE ANTILLES—PART II
(LEPIDOPTERA: ARCTIIDAE: PERICOPINAE)

Ba: Topp

Systematic Entomology Laboratory, IIBHI, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Sixteen Pericopinae species reported from the Antilles are
treated. Thirteen species, including Stenognatha flinti, n. sp., are illustrated.
Ira gundlachiana Neumoegen is transferred to Syntomidopsis and is thus
included in the subfamily.

The Arctiidae of the Antilles represent a fairly large group of species and
remain mostly unstudied. In the course of my studies of the noctuoid moths
of the Antilles, I have checked collections for material, collected on seven
of the Islands, attempted to identify the material located in collections or
recently collected, reviewed the literature pertaining to previous studies,
tried to recognize undescribed species, etc. Unfortunately, visual problems
in 1979 necessitated retirement and brought detailed study of the Arctiidae
to a halt. Since that time, I decided to publish illustrations of the Antillean
species available to me, and to indicate what problems are unresolved.
Much more work is required to characterize the various subfamily groups.
At the present time classification is based mainly on venation. Characters
of the tympanum, genitalia, thoracic furrows, and other systems need to be
examined and combined with venational characters to develop a sound clas-
sification. All genera and species should be studied because several at pres-
ent appear to be misplaced.

Syntomidopsis gundlachiana (Neumoegen), NEw COMBINATION
Fig. |

Ira gundlachiana Neumoegen, 1890: 64.

Length of forewing.—Male, 12 mm.

This species has not been included in the Pericopinae previously. When
described, Neumoegen proposed a new genus, /ra, 1890, but that generic
name is preoccupied by Ira Walker, 1866 (Geometridae). The species is
close to the following species, but the male of S. variegata (Walker) has

316 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

some longitudinal black marks on the base of the forewing, has three distinct
white spots in the black apex of that wing, and the white spot of the tornal
angle is much more distinct. The only specimen I have examined is the type
male. The female sex is unknown to me. The type-specimen is from Plan-
tation Jagey, S.E. Cuba.

Syntomidopsis variegata (Walker)
Figs. 2, 3

Eudule variegata Walker, 1854: 380.
Syntomidopsis variegata (Walker): Hering, 1925: 452.
Syntomidopsis variagata (sic) (Walker): Bryk, 1931: 47.

Length of forewing.—Male, 15 mm; female, 17 mm.

Fifteen examples from Jamaica in the collection of the National Museum
of Natural History have been examined. The species differs from S. gund-
lachiana as explained under that species, and it is also slightly larger. The
pink coloration of the wings of both species separate them from all other
Pericopinae species I know.

Stenognatha cinda (Schaus)
Figs. 4, 5

Hyalurga cinda Schaus, 1938: 508.
Stenognatha cinda (Schaus): Lane and Watson, 1975: 109.

Length of forewing.—Male, 20 mm; female, 24.5 mm.

Lane and Watson (1975: 109-117) stated how the species of the genus
known to them differed from each other. In total I have examined four
examples. The pair figured here are the types and are in the National Mu-
seum of Natural History. The other two are in the British Museum (Natural
History), London, England. The type-locality is Las Animas, Pinar del Rio,
Cuba. The maculation of the forewing and coloration of the hindwing as
shown in the figures is distinctive for each species of this genus.

Stenognatha flinti Todd, NEw SPEcIES
Fig. 6

Length of forewing.—Female, 23.5 mm.

Type.—Holotype, 2°, 12 km N. of Constanza, Dominican Republic, 6-18
June 1969, Flint and Gomez, in the National Museum of Natural History.

This female and the species described by Lane and Watson as S. toddi
in 1975 were known to me before Don R. Davis went to Jamaica and to
Hispaniola in 1973. I showed him examples including the unique type-spec-
imen of this species and asked that he try to get males on Hispaniola, but
he collected no examples there, so I know only the type of this species. The
hindwing of this species is so different from that of the other species that
I decided to describe it even though the male is not known. The hindwing

VOLUME 84, NUMBER 2 317

“ eA as : ai
Figs. 1-8. Dorsal view of adults. 1, Syntomidopsis gundlachiana, type male, Cuba. 2, S.
variegata, male, Jamaica. 3, S. variegata, female, Jamaica. 4, Stenognatha cionda, male,
Cuba. 5, S. cionda, female, Cuba. 6, S. flinti, type female, Dominican Republic. 7, S. toddi,
male, Jamaica. 8, S. toddi, female, Jamaica.

is mostly smoky gray except for a small yellow patch on the anal margin
and the fringe is completely white and contrasts with the rest of the wing.

Stenognatha toddi Lane and Watson
Figs. 7, 8

Stenognatha toddi Lane and Watson, 1975: 114.

318 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Length of forewing.—Male, 23.5 to 29 mm; female, 28.5 to 33 mm.

This species is known only from Jamaica. Both sexes, male (Fig. 7) and
female (Fig. 8), are easily recognized by the maculation and coloration of
the wings. Five males and one female collected by Davis in Jamaica, in-
cluding the holotype, have been examined. In addition I examined another
male in the British Museum (Natural History) collected before 1973. The
specimens figured in this paper are the ones figured by Lane and Watson,
1975, in figures 11-14.

Hyalurga vinosa (Drury)
Figs. 9-15

Sphinx vinosa Drury, 1773, 1: 47, pl. 23, fig. 4.
Ephestris vinosa (Drury): Hubner, 1820: 178.

Dioptis (Hyrmina) vinosa (Drury): Walker, 1854: 332.
Hyalurga vinosa (Drury): Herrich-Schaeffer, 1866: 131.
Lauron vinosa (Drury): Dewitz, 1877: 95.

Lauron domingonis Butler, 1876: 120.

Dioptis ergolis Walker, 1854: 335.

Hyalurga ergolis (Walker): Hering, 1925: 450.
Nectua horologica Goeze, 1781: 242.

Lauron horologica (Goeze): Kirby, 1892: 411.
Hyalurga horologica (Goeze): Hering, 1925: 450.
Dioptis rica Hubner, 1825: 31.

Hyalurga rica (Hubner), Dewitz, 1879: 170.

The many problems related to the species of this genus occurring in the
Antilles and the proper application of names are mostly unresolved. Bryk
(1931: 41-46) listed 42 species, and Antillean localities are given for four
species. The type of one species, H. rica Htibner, is from Cuba. Genitalia
of the types of all species need to be examined and specific characters, if
any in either sex, should be noted. In addition, the mimetic relationships,
if any, to certain ithomiid butterflies in Central and South America should
be studied. For this species, I have followed previous workers in using the
name H. vinosa (Drury) and accepting certain synonyms; I have not
checked them. A total of 176 specimens was available for study. Food plants
reported for this species by earlier workers are: Heliotropium indicum L.,
H. angiospermum Murray, and Tournefortia hirsutissima L.

Hyalurga halizoa Druce
Fig. 16
Hyalurga halizoa Druce, 1907: 308.

Three specimens of this species from Jamaica have been identified and
studied among material in the National Museum of Natural History in Wash-
ington. The four small white spots composing the transverse band of the

VOLUME 84, NUMBER 2 319

Figs. 9-16. Dorsal view of adults. 9-15, Hyalurga vinosa variants. 9, From Cuba. 10, From
Cuba. 11, From Puerto Rico. 12, From Jamaica. 13, From Puerto Rico. 14, From Dominica.
15, From Jamaica. 16, H. halizoa, Jamaica.

forewing and the orange scaling over much of the forewing make this species
easy to recognize. I have not studied the genitalia.

Hyalurga diastilha Hering ?
Hyalurga diastilha Hering, 1925: 451, pl. 65, row f.

This species is doubtfully recorded because I have not seen a specimen
that agrees with the illustration in Seitz (1925: pl. 65, row f). That illustration

320 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

shows a very narrow transverse bar on the forewing, but the part next to
the costal margin is bent basad in a crescent shape. In the series of Hyalurga
specimens in the National Museum of Natural History there are eleven
examples with a straight very narrow transverse bar which is not curved
basad just before the costal margin. Unfortunately I have no illustration of
this form for this paper. The female type in the Tring Museum needs to be
checked to determine whether the illustration in Seitz is correct, if both
wings are marked the same way, and whether the ferhale genitalia agree
with genitalia of a female with the straight bar from Jamaica in the National
Museum of Natural History. If this identification, based on a female, seems
correct, the male genitalia of the National Museum specimens need to be
examined to learn whether diastilha is different from H. vinosa and H.
halizoa.

Are marginata (Drury)
Bige7

Bombyx marginata Drury, 1773: 38.
Are marginata (Drury): Walker, 1855: 758.

Length of forewing.—Male, 23 to 26 mm; female, 26 to 30 mm.

This species is readily identified by the maculation. The shape of the black
discal spot and the pale costal margin of the forewing are characteristic.
The sexes are marked the same, but the females are slightly larger. I have
examined 42 specimens, all from Jamaica. The subfamily placement of this
genus and species needs to be checked.

Ctenuchidia virgo (Herrich-Schaeffer)
Figs. 18, 19

Ctenucha virgo Herrich-Schaeffer, 1855, 1: fig. 301.
Ctenuchidia virgo (Herrich-Schaeffer): Grote, 1865: 227.

Length of forewing.—Male, 16 mm; female, 19 mm.

I have not resolved any problems connected with this genus. I have seen
only 37 specimens, 28 of them from Puerto Rico. Other examples are from
Cuba and Jamaica. More material needs to be studied, especially examples
from other islands. Genitalia should be studied, and also an attempt should
be made to determine if there exists differences in maculation in the different
island populations.

Ctenuchidia fulvibasis Hering ?
Crenuchidia fulvibasis Hering, 1925: 447.
Centuchidia virgo fulvibasis Hering: Forbes, 1930: 42.

I have not identified this name. The male type in the Berlin Museum needs
to be examined and the genitalia studied. It is quite possible that Forbes

VOLUME 84, NUMBER 2 321

Figs. 17-24. Dorsal view of adults. 17, Are marginata, Jamaica. 18, Ctenuchidia virgo,
male. 19, C. virgo, female. 20, Composia credula, male. 21, C. credula, female. 22, Eucyane
gundlachia, male. 23, E. gundlachia, female. 24, Composia fidelissima, male.

322 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

‘
j
|
}

Figs. 25-27. Dorsal view of adults. 25, Composia fidelissima, female. 26, Sphaeromachia
cubana, male. 27, S. cubana, female.

was correct in treating fulvibasis as a race of C. virgo, but I have considered
the specimens I have from Jamaica as typical virgo.

Ctenuchidia interrupta Hering ?
Ctenuchidia interrupta Hering, 1925: 448.

I have not seen examples. The type, a male, is in the Tring Museum,
England.

Composia credula (Fabricius)
Figs. 20, 21

Bombyx credula Fabricius, 1775: 584.
Composia credula (Fabricius): Htibner, 1820: 179.
Phal.{aena\ Noct.{ua] sybaris Cramer, 1775: 112.

Length of forewing.—Male, 24 mm; female, 29 mm.

This species is easily recognized by the maculation, especially the nu-
merous white spots. The species occurs throughout the Greater Antilles,
and south to Brazil on the continent. The continental distribution needs to
be studied and better recorded.

Composia fidelissima Herrich-Schaeffer
Figs. 24, 25
Composia fidelissima Herrich-Schaeffer, 1866: 131.
Composia fidellissima (sic) Herrich-Schaeffer: Kirby, 1892: 190.

VOLUME 84, NUMBER 2 323

Phaloesia olympia Butler, 1871: 290.

Length of forewing.—Male, 23 mm; female, 33 mm.

This species is easily identified by the maculation, as illustrated. The two
sexes are much alike, but the female is much larger. This species apparently
occurs from southern South America (Uruguay) into Central America (Hon-
duras and Mexico) across to Cuba, Florida, and the Bahama Islands. Gow-
dey (1926: 57) reported the species from Jamaica.

Sphaeromachia cubana (Herrich-Schaeffer)
Figss:' 26,627

Pericopis cubana Herrich-Schaeffer, 1866: 131.
Sphaeromachia cubana (Herrich-Schaeffer): Grote, 1867: 304.

Length of forewing.—Male, 30 mm; female, 36 mm.

This species is recognized by the maculation, coloration, and size. The
sexes are dimorphic, the pale areas of the wings being of a different color.
In the male the color is dark yellow or yellowish orange, in the female, pale
whitish green. The tip of the abdomen of the male is black, but not so in
the female. The trivial name implies the species occurs in Cuba. It also
occurs in Mexico. I examined only five specimens.

Eucyane gundlachia Schaus
Figs.22,.23

Eucyane gundlachia Schaus, 1904: 138.

In the original description Schaus stated ‘*. . . quite similar to Napata
chalybea Hbn., except in neuration.’’ It does look like a species of Cte-
nucha. Two syntypes from Baracoa, Cuba are figured in this work. The
subfamily placement should be checked.

LITERATURE CITED

Bryk, F. 1931. Pericopinae. Jn Strand, Lepidopterorum Catalogus, Pt. 45, Berlin: Junk. 57 pp.
Butler, A. G. 1871. Descriptions of some new species of Lepidoptera. Ann. Mag. Nat. Hist.
(4) 8: 282-291.

. 1876. On the Lepidoptera referred by Walker to the genus Dioptis of Hiibner. Cistula

Entomol. 2: 107-121.

Cramer, P. 1775. Di uitlandsche kapellen voorkomende in de drie waerelddeelen Asia, Africa
en America (Papillons exotiques des trois parties du monde, l’Asie, l'Afrique et
lAmerique) V. 1. Amsterdam: Baalde and Utrecht: Wild. 155 pp., 96 pls. pp. 1-132,
1775; pp. 133-155, 1776.

Dewitz, H. 1877. Dammerungs—und Nachtfalter von Portorica, gesammelt von Herrn Consul
Krug. Mitt. Muench. Entomol. Ver. 1: 91-96.

—. 1879. Naturgeschichte cubanischer Schmetterlinge/Nach beobachtungen des Herrn
Dr. Gundlach bearbeitet. Z. Ges. Naturwiss. 52: 155-174, 1 pl.

Druce, H. 1907. Descriptions of new species of Heterocera belonging to the families Syn-
tomidae, Hypsidae, Cyllopodidae, Dioptidae and Eratinidae. Ann. Mag. Nat. Hist. (7)
19: 299-371.

324 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Drury, D. 1773. Illustrations of Exotic Entomology. Vol. 2. London: Bohn. 90 pp. + ix, 50
pls.

Fabricius, J. C. 1775. Systema Entomologiae . . . , Lipsiae Kovte. 832 pp. + xxx.

Forbes, W. T. M. 1930. Insects of Porto Rico and the Virgin Islands—Heterocera or Moths
(excepting the Noctuidae, Geometridae and Pyralidae). Scientific Survey of Porto Rico
and the Virgin Islands, V. 12, pt. 1. New York: New York Academy of Sciences.
Pericopidae, pp. 40-43.

Goeze, J. A. E. 1781. Entomologische Beytrage zu des Ritter Linne zueliten Ausgabe des
Natursystems. Leipzig. Weidmann. Vol. 3, Theil 3, 439 pp.

Gowdey, C. C. 1926. Catalogus Insectorum Jamaicensis. Jamaica Dep. Agric. Entomol. Bull.
No. 4, Part 1, pp. 1-114.

Grote, A. R. 1865. Notes on the Bombycidae of Cuba. Proc. Entomol. Soc. Phil. 5: 227-255.

. 1867. Notes on the Zygaenidae of Cuba. Proc. Entomol. Soc. Phil. 6: 297-329.

Herrich-Schaeffer, G. A. W. 1855. Sammlung neuer oder wenig bekannter Ausseuropaischer
Schmetterlinge. Regensburg, Manz. Plates 49-78. Figs. 259-451.

—. 1866. Schmetterlinge aus Cuba. Corresp.—Blatt. Zool._Min. Ver. Regensberg 20:
131-132.

Hering, M. 1925. In Seitz, Die Gross-schmetterlinge der Erde, Pericopinae. 6: 423-455, pl.
60-65.

Hiibner, J. 1816 [1826]. Verzeichniss bekannter Schmettlinge (sic), pp. 1-431 + 1-72.

. 1825 [1827-1831]. Zutrage zur Sammlung Exotischer Schmetterlinge. Drittes Hundert.

Augsburg [no publisher listed]. 40 pp. + 8 unnumbered pages, 33 plates [1824-1825].

Figs. 401-600.

Kirby, W. F. 1892. A synonymic catalogue of Lepidoptera Heterocera. London: Gurney and
Jackson. 951 pp.

Lane, M. A. and A. Watson. 1975. A revision of the genus Stenognatha Felder (Lepidoptera:
Arctiidae: Pericopinae). J. Nat. Hist. 9: 107-117.

Neumoegen, B. 1890. New beauties from near and far. Entomol. Am. 6: 61-64.

Schaus, W. 1904. New species of American Heterocera. Trans. Am. Entomol. Soc. 30:

135-178.

. 1938. On new species of American Heterocera. Ann. Mag. Nat. Hist. (11) 2: 506-509.

Walker, F. 1854. List of the specimens of lepidopterous insects in the collection of the British

Museum, Part 2. London: Trustees British Museum. pp. 279-581.

. 1855. List of the specimens of lepidopterous insects in the collection of the British

Museum, Part 3. London: Trustees British Museum. pp. 583-774.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 325-331

A KEY TO THE SPECIES IN HYALOMYZUS
(HOMOPTERA: APHIDIDAE) IN NORTH
AMERICA, WITH THE DESCRIPTION
OF A NEW SPECIES!

CLYDE F. SMITH

Department of Entomology, North Carolina State University, Raleigh,
North Carolina 27650.

Abstract.—Keys are given for the apterous and alate viviparae of Hyalo-
myzus Richards known to occur in North America and Puerto Rico. All of
the morphs of Hyalomyzus mitchellensis, n. sp., are described. They were
collected on Hypericum mitchellianum Rydberg at Mt. Mitchell, N.C.

Richards (1958) described the genus Hyalomyzus and characterized it as
follows: Alate viviparae with many secondary rhinaria on antennal segments
Ill, IV, and V, and without a dorsal sclerotic patch on the abdomen; ap-
terous vivipara with the dorsum of the abdomen strongly wrinkled on seg-
ments I-IV; cornicles swollen, generally imbricated. Other characters are
similar to Myzus Passerini. Nielsson and Habeck (1971) discussed the re-
lationship of Hyalomyzus with closely related genera, and placed H. collin-
soniae (Pepper, 1950) as a synonym of H. eriobotryae Tissot (1935). Eastop
and Hille Ris Lambers (1976) and Smith and Parron (1978) considered H.
collinsoniae to be a valid species.

Measurements are in mm in the following description and key.

Hyalomyzus mitchellensis Smith, NEw Species
Figs. 1A—D, 2A
Fundatrix (Fig. 1A).—Color of living material, yellowish green, Ist-instar
nymphs, yellowish. Color of cleared specimens, dusky on antenna beyond
mid-section of antennal segment III, tip of tibiae, tarsi, siphunculi, cauda,
anal plate, sclerotic areas on integumental sutures of abdomen.
Measurements (4 specimens).—Body length, 1.45-1.77; head width,

1 Paper number 6939 of the Journal Series of the North Carolina Agricultural Research
Service, Raleigh, N.C. 27650.

326 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(.33-0.40; length antennal segment III, 0.19-0.25, IV, 0.11-0.17, V,
0.15—0.16, VI, 0.09-0.13 + 0.11-0.16; rostral IV + V, 0.08—0.09, metatibia,
(.61—0.68: metatarsomere II, 0.08-0.11; siphunculus, 0.31-0.36; cauda,
0.16—0.17.

Some specimens collected May 12, 1969, with 2-6 sensoria on metatibia.
Antenna without secondary rhinaria, head rugose with only slight indication
of antennal tubercles. Siphunculi slightly imbricated. Cauda bearing 4 setae.
Tarsal chaetotaxy, 3-3-3.

Apterous vivipara (Fig. 1B).—Color of living specimens, pale to dark
green with conspicuous black siphunculi. Cleared specimens, dark on an-
tennal segments IV, V, and VI, siphunculi, cauda, anal plate, tip of tibiae,
and metatarsomere II. Dusky on head, rostrum, and sclerotic areas on in-
tersegmental areas of dorsum of abdomen.

Measurements (first measurement is that of holotype) (range of 9 speci-
mens).—Body length, 1.35 (1.34-1.90); head width, 0.36 (0.36—0.42); length
of antennal segment III, 0.26 (0.21-0.31), IV, 0.20 (0.17-0.28), V, 0.17
(0.15—0.23), VI, 0.10 (0.09-0.11) + 0.20 (0.18—0.21); rostral IV + V, 0.08
(0.06—0.09); metatibia, 0.66 (0.62—0.88); metatarsomere II, 0.09 (0.08—0.11);
siphunculi, 0.33 (0.31-0.39); cauda, 0.17 (0.12—0.18).

Cauda bearing 4 (3-5) setae, tarsal chaetotaxy 3-3-3. Abdomen strongly
wrinkled on dorsal surface.

Alate vivipara.—Color of cleared specimens, dark on head, antenna, legs,
thorax, siphunculi, cauda, and anal plate. Dusky sclerotic areas on inter-
segmental area on dorsum of abdomen.

Measurements (4 specimens).—Body length, 1.42—1.58; head width,
0.36-0.46; length antennal segment III, 0.43-0.50, IV, 0.28-0.32, V,
0.21-0.26, VI, 0.12-0.13 + 0.22-0.28; rostral IV + V, 0.07-0.09; metatibia,
0.75-0.98; metatarsomere II, 0.09-0.11; siphunculi, 0.30—-0.33; cauda,
0.14-0.17. Secondary rhinaria on antennal segment III, 19-28, IV, 13-21,
V, 3-8.

Cauda with 4-5 setae, tarsal chaetotaxy 3-3-3 (2). Cubitus of hindwing
present or absent. Venation of forewing (Fig. 2A), dusky but very slight
fuscus area on margins of veins.

Apterous male (Fig. 1D).—Color of living specimens pale, dark green to
brownish, dark on tip of abdomen; siphunculi brown black. Color of cleared
specimens dusky to dark on head, antenna, legs, siphunculi, cauda, anal
plate, and sclerotic areas on abdomen.

Measurements (4 specimens).—Body length, 0.65-0.88; head width,
0.22-0.28; length antennal segment III, 0.13-0.18, IV, 0.09-0.11, V,
0.09-0.11, VI, 0.07-0.08 + 0.11-0.13; rostral IV + V, 0.06; metatibia,
0.35—0.41; metatarsomere I, 0.06-0.08; siphunculi, 0.19-0.20.

Antennal segment III with 7-13 secondary rhinaria, IV, 4-8; V, 2-3; VI,
0. Antennal segment HI and IV may not be distinctly separated. Dorsum of
abdomen slightly wrinkled.

VOLUME 84, NUMBER 2 327

Fig. 1. Hyalomyzus mitchellensis from Hypericum mitchellianum, Mt. Mitchell, N.C. (all
figures the same magnification). A, Fundatrix, collec. 69-111, May 12, 1969. B, Apterous

vivipara (holotype), collec. 79-4, June 19, 1979. C, Apterous ovipara, collec. 67-417, October
11, 1967. D, Apterous male, collec. 67-417, October 11, 1967.

328 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Apterous ovipara (Fig. 1C).—Color of living specimens, dark green to
brownish, dark on tip of abdomen, siphunculi brownish black. Color of
cleared specimens, similar to apterous male except abdomen usually not as
dark and sclerotic areas on intersegmental areas on dorsum of abdomen
prominent.

Measurements (5 specimens).—Body length, 0.96—-1.32; head width,
0.29-0.31; length antennal segment III, 0.10-0.17, IV, 0.09-0.12, V,
0.08—0.12, VI, 0.07-0.08 + 0.12-0.15; rostral IV + V, 0.06—0.07; metatibia,
0.37-0.43; metatarsomere II, 0.07-0.09; siphunculi, 0.20-0.27; cauda,
0.09-0.11.

Antennae without secondary rhinaria. Metatibia with numerous sensoria
on the basal 34, sensoria confined primarily to the ventral portion. Cauda
with 4 setae.

Eggs.—Deposited on tips of shoots. Eggs white when first deposited,
becoming dark olive green to black later.

Type-locality —Mt. Mitchell, North Carolina.

Types.—Holotype from collection 79-4. Apterous specimen no. 4 at 6
o'clock position on slide, collected 20 June 1979 on Hypericum mitchellian-
um Rydberg, Mt. Mitchell, North Carolina by C. F. Smith and C. K. Smith.
All other specimens listed below are paratypes. The holotype and paratypes
are in the National Museum of Natural History, Washington, D.C. (USNM);
paratypes in the collections of USNM; Canadian National Collection, Ot-
tawa; British Museum (Natural History), London; North Carolina State
University; and the author.

Collections.—All known specimens of this species were collected on Hy-
pericum mitchellianum, on Mt. Mitchell, North Carolina along the “‘Camp
Alice Trail,’ 9 September 1963, collection 63-64, C. F. Smith, C. K. Smith,
J. O. Pepper, A. N. Tissot (2 slides); 11 October 1967, collection 67-417, C.
F. Smith (7 slides); 10 September 1968, collection 68-208, C. F. Smith, M.
G. Robertson (1 slide); 12 May 1969, collection 69-11, C. F. Smith, M. G.
Robertson (15 slides); 11 June 1969, collection 69-168, M. G. Robertson, C.
F. Smith (11 slides); 11 June 1976, C. F. Smith, collection 76-39 (18 slides);
12 August 1976, collection 76-46, C. F.Smith, M. Cermeli (10 slides); 20
September 1976, collection 76-56, C. F. Smith, C. K. Smith (5 slides); 10
August 1978, collection 78-258, C. F. Smith, C. K. Smith (3 slides); 12
October 1978, collection 78-260, C. F. Smith, C. K. Smith (4 slides); 12
October 1978, collection 78-261, C. F. Smith, C. K. Smith (1 slide); 19 June
1979, collection 79-4, C. F. Smith, C. K. Smith (32 slides).

Biology.—Hyalomyzus mitchellensis apparently spends its entire life
cycle on Hypericum mitchellianum at the base of leaf axels and flower buds,
and on stems of its host under rocks. Alate viviparae were abundant June
1979, at the time of the other collections alates were absent or very scarce.
Males and oviparous females were collected 11 October. Fundatrices were

VOLUME 84, NUMBER 2 329

~~,

mom

Fig. 2. Wings of alate viviparae. A, Hyalomyzus mitchellensis from Hypericum mitchel-
lianum, Mt. Mitchell, N.C., collec. 79-4, June 19, 1979. B, H. collinsoniae from crab apple
(Pyrus augustifoliae), Laurel Springs, N.C., collec. 6-247, July 4, 1966.

collected on 12 May, some specimens had slightly swollen metatibia bearing
2-6 sensoria.
Etymology.—Named for Mt. Mitchell, the type-locality.
Discussion.—The alate and apterous viviparae of Hyalomyzus mitchel-
lensis can be separated from other species of Hyalomyzus by characters
given in the key. I do not have specimens of the other morphs of the other
species of Hyalomyzus.

KEY TO APTEROUS VIVIPARAE OF HYALOMYZUS

1. | Processus terminalis shorter than antennal segment III .......... 2
- Processus terminalis equal to or longer than antennal segment III

aie! (et Chet a elie) etches ice) wc wv a 6 8 © 8.0. 6 6, 6) ens © Skee fe afulu a ta! O16 6 e 8 is R78, eh Ue ie See BAe ee Bp ee ae

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

_ Abdomen without sclerotic spots. Siphunculi and antennal seg-

ments IV—VI not distinctly darker than base of antennal segment
III. On Crataegus, Aster, Hypericum ... sensoriatus (Mason) (1940)
Abdomen with sclerotic spots (Fig. 1B). Siphunculi and antennal
segments IV—VI distinctly darker than base of antennal segment
Ill (Fig. 1B). On Hypericum mitchellianum pe 2 eee
Me SOA es sraintna: Bast mitchellensis, new species

. Siphunculi dark, at least distal 42. Not on Monarda ............. 4
Sipuncult pale. On Monarda 7 7.5.2.0. -- monardae (Davis) (1911)
WNOE On JUSSIGEG: 65 sacird cnn Dae Se Oe ate to oe 3
On) JUSSIMCE a 2s Ailes Leper ee jussiaeae Smith (1960)
. Antennal segments I and II pale. Not on Collinsonia ............ 6

Antennal segments I and II dark. On Collinsonia .............
ee Re See eh a Pee ee RM Re ccc collinsoniae Pepper (1950)

. Length of antennal segment III, 0.27-0.40, IV, 0.18-0.29, V,

0.202028, VIE ‘0: 1020.13" 290.49 = 0 ce. ea
Br UE RU ae ae faaiee ae ee oer tissoti Nielsson and Habeck (1971)
Length of antennal segment III, 0.40-0.50, IV, 0.33-0.43, V,
0:2820:33> VIE 0: N=0s13s-- 046-055) 2a. eriobotryae Tissot (1935)

KEY TO ALATE VIVIPARAE OF HYALOMYZUS

Processus terminalis less than 3x length of base of antennal seg-

Ment VT ois dees nid a Se eehtere A ew py
Processus terminalis more than 3x length of base of antennal
Ssepment VIO s. 02. cea a a ciSe gals tie nie ee ieee ieee ee 3
. Secondary rhinaria on antennal segment III, 19-23, IV, 15-18, V,
3-5. On Hypericum mitchellianum ..... mitchellensis, new species
Secondary rhinaria on antennal segment III, 45-62, IV, 27-43, V,
11-17. On Crataegus, Aster, Hypericum ...... sensoriatus (Mason)

. Alate vivipara with antennal segment V with 0-11 secondary rhi-

naria. Anal vein of forewing may or may not be distinctly bor-
dered: (Pigs 2), fviciy Ape suk OR oe aio crate eee ie Cee eee 4
Alate vivipara with antennal segment V with 10-20 secondary
rhinaria. Anal vein of forewing not distinctly bordered ........
Beas « d ivglay Megalo Bip <cqactiteet eet Benes ale pike tees aes eriobotryae Tissot

. Antennal segment III usually with 20 or more secondary rhinaria.

Noton Monarda 5... 22+ Pinna gded ont sa ene 2 a eee 5)
Antennal segment III with 8-12 secondary rhinaria. On Monarda
54513 Sea Ee ERR nee DE a ee ch SR monardae (Davis)

. Antennal segment III with 11-40 secondary rhinaria. Anal vein of

forewing not distinctly borderediay. 2G. heel te ee eee 6
Antennal segment III with 40-52 secondary rhinaria. Anal vein of
forewing distinctly bordered (Fig. 2B) ........ collinsoniae Pepper

VOLUME 84, NUMBER 2 331

6(5). Antennal segment III with 11-25 secondary rhinaria. On Cratae-

BUS eSPp=, Lsnerdia intermedia, Drosera Spo. ecn acs. < 04 eee
eee ie ee ees ese tre atta tissoti Nielsson and Habeck

_ Antennal segment III with 26-40 secondary rhinaria. On Jussiaea
BT erage Cee erate tom Seth ne ae hee ee eee jussiaeae Smith

LITERATURE CITED

Davis, J. J. 1911. Williams’ ‘“‘The Aphididae of Nebraska;’’ a critical review. Univ. Stud.
(Nebr.) 11: 253=291.

Eastop, V. F. and D. Hille Ris Lambers. 1976. Survey of the World’s Aphids. Dr. W. Junk
b.v., The Hague. 573 pp.

Mason, P. W. 1940. A revision of the North American aphids of the genus Myzus. U.S. Dep.
Agric. Misc. Publ. 371: 1-30.

Nielsson, R. J. and D. H. Habeck. 1971. The genus Hyalomyzus (Homoptera: Aphididae),
with the description of a new species. Ann. Entomol. Soc. Am. 64: 883-887.

Pepper, J. O. 1950. Six new aphids from Pennsylvania. Fla. Entomol. 33:3-15.

Richards, W. R. 1958. A new aphid genus (Homoptera: Aphididae). Fla. Entomol. 41: 169-172.

Smith, C. F. 1960. A new species of Aphidae: Homoptera from Puerto Rico. J. Agric. Univ.
Puerto Rico 44: 157-162.

Smith, C. F. and C. S. Parron. 1978. An annotated list of Aphididae (Homoptera) of North
America. N.C. Agric. Exp. Stn. Tech. Bull. No. 255: 1-427.

Tissot, A. N. 1935. A new Myzus from Florida. Fla. Entomol. 18: 49-52.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 332-336

TWO NEW SPECIES OF NEPHASPIS CASEY
(COLEOPTERA: COCCINELLIDAE) FROM
TRINIDAD AND COLOMBIA

RoBERT D. GORDON

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Two new species, Nephaspis bicolor and N. namolica, are
described from Trinidad and Colombia, and reference is made to whitefly
predation by members of the genus. Nephaspis amnicola Wingo is reported
as being established in Hawaii.

Nephaspis is a Western Hemisphere genus first recognized by Casey
(1899). Gordon (1972) discussed the generic affinities and provided a key to
the four species recognized at that time. One additional species was later
described from Trinidad and a revised key to the species prepared (Gordon,
1978). Since then, two more undescribed species have been discovered, and
they are added to the classification herein. One of these species is native to
Trinidad, but has been introduced into Hawaii in an effort to control the
spiraling whitefly, Aleurodicus dispersus Russell. A request from S. Higa,
Plant Pest Control Branch, Department of Agriculture, Honolulu, Hawaii,
for a determination of this species of Nephaspis led to the realization that
it was undescribed. Another undescribed species of Nephaspis was received
from Fred Bennett of the Commonwealth Institute of Biological Control,
Curepe, Trinidad. These specimens were collected in Colombia feeding on
a species of whitefly on guava.

In addition to the new species now being released in Hawaii, Nephaspis
amnicola Wingo now occurs there, having been introduced into Hawaii from
Honduras and Trinidad in 1979 for control of the spiraling whitefly.

Type-materials of the two new species are deposited in the collection of
the National Museum of Natural History, Washington, D.C. (USNM) and
that of Fred Bennett, CIBC.

I thank Linda Lawrence, Systematic Entomology Laboratory, USDA,
staff artist, for preparing the illustrations used herein.

333

VOLUME 84, NUMBER 2

oes

a ET

Male genitalia of Nephaspis bicolor.

Figs. 1-3.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

nm

Male genitalia of Nephaspis namolica.

Figs. 4-6.

VOLUME 84, NUMBER 2 335

Nephaspis bicolor Gordon, NEw Species
Figs. 1—3

Holotype.—Male, length 1.40 mm, width 1.0 mm. Black; head, pronotum,
mouthparts, legs, and apical 2 abdominal sterna yellow. Head finely punc-
tured, punctures separated by I-3x a diameter; pronotal punctures equal
in size to head punctures, separated by a diameter or less. Ventral surface
coarsely punctured, punctures separated by a diameter or less medially,
contiguous laterally. Genitalia as in Figs. 1-3.

Allotype.—Female, length 1.45 mm, width 1.05 mm. Similar to male ex-
cept head black, pronotum black except lateral 4% yellow.

Variation.—Length 1.40 to 1.50 mm, width 1.0 to 1.10 mm. Head of fe-
male often entirely yellow with vertex black.

Type-material.—Holotype (USNM 100004) and 10 paratypes; Trinidad,
St. Joseph, Apr 16, 1980, R. Burkhart, on guava. Allotype and 5 paratypes;
Trinidad, Curepe, May 28, 1980, R. Burkhart, A. dispersus on Guava and
Candlestick.

Comments.—This species has male genitalia most similar to those of N.
amnicola Wingo, and it will key to that species in Gordon (1978). However,
the sipho of N. bicolor is straight before the apex, not sinuate as in N.
amnicola. The male color pattern is not diagnostic for this species in any
way, but the female pronotum with the wide yellow area on each side is
unique and will separate that sex from females of all other species. The
specific epithet refers to the pronotal color pattern of the female.

Nephaspis namolica Gordon, NEw SPECIES
Figs. 4—6

Holotype.—Male, length 1.40 mm, width 0.95 mm. Black; pronotum yel-
low with median black spot at base; head, mouthparts, legs, and apical 3
abdominal sterna yellow. Head finely punctured, punctures separated by
1-3 x a diameter; pronotal punctures slightly larger than on head, separated
by | or 2 times a diameter; punctures on elytron coarse, separated by less
than a diameter to twice a diameter. Ventral surface coarsely punctured,
punctures separated by less than a diameter. Genitalia as in Figs. 4-6.

Allotype.—Female, length 1.35 mm, width 0.95 mm. Similar to male ex-
cept vertex of head black, pronotum black except anterolateral angle nar-
rowly yellow.

Variation.—Length 1.35 to 1.45 mm, width 0.95 to 1.0 mm. Black spot
on male pronotum confined to basal 2, or extending to midpoint of prono-
tum.

Type-material.— Holotype (USNM _ 100005), allotype, and 10 paratypes;
Colombia, Palmira (V), IV-79, F. Garcia, hojas guayobo (leaves of guava),
predator mosca blanca (whitefly predators).

Comments.—Nephaspis namolica has male genitalia very similar to those

336 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of N. amnicola and will key to that species in Gordon (1978). The dorsal
surface of the basal lobe in N. namolica is strongly sinuate in lateral view,
and the shape of the basal lobe in ventral view is broader and more abruptly
tapered before the apex. In N. amnicola, the dorsal surface of the basal
lobe is feebly sinuate, and the shape of the basal lobe in ventral view is
slender and tapered to the apex. The specific epithet is an anagram.

LITERATURE CITED

Casey, T. L. 1899. A revision of the American Coccinellidae. J. N.Y. Entomol. Soc. 7: 71-169.

Gordon, R. D. 1972. A review of the genus Nephaspis Casey and a comparison with the genus
Clitostethus Weise (Coleoptera: Coccinellidae). Rev. Agric. (Piracicaba) 47: 145-154.

———. 1978. West Indian Coccinellidae II (Coleoptera): some scale predators with keys to
genera and species. Coleopt. Bull. 32: 205-218.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 337-341

NEW SYNONYMIES AND A NEW COMBINATION IN THE
NORTH AMERICAN MIRIDAE (HEMIPTERA)

THOMAS J. HENRY

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Four species of Miridae (Hemiptera) from the eastern United
States are shown to be synonyms: Phytocoris discoidalis Henry, 1974 = P.
dreisbachi Knight, 1974; Plagiognathus inopinus Knight, 1926 = P. albatus
Van Duzee, 1915; Rhinocapsus miniatus Knight, 1923 = R. rubricans Pro-
vancher, 1887; and Texocoris secludis Schaffner, 1974 = Parthenicus nigrel-
lus Knight, 1939. The monotypic genus Texocoris Schaffner, 1974 is retained
to accommodate nigrellus (new combination).

A forthcoming paper on the Miridae associated with ericaceous plants in
eastern North America has prompted me to provide information showing
that Rhinocapsus miniatus Knight is a junior synonym of R. rubricans (Pro-
vancher). Additional synonymies and a new combination involving three
other species from the eastern United States are also presented. Unless
otherwise noted, all specimens examined are deposited in the collections of
the Pennsylvania Department of Agriculture, Harrisburg, or the National
Museum of Natural History, Washington, D.C.

Phytocoris dreisbachi Knight

Phytocoris dreisbachi Knight, 1974: 125; Kelton, 1980: 177.
Phytocoris discoidalis Henry, 1974: 187; Henry, 1979: 9. New Synonymy.

Knight (1974) described 15 new species of the Phytocoris junceus group.
That same year I described the new species discoidalis (Henry, 1974). Be-
cause my species belonged to the junceus group, I used Knight's key and
found that discoidalis runs to couplet 4 with dreisbachi. Recently, after
examining Knight’s type-specimens, I found that discoidalis is conspecific
with dreisbachi. Because Knight’s description appeared a few months prior
to mine, I must recognize P. discoidalis as a junior synonym of P. dreis-
bachi.

Phytocoris dreisbachi, originally described from Michigan, is now known

338 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

to occur in Pennsylvania, Virginia, Wisconsin (Henry, 1979), and Alberta
and Manitoba (Kelton, 1980). This species has been collected on Pinus
banksiana Lamb. and P. virginiana Mill.

Plagiognathus albatus (Van Duzee)

Psallus albatus Van Duzee, 1915: 116.

Plagiognathus albatus: Van Duzee, 1916: 46; Van Duzee, 1917: 410; Knight,
1923: 445; Blatchley, 1926: 943; Knight, 1941: 36; Froeschner, 1949: 160;
Davis, 1955: 132; Carvalho, 1958: 94; Wheeler and Henry, 1977: 153;
Wheeler, 1980: 354 (see for review of economic literature).

Plagiognathus inopinus Knight, 1926: 11; Blatchley, 1926: 942; Carvalho,
1958: 103. NEw SYNONYMY.

Plagiognathus albatus often has been implicated as a pest of sycamore
or American plane, Platanus occidentalis L. Wheeler (1980) studied the life
history and gave evidence that P. albatus caused necrotic lesions and even-
tual holes in the foliage of London plane, Platanus x acerifolia. This com-
mon eastern North American mirid occurs wherever Platanus species grow
(Wheeler, 1980).

Plagiognathus inopinus, the only other member of this genus recorded
from Platanus spp. (Knight, 1926), was found to be common on street plant-
ings of London plane at Harrisburg, Pennsylvania (the type-locality for this
species). Having had the advantage to collect in Harrisburg for eight
years, I was able to observe a close relationship between inopinus and
albatus. Plagiognathus inopinus is a uniformly dark species; P. albatus
is largely pale with darker markings. Continued collecting in this area
showed that besides the typically dark inopinus and the much lighter colored
albatus, at least three intermediate color morphs were present concurrently.
All morphological structures are the same, including color and lengths of
the antennal segments and the distinct spots on the pale femora. Male gen-
italia offer no distinctions. I have examined Knight’s holotype and conclude
that inopinus is only a dark extreme of albatus.

Rhinocapsus rubricans (Provancher)

Plagiognathus rubricans Provancher, 1887: 154.

Microphylellus rubricans: Van Duzee, 1912: 323.

Rhinocapsus rubricans: Van Duzee, 1917: 409; Carvalho, 1958: 140; Kelton,
1968: 1075 (lectotype designation); Kelton, 1980: 306.

Rhinocapsus miniatus Knight, 1923: 460; Blatchley, 1926: 926. New Syn.
ONYMY.

Rhinocapsus miniatus, described from a single male taken at Lakehurst,
New Jersey, is one of several species of Miridae in North America that has
remained obscure since its original description. Over the past eight years,

VOLUME 84, NUMBER 2 339

Table 1. Comparison of the length of the second antennal segment and the basal width of
the pronotum for specimens of Rhinocapsus rubricans (in mm).

Male Female

Length of Length of

Antennal Basal Width Antennal Basal Width

Segment II Difference of Pronotum Segment II Difference of Pronotum
1.18 Folle 1.06 1.14 + .048 1.10
1.36 +.06 1.30 1522 srl 1.08
1.14 .00 1.14 1.20 +.02 1.18
1.34 +.04 1.30 1.04 = {I 1.06
1.26 +.06 1.20 22 = IP 1.34
1.30 S361? 1.18 1.08 + .04 1.04
1.20 +.08 et 1.18 — .08 1.26
1.30 +.06 1.24 1.18 00 1.18
1.32 +.04 1.28 1.18 a 1.28
1.36 +.06 1.30 1.20 —.08 1.28
1.30 + .06 1.24 1.16 —.10 1.26
ES2 +.02 1.30 1.24 — .04 1.28
1.34 sea, 1.12 1.18 —.08 1.26
1.36 ame IW) 1.26 1.26 — .04 1.30
1.24 +.06 1.18 1.14 +.04 1.10

4 Differences (in mm) of measurements with **+”° indicating that the second antennal seg-
ment is longer than the basal width of the pronotum and **—”’ indicating that the second
antennal segment is shorter.

I have obtained numerous specimens of Rhinocapsus species, including
examples from southern New Jersey. In an attempt to identify this material,
I found that Knight’s (1923: 459) key for separating miniatus from rubricans
is unworkable.

Knight presented only two distinguishing characters for recognizing these
two species: (1) Length of second antennal segment less than or subequal
to basal width of pronotum (rubricans) or second antennal segment distinct-
ly greater than basal width of pronotum (miniatus); and (2) size larger
(4.3-4.6 mm for rubricans) or size smaller (3.4 mm based on the holotype
of miniatus).

In measuring 15 males and 15 females of rubricans, | discovered that, in
all but one specimen, the lengths of male second antennal segments are
greater than the basal width of the pronotum; in females the lengths of the
second segments were, in all but five examples, less than the basal width
of the pronotum (Table 1). For the same number of specimens, males of
rubricans averaged 4.26 mm in body length, with a range of 4.00-4.67 mm,
and females averaged 3.86 mm, with a range of 3.25-4.33 mm.

It is apparent that rubricans males in most cases will key to miniatus and
females to rubricans. Perhaps, in assembling his key, Knight used females
of rubricans for his comparison to miniatus (a male). Also, the size of

340 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

females frequently falls into the small-size range diagnosed for miniatus.
Although no males were found that measured less than 4.00 mm, it is within
the bounds of variation for such smaller specimens to occur, especially if
the more harsh conditions of the Lakehurst-New Jersey Pine Barrens region
are considered. I have remeasured the holotype of miniatus and find it to
be 3.64 mm in length (by my ocular micrometer), rather than 3.4 as given
by Knight.

Knight also commented that miniatus was more uniformly red than ru-
bricans (which is typically fuscous on the hemelytra). An examination of
his holotype suggests that the specimen is teneral; the color is light red and
the metafemora are collapsed. Reared adults (teneral) of rubricans, also, are
lighter or more uniformly reddish. Other than the slight size difference, ten-
eral males of rubricans are indistinguishable from miniatus; thus, I consider
miniatus a junior synonym of rubricans.

Texocoris nigrellus (Knight), NEw COMBINATION

Parthenicus nigrellus Knight, 1939: 23; Froeschner, 1949: 166; Carvalho,
1958: 123, Akingbohungbe et al., 1972: 11; Akingbohungbe et al., 1973:
14; Henry and Smith, 1979: 214.

Texocoris secludis Schaffner, 1974: 283. NEw SYNONYMY.

While examining Miridae in the Texas A&M University insect collection,
I recognized that specimens identified as Texocoris secludis were very sim-
ilar to Parthenicus nigrellus. Further examination of Schaffner’s (1974) fig-
ures of male genitalia and comparison of type-material of both species con-
firmed that sec/ludis is a junior synonym of nigrellus. Because nigrellus does
not belong in the genus Parthenicus Reuter, I find it necessary to retain the
monotypic genus Texocoris Schaffner to accommodate nigrellus.

Texocoris nigrellus has been reported from Georgia, Illinois, lowa, Texas
(Knight, 1941), and Missouri (Froeschner, 1949). Akingbohungbe et al.
(1972) added Wisconsin and listed bloodroot as the host. Recently, I iden-
tified a series of nigrellus collected in Missouri (Columbia, Boone Co., 31
May 1981) on a fruiting mulberry tree, Morus sp.

ACKNOWLEDGMENTS

I thank A. G. Wheeler, Jr. (Bureau of Plant Industry, Pennsylvania De-
partment of Agriculture, Harrisburg) for lending specimens of Plagiogna-
thus albatus and Rhinocapsus rubricans and Robert Blinn (Department of
Entomology, University of Missouri, Columbia) for allowing me to report
his collection of Texocoris nigrellus.

LITERATURE CITED

Akingbohungbe, A. E., J. L. Libby, and R. D. Shenefelt. 1972. Miridae of Wisconsin (He-
miptera: Heteroptera). Univ. Wis.—-Madison Coll. Agric. Life Sci. Res. Div. Res. Bull.
R 2396, 24 pp.

VOLUME 84, NUMBER 2 341

—. 1973. Nymphs of Wisconsin Miridae (Hemiptera: Heteroptera). Univ. Wis.-Madison
Coll. Agric. Life Sci. Res. Div. Res. Bull. R 2561, 25 pp.

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America. Nature Publ. Co.,
Indianapolis, Indiana. 1116 pp.

Carvalho, J. C. M. 1958. Catalog of the Miridae of the world. Arq Mus. Nac., Rio de J. Part
II. Phylinae 45: 1-216; Part III. Orthotylinae 47: 1-161.

Davis, N. T. 1955. Morphology of the female organs of reproduction in the Miridae (Hemip-
tera). Ann. Entomol. Soc. Am. 48: 132-150.

Froeschner, R. C. 1949. Contribution to a synopsis of the Hemiptera of Missouri. Pt. [V. Am.
Midl. Nat. 42: 126-188.

Henry, T. J. 1974. Two new pine-inhabiting Phytocoris from Pennsylvania (Hemiptera: Mi-
ridae). Entomol. News 85: 187-191.

———. 1979. Descriptions and notes on five new species of Miridae from North America
(Hemiptera). Melsheimer Entomol. Ser. 27: 1-10.

Henry, T. J. and C. L. Smith. 1979. An annotated list of the Miridae of Georgia (Hemiptera-
Heteroptera). J. Ga. Entomol. Soc. 14: 212-220.

Kelton, L. A. 1968. On the Heteroptera in the Provancher collection (Hemiptera). Nat. Can.
(Que.) 95: 1065-1080.

—. 1980. The plant bugs of the Prairie Provinces. Agric. Can. Publ. 1703, Ottawa. 408
pp.

Knight, H. H. 1923. Family Miridae (Capsidae), pp. 422-655. Jn Britton, W. E., ed., Hemiptera
of Connecticut. Conn. State Geol. Nat. Hist. Surv. Bull. 34.

—. 1926. Descriptions of four new species of Plagiognathus from the eastern United
States (Hem., Miridae). Entomol. News 37: 9-12.

—. 1939. Three new species of Miridae from North America. Bull. Brooklyn Entomol.
Soc. 34: 21-23.

———. 1941. The plant bugs, or Miridae, of Illinois. Bull. Ill. Nat. Hist. Surv. 22, 234 pp.

. 1974. A key to species of Phytocoris Fallén belonging to the Phytocoris junceus group

of species (Hemiptera, Miridae). lowa State J. Res. 49: 123-135.

Provancher, L. 1886-98. Petite faune entomol. du Canada. Les Hemipteres 3, 354 pp.

Schaffner, J. C. 1974. Texocoris secludis, anew genus and species of Orthotylinae from Texas
(Heteroptera: Miridae). J. Kans. Entomol. Soc. 47: 281-284.

Van Duzee, E. P. 1912. Synonymy of the Provancher collection of Hemiptera. Can. Entomol.
44: 317-332.

—. 1915. New genera and species of North American Hemiptera. Pomona J. Entomol.
Zool. 7: 109-121.

— . 1916. Checklist of the Hemiptera of America north of Mexico. New York (N.Y.

Entomol. Soc.). 111 pp.

. 1917. Catalog of the Hemiptera of America north of Mexico. Univ. Calif. Publ. Ento-

mol. 2: 1-902.

Wheeler, A. G. 1980. Life history of Plagiognathus albatus (Hemiptera: Miridae), with de-
scription of the fifth instar. Ann. Entomol. Soc. Am. 73: 354-356.

Wheeler, A. G., Jr. and T. J. Henry. 1977. Rev. Modestus Wirtner: Biographical sketch and
additions and corrections to the Miridae in his 1904 list of western Pennsylvania He-
miptera. Great Lakes Entomol. 10: 145-157.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 342-348

A CO, PLATFORM TRAP FOR TAKING ADULT
DERMACENTOR ANDERSONI (ACARI: [IXODIDAE)

R. B. Eaps, G. C. SMITH, AND G. O. MAUPIN

Vector-Borne Diseases Division, Center for Infectious Diseases, Centers
for Disease Control, Public Health Service, U.S. Department of Health and
Human Services, P.O. Box 2087, Fort Collins, Colorado 80522.

Abstract.—A simple, light, and inexpensive CO, platform tick trap is
described, and its effectiveness in attracting and collecting Dermacentor
andersoni Stiles is evaluated. Tests utilized 8 traps set equidistant from a
central tick release point. In 1979, traps were placed at distances of 6 m, 9
m, and 12 m on different dates. At 6 m, 76% of the ticks were recovered in
72 h, 40% in the Ist 24 h. At 9 m, only 3% of the released ticks were taken
in 48 h, and 2% were trapped at 12 m in 72 h. In 1980, simultaneous tests
were performed at 6-m and 9-m distances, with an additional 6-m test at a
later date. In the 6-m tests, 50% of the ticks were captured in 24 h and 62%
in 48 h. One of the 6-m tests was continued for 72 h, during which time 82%
of the released ticks were taken. In the 9-m test, tick captures were 2% in
24 h, 15% in 48 h, and 23% in 72 h.

The Rocky Mountain wood tick, Dermacentor andersoni Stiles, is an
important vector and pest species in the Rocky Mountain region. Colorado
tick fever is the most prevalent disease with which the species is associated;
some 150-250 cases are reported each year in Colorado, chiefly in Larimer
and Boulder Counties (Goodpasture et al., 1978). Actual prevalence of the
disease is difficult to assess, as it is considered to be underreported.

Vector-Borne Diseases Division personnel have conducted studies on the
natural history of Colorado tick fever in Colorado. In connection with these
studies, a CO, platform trap has been developed to study the seasonal host-
seeking activities of unfed, adult D. andersoni. Garcia (1962) first demon-
strated the effectiveness of CO, emitted by dry ice in attracting certain
species of ticks in the field. Miles (1968) devised a CO, trap to collect ticks
and fleas from mammal burrows. Wilson et al. (1972) reviewed the literature
dealing with the utilization by ticks of chemoreceptors in host-seeking. They
also described a CO, platform trap effective in collecting all stages of Am-
blyomma americanum (Linnaeus) and demonstrated the advantages of this

VOLUME 84, NUMBER 2 343

Fig. 1. CO, platform trap for ticks.

type of trap over flagging with white flannel cloth to study unfed (flat) tick
populations. Our trap is discussed here, with an evaulation of its effective-
ness in attracting adult D. andersoni.

MATERIALS AND METHODS

The trap (Fig. 1) differs from the Wilson model primarily in the type of
CO.-holding container, and in that it is in two pieces rather than one. A 1-
liter Boston Mailer®! of moulded styrene foam for holding dry ice is inserted
in a central, octagonal opening in a fiberglass shell platform to complete the
trap. The container holds ca. 650 g of chipped dry ice, an amount sufficient
to operate a trap in shade or partial shade for 24 h even at the upper end of
the summer air temperature range in the study area (ca. 32°C).

To activate a trap, a base is anchored firmly in litter or top soil on the
ground. The container halves are filled with small chunks of dry ice and
sealed with masking tape. A charged container is inserted into the central
Opening in a base and 5-cm masking tape is placed around the container.

1 Use of trade names is for identification only and does not constitute endorsement by the
Public Health Service or by the U.S. Department of Health and Human Services.

344 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ale

RELEASE
ae

Ale #8
ae “fhuticecttam

Fig. 2. Design to evaluate the effectiveness of CO, platform traps in attracting adult Der-
macentor andersoni.

Only the proximal 2.5 cm of the tape is attached. The outer 2.5 cm is
unattached and entangles ticks crawling up the sloping sides of the platform.

The design for the placement of the CO, platform traps, under field con-
ditions, to attract adult D. andersoni is shown in Fig. 2. Releases of adult
D. andersoni marked with a small spot of HiGlo fluorescent paint® were
made at the center of the 8 equidistant CO, traps. Separate experiments
were made in 1979 with the traps at 6 m, 9 m, and 12 m from the center.
The ticks were obtained by flagging in the vicinity of the release sites,
marked, and held in the laboratory for a week at 4°C prior to the first release
experiment. Sufficient numbers of ticks were not available to permit simul-
taneous testing of the tick responses at the three distances. The same ticks
were used at all three distances. Between tests they were held at 4°C, with
negligible mortality; diminished vigor was not observed. In 1980, the traps

VOLUME 84, NUMBER 2 345

Table 1. Adult Dermacentor andersoni response to CO, platform traps, 1979.
9 Meters
6 Meters 200 Marked 12 Meters
245 Marked Ticks Released Ticks Released 255 Marked Ticks Released
Marked Recoveries Marked Recoveries Marked Recoveries

Trap May 16 17 18 June 5 June 6 May 22 23 24
Station 24h 48h 72 24h 48h 24h 48h 72h

1 0 4 1 0 | 0 l 0

2 0 17 16 0 I 0 0 0

3 0 18 14 l l 0 0 0

4 0 8 2 l 0 l 0 0

5 0 l 0 0 0 0 0 0

6 l 1 0 0 0 ] 0 0)

7 87 3 3 0 0 0 l 0

8 9 0 1 0 0 l 0 0
Totals 97 52 37 2 3 3 2 0

% available ticks

recaptured 39.6 15)5II 38.5 1.0 1.5 12 0.8 0

were tested simultaneously in different grids at 6 m and 9 m from the release
point, and a third test at 6 m was done 12 days later. Insufficient numbers
of ticks were taken by flagging near the release sites for the tests, and ca.
50% of the test ticks were taken by flagging at the Moraine Campground,
Rocky Mountain National Park. All were marked and held 1-2 weeks at 4°C
prior to the tests.

In each test the 8 traps were charged with CO, ca. 0800, just prior to the
release of the marked ticks. The traps were checked at 24-h intervals at ca.
0800, trapped ticks were removed from the experiment, and the traps were
recharged with CO,. This was repeated for the desired number of days in
each test period.

Temperature and relative humidity at the study site were recorded con-
tinuously during the tests on 7-day charts with a Belfort Hygrothermo-
graph.® An anemometer was available for continuous recording of wind
speed during the last of the tests.

STUDY AREA

The study site, on the Cherokee Christian Ranch, is 40 km NW of Fort
Collins, within the Roosevelt National Forest, along the east slope of the
front range of the Rocky Mountains in north-central Larimer County, Col-
orado (40°53'N lat., 105°28’W long.). Marr (1961) has described front range
ecosystems and Marr et al. (1968) have characterized the climate. Soils are
derived mainly from granite, schist-gneiss, and undivided metamorphic and
igneous rocks, with occasional sedimentary formations at lower elevations

346 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Adult Dermacentor andersoni response to CO, platform traps, 1980.

6M
6 Meters 9 Meters 250 Make

179 Marked Ticks Released 179 Marked Ticks Released Ticks Released
Marked Recoveries Marked Recoveries Marked Recoveries
Trap May 27 28 29 May 27 28 29 June 10 June 11

Station 24h 48h 72h 24h 48h 72h 24h 48h

l 22 l l 0 l 2 9 !

2 13 2 1 0 0 0 2 0

3 20 0 0 0) 0 0 1 1

4 0 0 0 0 0 l i l

5 0 0 0 ] 0 l 1] l

6 l 0 l l 3 2 5 l

7 29 16 5 0 8 4 47 14

8 24 8 2 2 11 4 41 4

Totals 109 Pal 10 4 23 14 107 24

% available ticks

recaptured 60.9 38.6 233 22 Bio 92 42.8 16.8

(Lovering and Goddard, 1950). Dominant plants on the hillsides are pon-
derosa pine, Pinus ponderosa; mountain mahogany, Cercocarpus montan-
us; and bitter-brush, Purshia tridentata.

Cherokee Christian Ranch was chosen for this study because it is a per-
sistent focus of Colorado tick fever. For several years, an annual attack
rate of 3-5 cases per 1000 camp days has been documented (Poland, un-
published data, Vector-Borne Diseases Division Annual Reports 1975-79).
The site at which the ticks were released to test the effectiveness of the CO,
traps was at an elevation of ca. 2179 m (7150 ft.). The immediate release
area is flat, without rocky outcroppings, and with scattered ponderosa pines;
small bitter-brush bushes and short grasses offer little obstruction to tick
dispersal.

RESULTS

The first of the tests of the effectiveness of the CO, traps in attracting D.
andersoni was done for three 24-h trapping periods May 16, 17, and 18,
1979, with the 8 traps placed 6 m from the central release point (Fig. 2).
This point was continuously in the shade of a small ponderosa pine which
provided several inches of litter for tick cover.

As shown in Table |, at 6 m almost 40% of the marked ticks released May
15 were recovered in 24 h. All of the ticks were taken at trap stations 6, 7,
and 8. The 97 captured ticks were removed from the experiment, and the
traps were recharged with CO,. After 48 h, 52 of the remaining 148 ticks

VOLUME 84, NUMBER 2 347

(35%) were captured. Ticks were taken from all stations except number 8. How-
ever, the majority were from stations 1, 2, 3, and 4. After 72 h, 37 of the
remaining 96 ticks (39%) were captured, in essentially the same traps that
took ticks at the 48-h interval. Since 76% of the marked ticks were recovered
at a distance of 6 m in three consecutive trap days, the trap distances were
increased to 12 m on the second test run May 22, 23, and 24. Catches were
poor. Three of 255 were taken after 24 h (1%); two of the remaining 252
(.8%) were trapped after 48 h; no ticks were on the traps after the third
24-h trapping period.

With only 5 of 255 (2%) ticks recovered at the 12-m distance in three
consecutive trap days, a third test was run at 9 m June 5 and 6. Results
differed little from those obtained at the 12-m distance. Five ticks of the 200
marked specimens released (3%) were trapped.

Additional evaluation studies were made of the CO, platform traps in 1980
in the same study area. Simultaneous releases were made at distances of 6
m and 9 m May 26, and trapping was conducted May 27, 28, and 29. At 6
m, 109 of 179 ticks (61%) were recovered in 24 h; 27 of the remaining 70
(39%) were taken at 48 h; and 10 of the remaining 43 (23%) were on the
traps at 72 h (Table 2). Of the 179 marked ticks released at 6 m, 146 (82%)
were recovered at 6 m in three consecutive trap days. As in 1979, the traps
were recharged with CO, at 24-h intervals.

Four of 179 marked ticks released 9 m from the traps were taken in 24 h
(2%), 23 (13%) of the remaining 175 ticks in 48 h, and 14 (9%) of the re-
maining 152 ticks at 72 h. Forty-one of the 179 ticks released (23%) were
trapped at 9 m in three consecutive trap days.

An additional release with the 6-m traps was made of 250 marked ticks
June 10, and the traps operated for two consecutive trap days. As shown
in Table 2, 107 ticks (43%) were trapped in 24 h, and 24 of the remaining
143 (17%) were taken on the second day (48 h).

Temperature and humidity readings were similar during the tests in both
years, with the days cool and sunny and the nights at or near freezing. The
average daily high was 24°C, the low 3°C, and the mean 13.5°C. Ambient
humidity high averaged 94.6% (night), low 22.3% (day), and the mean
58.4%.

DISCUSSION

This tick trap apparently depends to a considerable extent on the wind to
disseminate the sublimating dry ice. In this respect, the 6-m release May
16-18, 1979, is especially interesting. May 16 the ticks were captured in trap
stations 6, 7, and 8, suggesting the wind was from the east, southeast. May
17 and 18 the ticks were taken mainly in stations 1, 2, 3, and 4, indicating
a wind shift to south, southwest. Wind velocity was recorded only during

348 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the 6-m tests June 10 and 11, 1980. Tick recoveries (Table 2) were obtained
when the wind was light (1.6—4.8 km/h) during the day and gentle (6.4-9.6
km/h) at night.

Comparing the efficacy of the CO, platform trap described here and the
Wilson model in attracting ticks is difficult. Different species of ticks were
tested, and evaluation techniques differed. The Wilson trap released | kg
of dry ice over a 7-day period, while our trap released ca. 650 g in 24 h. In
seven days, the Wilson trap took 5% of the marked A. americanum ticks
released at 6.1 m from the trap, an average of 7.5% at 9.4 m and an average
of 3.5% at 12 m. As shown in Tables | and 2, our trap was much more
effective in taking D. andersoni at release distances of 6 m.

We believe that this CO, platform trap is a useful tool in studying the
host-seeking activities of adult D. andersoni. It is of light weight, and fairly
large numbers can be carried over rough terrain without difficulty. For op-
timal sampling effectiveness, trap stations should probably be 12 m apart,
to avoid competition in attracting ticks.

ACKNOWLEDGMENTS

Jack Poland and the other officers of the Meadow Lark Church of Christ
were most generous in allowing us to carry on this study on church property.
Numerous persons in VBDD have assisted in the field, including Sue Taylor,
E. G. Campos, Richard Bolin, and Leon Carter. Bruce Francy provided
technical assistance throughout the study.

LITERATURE CITED

Garcia, R. 1962. Carbon dioxide as an attractant for certain ticks (Acarina; Argasidae and
Ixodidae). Ann. Entomol. Soc. Am. 55(5): 605-606.

Goodpasture, H. C., J. D. Poland, D. B. Francy, G. S. Bowen, and K. A. Horn. 1978.
Colorado tick fever: Clinical, epidemiologic and laboratory aspects of 228 cases in Col-
orado in 1973-74. Ann. Intern. Med. 88(3): 303-310.

Lovering, T. S. and E. N. Goddard. 1950. Geology and ore deposits of the Front Range,
Colorado. U.S. Geol. Surv. Prof. Paper 223: 1-319.

Marr, J. W. 1961. Ecosystems of the east slope of the Front Range in Colorado. Univ. Colo.
Biol. Ser. 8, Univ. Colo. Press, Boulder, Col. 134 pp.

Marr, J. W., J. M. Clark, W. S. Osburn, and M. W. Paddock. 1968. Data on mountain
environments. III. Front Range, Colorado, four climax regions, 1959-64. Univ. Colo.
Biol. Ser. 29, Univ. Colo. Press, Boulder, Col. 181 pp.

Miles, V. I. 1968. A carbon dioxide bait trap for collecting ticks and fleas from animal burrows.
J. Med. Entomol. 5: 491-495.

Wilson, J. G., D. R. Kinzer, J. R. Sauer, and J. A. Hair. 1972. Chemo-attraction in the lone
star tick (Acarina: Ixodidae): 1. Response of different developmental stages to carbon
dioxide administered via traps. J. Med. Entomol. 9: 245-252.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 349-365

COEXISTENCE IN TWO SPECIES OF HOLCOCEPHALA
(DIPTERA: ASILIDAE) IN A MARYLAND
HABITAT: PREDATORY BEHAVIOR

A. G. SCARBROUGH

Department of Biological Sciences, Towson State University, Baltimore,
Maryland 21204.

Abstract.—Coexistence was studied in two species of Holcocephala in
a Maryland habitat. General behavior patterns of H. abdominalis (Say) and
H. calva (Loew) were similar although each species exhibited significant
variations. Coexistence is possible because the species differed in their se-
lection of prey and foraging and feeding patterns.

The three Nearctic species of Holcocephala, abdominalis (Say), calva
(Loew), and fusca Bromley, are usually found in mesic habitats and range
from the eastern United States west to eastern Texas, Kansas, and Ne-
braska (Martin and Wilcox, 1965). They typically forage from tips of dead
twigs or leaves of rank vegetation, located near or along margins of streams,
fields, and woods (Baker and Fischer, 1975; Brimley, 1922; Bromley, 1931,
1946, 1950a, b; McAtee and Banks, 1970; Hull, 1962; Scarbrough, 1974;
Johnson, 1976; Dennis, 1979). They sometimes forage from edges of wooded
areas where vegetation patterns change abruptly, e.g. clearings, trails, and
streams, but typically avoid dark thickly vegetated areas of forest (Scar-
brough, 1974; Dennis, 1979). Holcocephala abdominalis is sometimes found
foraging in dry fields, a short distance from streams (Scarbrough, 1974) and
forest margins (Johnson, 1976), especially when the fly is abundant. Hoi-
cocephala calva and H. fusca appear to restrict selection of perches to
vegetation along tree lined margins of trails, footpaths, lawns, and streams
(Scarbrough, 1974; Dennis, 1979).

Holcocephala abdominalis and H. calva are sometimes found within a
common habitat, with the former species being apparently more abundant
(Baker and Fischer, 1975; Bromley, 1931; McAtee and Banks, 1920). Dennis
(1979) suggested that differences in population densities of the two coexist-
ing species may result from interspecific competition. Coexistence between
interacting populations occurs when the species involved exploit resources
differentially, with their realized niches being dissimilar in at least one im-

350 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

portant dimension (MacArthur, 1958; DeBach, 1966). If this principle is
operable then it can be assumed that one or both Holcocephala species
have adjusted, e.g. partitioned resources, in such a way as to reduce com-
petition by more effectively exploiting the available resources. The purpose
of this study was to examine coexistence of H. abdominalis and H. calva
in a Maryland habitat in terms of resource partitioning involving foraging
behavior, prey selection, and predation between the two species.

HABITAT AND METHODS

This study was conducted in a mesic hardwood picnic area, located near
York Road in the northeast section of the Towson State University campus,
during July, August, and September of 1973 through 1977. The linear 2.2 ha
plot has steep margins, which drop 8 to 10 m in elevation from the sur-
rounding campus, and is traversed by a stream and several footpaths.
Woody vegetation in the study area consisted of a mixed stand of bottom
land trees, with Liriodendron tulipifera L., Platanus occidentalis L.,
Quercus spp., Prunus serotina Ehrh., and Gleditsia triacanthos L. along
the sloped margins; and with Juglans nigra L. and scattered Ulmus amer-
icana L., Fagus grandifolia Ehrh., Acer spp., and Pinus spp. predominating
on the flat. This overstory produced a continuous shaded condition during
the day, except for brief periods in the middle of the day and late afternoon.
A dense shrub growth of Vitis sp., Rubus sp., Lonicera sp., Lindera benzoin
Blume, and herbaceous plants occurred along the steep slopes, stream and
less disturbed areas. Temperatures and relative humidities ranged from 14°C
and 99% in early morning to 36°C and 28% in mid-afternoon.

Prey selection and predation by the two species was determined via sur-
veying flies along a standard route of ca. 100 m and following individual
flies for 1-2 h periods. These techniques were necessary in order to identify
the timing of major behavior patterns and for subsequent analysis of indi-
vidual patterns. Prey samples were taken hourly per day during the survey
and later measured for total body length (Scarbrough and Sraver, 1979).
Further information on types and selection of prey was taken while individ-
uals were under surveillance, although prey were not taken. Surveillance of
flies was made under bright skies from 0600 to 2000 h (EST) during 1974
and 1975. Flies were selected randomly for surveillance (see Scarbrough
and Norden, 1977), marked on their thoracic notum with fast drying enamel
paint and released. They were then followed to a perch where a 15 min
interval elapsed before data was taken. As indicated above, most flies were
observed for two consecutive hours during which time their complete be-
havioral repertoire was recorded. Males, however, were sometimes difficult
to follow when they were actively searching for mates. Consequently males
were followed for only one hour. At the end of each observation period,

VOLUME 84, NUMBER 2 351

additional flies were selected and followed for the subsequent period. Data,
resulting from the observation of adults of each species, were then compiled
to form a theoretical 14 h ten-day period. A combined total of 140 females
and 198 males were monitored. Temperatures and relative humidities were
taken approximately 1.5 m above ground level and adjacent to each perched
fly each observation hour.

RESULTS AND DISCUSSION

Foraging behavior.—Data generated by this study showed that the general
foraging behavior of the two Holcocephala species was similar, and essen-
tially agreed with that reported in earlier studies (Dennis, 1979; Johnson,
1976). Both species usually foraged from shaded vegetation along tree line
margins of the study site, utilizing perches between 0.2 and 5.0 m above
ground level. However, H. abdominalis differed in that it was invariably
more abundant and foraged from perches on low vegetation (<1.0 m),
whereas H. calva was more abundant and foraged from perches on taller
vegetation (>1.5 m). Holcocephala abdominalis also differed from H. calva
in that as its population density increased in late August, it dispersed into
the open, dry sunlit clearings and foraged from flower spikes and leaf tips
of Plantago lanceolata L. and other herbaceous vegetation (0.2 to 0.4 m).
These results agree with other reports (Scarbrough, 1974; Johnson, 1976)
wherein H. abdominalis has been found resting on and foraging from weedy
vegetation in dry fields which border streams and wooded areas. Dispersion
to adjacent areas is apparently initiated by increased interactions between
increased densities of conspecifics and congenerics for limited perches (un-
published data). Holcocephala calva, like H. fusca (Dennis, 1979), was
rarely found utilizing perches away from the edges of these clearings, but
usually rested on and foraged from perches in the aforementioned locations.

Asilids, except those of the Leptogasterinae, typically forage under
brightly lit conditions. Holcocephala fusca in Virginia (Dennis, 1979) and
H. abdominalis in Indiana (Johnson, 1976) behave similarly in that they
usually foraged and captured prey in open sunlit areas. In an earlier study,
I found several specimens (>300) of H. abdominalis foraging from sunlit
weedy vegetation throughout an abandoned pasture (20,235 m*) at Allerton
Park, Ill. (unpublished data). However, at the Towson study site and others
in Maryland, neither species consistently foraged nor captured prey in direct
sunlight. Both species usually foraged from shaded sites and captured prey
which were ‘‘back lighted’’ against a bright sky, often without venturing
into direct sunlight. The exception to the latter occurred in late afternoons
when forage sites were sunlit for short periods and temperatures were high
(Fig. 1). These sites were also sunlit for short periods in early morning, but
temperatures and foraging activities were correspondingly low. Holcoceph-

352 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ip)

— es
=]

O Shin teenate

=]

LL

; .

< A
:
O

LL

a

H.abdominalis
4 393 N=474

4 366 N=337

06-0700 10-1100 14-1500 18-1900

Ts le i

Fig. 1. Comparisons of intra- and interspecific foraging behavior per two hour period during
a 14 h day for two species of Holcocephala in Maryland.

VOLUME 84, NUMBER 2 353

ala abdominalis differed from H. calva in that its foraging sites were sunlit
more often and for longer periods and, hence, it foraged more frequently
under sunlit conditions.

Length of forage flights of the two species varied (R = 3-150 cm), al-
though most prey (90.5%) were captured between 15 to 100 cm from the
forage site. Holcocephala abdominalis captured most prey (88.2%) between
20 and 60 cm of the forage site, whereas H. calva captured most prey
(92.1%) between 40 and 100 cm. As reported in other studies of Hol-
cocephala (Dennis, 1979; Johnson, 1976), prey were captured in air either
in front of the forage site or to one side of it, with the predator usually
returning to the same forage site. Both species foraged in an oblique path,
with the greatest rise occurring near the interception point. Thus, prey were
invariably captured above the forage site. Upon capturing the prey, the
predator’s flight path looped above, and sometimes behind the previous
perch where it came to rest. If the interception point was far to one side of
the forage site, the fly did not always compensate in the return flight, missing
the previous perch but landing at a nearby site. The latter usually occurred
when prey were large (>3 mm) or the distance to the prey was unusually
long (>100 cm).

Figure 2 shows foraging patterns of Holcocephala expressed as propor-
tions of total flights. Foraging activities between species (y? = 75.1,
P < .001) were significantly different, with each species showing charac-
teristic foraging patterns. Both species foraged during each observation pe-
riod, with the lowest proportion of flights occurring during the first and last
periods of the day. The foraging pattern of H. calva peaked during the early
afternoon, but had decreased significantly by the last observation period. In
contrast, the peak foraging period of H. abdominalis was short and occurred
in late afternoon, following a consistent but lower level of foraging. The
latter species also exhibited a significant decrease in foraging by late after-
noon, but the decrease was less than that for H. calva. Furthermore, a
greater proportion of the foraging flights were made earlier in the day (Z' =
3.08, P < .05) by H. abdominalis than H. calva, producing a slightly greater
rapid rise in foraging activities.

Foraging patterns of conspecifics (x? = 17.6, P < .01, H. a.; x”? = 35.6,
P— 001, H. c:) and. congenerics (x? =°33.1, P.< .001, 65 y* = 42.1,
P < .001, 2) also differed significantly (Fig. 1). Males of both species
showed abrupt, rapid increases in foraging flights which exceeded that for
conspecific females. However, the peaks produced by male activity differed
in time and duration from that of H. calva, occurring earlier in the day, but
extending for longer periods in late afternoon. Female H. calva foraged

1 Differences between proportions (Z) from Zar (1974).

354 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

IME

RE -E? DAN Ge FT

%

06-0700 10-1100

calva

abdominalis

H. calva

a 2 N=2516m

4 66 N=1672min

H. abdominalis
4 $2 N= 2294min

4 66 N=1524min

14-1500

1, hale ee SE

Fig. 2. Comparisons of intra- and interspecific feeding behavior per two hour period during

a 14 h day for two species of Holcocephala in Maryland.

18-1900

VOLUME 84, NUMBER 2 355

Table 1. Foraging responses of two species of Holcocephala toward potential prey. Data
for conspecific males and females within a column are not significantly different. Data between
congeneric sexes or totals for species within a column are significantly different at P < .01
level.

Captured Non-Captured Investigated
Species No. % No. % No. %
———
H. abdominalis 350 43.9 26 3)53) 421 52.8
Males 142 42.1 13 3.9 182 54.0
Females 208 45.2 13 2.8 239 52.0
H. calva 452 37/7 25 2-1 721 60.2
Males 179 37.9 5 1.0 294 61.5
Females 273 Sip 20 2.8 427 59.3

more frequently in mid- to late afternoon following a steady increase during
the earlier part of the day. In contrast, female H. abdominalis showed an
increase in foraging until 1000 h, after which no significant change occurred.

When potential prey flew near specimens of either perched species, their
movements usually stimulated them to chase. Upon approaching the poten-
tial prey in flight, the asilids either rejected the prey item without physical
contact or attempted to capture it. Table | shows the responses of Holco-
cephala to potential prey. Foraging responses between conspecific males
and females were similar (? = 1.21, P < .05, H. a.; x? = 3.14, P < .0S,
H. c.), whereas significant differences (y? = 11.63, P < .01) existed be-
tween species. Holcocephala abdominalis successfully captured (x? = 7.93,
P < .01) agreater proportion of prey, but investigated (y? = 10.59, P < .01)
fewer potential prey than did H. calva. Both species immobilized most prey
that they captured. Some large prey were released or dropped without being
immobilized during the return flight to the forage perch. These data suggest
that perched H. abdominalis may be more capable of recognizing capturable
potential prey than H. calva.

Feeding behavior. —Prey were usually impaled upon the hypopharynx of
the predator at the time of capture, although large prey (>3.4 mm, e.g.,
Reticulitermes flavipes (Kollar) and Ponera pennsylvanica Buckley) were
manipulated with all six tarsi near the forage site before insertion. The
hypopharynx was usually inserted in the dorsum of the prey, normally in
the thorax. The eyes and tip of the abdomen were sometimes used when
prey were small (<2 mm) or following manipulation and after the asilid had
been feeding for extended periods. Apparently the choice of the site for
hypopharynx insertion was related to the manner in which the predator
captured its prey. Both asilids usually captured prey by grasping the margin
of the thorax, resulting in the largest part of the prey’s body being directly
below the predator’s head.

356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Additional activities exhibited by both species while feeding included
grooming, head movements, and manipulating prey. Perched asilids may
groom any part of their body at any time. However, while feeding, grooming
was limited to rubbing the hindlegs together or over the surface of wings
and abdomen. Grooming of the more anterior parts of the body usually
followed feeding. Rapid head and body movements directed toward moving
objects suggested that the asilids detected both crawling and flying insects
and other arthropods in the vicinity of their perches. Prey were manipulated
while the asilids perched or hovered near the feeding site. During manipu-
lation, the position of the hypopharynx in the prey was adjusted, usually
involving its removal and insertion at a different site. The flies removed
prey from the hypopharynx by pushing it off with the foretarsi while perch-
ing, whereas all six legs were used during its removal and manipulation
while hovering. Hovers (x = 10s, R = 6-20, N = 66, H.c.;x =78,R =
4-17, N = 48, H. a.) varied in length, with longer times being associated
with larger prey. The asilids invariably returned to previous perches to
resume feeding following manipulation. Holcocephala calva, in contrast to
H. abdominalis, sometimes manipulated prey without removing it from the
hypopharynx by using one tarsus to ‘‘spin’’ it on the hypopharynx, or ad-
justing the depth of the hypopharynx by pushing the prey against the perch.
Holcocephala fusca (Dennis, 1979) also has been reported to hover and
manipulate prey.

While perching or feeding, the asilids were sensitive to moving shadows
that accompanied abrupt changes in air currents. Depending upon the
strength of the stimulus, they responded by flying to another location or by
moving their hindlegs in an alternating ‘‘kicking’’ motion. If the stimulus
occurred often, they retreated (ca. 2-3 cm) from the tip of a twig and re-
peated the ‘‘kicking.’’ If the stimulus was strong, such as that produced by
an approaching predator (e.g., Vespula spp., conspecifics and Araneida),
they invariably flew to a nearby perch. Flight was more frequent when the
stimulus was generated in front of the perched asilids. Leg motion and
retreating occurred more frequently than flight, especially when the stimu-
lation originated behind the asilids. Speed of leg movement was highly vari-
able, but was apparently related to air velocity. The asilids were induced to
kick at different frequencies either by blowing air upon them via a hollow
tube at different velocities or by waving a hand near them at different
speeds. Leg movement usually increased with either increased air current
velocity or increased frequency of waving.

Dennis (1979) and Johnson (1976) reported that Holcocephala removed
prey by either pushing it off the hypopharynx with a foretarsus or with-
drawing the hypopharynx while perched or flying, allowing the prey to drop.
Both species in this study also used these methods to remove prey at the
completion of feeding. Furthermore, when the latter method was used at a

VOLUME 84, NUMBER 2 357

Table 2. Mean feeding times per prey and mean number of prey fed upon per unit time for
two species of Holcocephala during 10 observation days in Maryland.

sss

; Time/prey

Species Sample (min) Prey/h/d Prey/d

H. abdominalis 430 8.58 Les 21.5
Male 172 R25 2A ee
Female 258 8.9] 1.8 258
H. calva 423 9.15 1.8 25.3
Male 148 8.80 se 18.48
Female 275 9.44 2:3 LyPH Ps

* Means followed by the same letter within a column are not significantly different. Remain-
ing data between congeneric sexes or species within a column are significantly different at
P < .05 level (¢t test).

perch, the asilid’s head was tilted forward approximately 90°, positioning
the hypopharynx in an almost 180° vertical plain to the ground. This behav-
ior apparently enhances the role of gravity in removing the prey.

Table 2 depicts the mean feeding time per prey and mean number of prey
fed upon per unit time for ten observational days. These data show that H.
calva fed for significantly longer periods per prey and fed upon larger num-
bers of prey per h and per day than H. abdominalis. Similar differences
were found between sexes of each species, with females feeding for longer
periods and upon larger numbers of prey than males. In contrast, males
differed from females in that they fed upon approximately equal numbers
of prey per unit time. Differences in mean feeding times and mean number
of prey are attributed to greater predator-prey size relationships and greater
energy demands between the sexes and the species.

Time utilized for feeding per 2 h period per day for both species of Hol-
cocephala varied significantly (x? = 152.4, P < .001), although each exhib-
ited a distinct feeding pattern (Fig. 2). Both species fed for shorter periods
during early morning, but showed increased feeding times by mid-morning.
Holcocephala calva fed for consistently longer periods (1000 to 1700 h, no
significant difference per 2 h period) through most of the day, although a
significant decrease (Z = 5.28, P < .05) occurred in late afternoon. In con-
trast, feeding times were less (x? = 4.9, P < .05) from 1000 to 1500 h for
H. abdominalis than for H. calva, but were significantly greater (y? = 69.1,
P < .001) from 1600 to 1900 h. Feeding times also decreased (Z = 16.7,
P < .001) in late afternoon for H. abdominalis but the decrease was signif-
icantly less than that for H. calva.

Conspecific asilids displayed significant variations (x° = 134.1, P < .001
36 vs. 9 H.a.; x? = 105.4, P < .001 od vs. 2 H. c.) in feeding proportions
during certain periods of the day (Fig. 2). Conspecific males utilized less
feeding time than conspecific females from 0600 to 1000 h, but feeding times

nw

58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Correlations between foraging and feeding behavior with two environmental pa-
rameters for two species of Holcocephala.

Temperature RH
Species Foraging Feeding Foraging Feeding
H. abdominalis 0.9124 0.746» —0.821° —().848°
H. calva 0.786" 0.861° —0.827 —().856°
a P ean
DyPa<<n()5):

increased significantly by the 10-1100 period. Feeding times of male H.
calva increased until 1500 h when they began to decrease. In contrast,
feeding times of male H. abdominalis did not change significantly between
10-1500 h, but were followed by a rapid rise by 16-1700 h and a subsequent
rapid decrease by 1900 h. Feeding times of females of both species tended
to level off with only minor fluctuations following the rise in early morning.
Feeding times of female H. calva decreased rapidly by 1900 h whereas those
of H. abdominalis remained relatively consistent with former periods.
Foraging and feeding patterns for both species were positively correlated
with temperature and negatively correlated with relative humidity (Table 3).
As air temperatures increased and relative humidities decreased, both for-
aging rates and feeding times increased (Figs. 1, 2). Both species utilized
less time for feeding and foraged less during early morning when tempera-
tures were lowest and relative humidities greatest. Also, H. calva showed
a significant decrease in foraging rates and feeding times in afternoon as
relative humidities increased. Activities remained relatively constant in

Table 4. Dietary composition and characteristics of prey captured by two species of Hol-
cocephala in Maryland. Predator lengths (mm) H. calva, * = 7.25 + 0.58; H. abdominalis,
xX = 5.81 + 0.64.

H. abdominalis H. calva
Prey No. % x SD No. % x¥ SD
Diptera 235 35.8 1.58 + 0.652 184 Sil 1:97 == 037562
Hymenoptera 223 34.0 2.36 + 0.87» 170 28.7 23622 19"
Coleoptera 106 16.2 1.54 + 0.598 59 10.0 1.88 + 0.674
Homoptera, Hemiptera 45 6.8 1.47 + 0.498 89 15.0 1:80)-s00772
Psocoptera 27 4.1 M67, ==10:422 72 Pr? 1.82 + 0.50?
Misc. 20 3.0 1.74 + 0.40” 18 3.0 1.89 + 0.47»

656 99.9 7/8) 22 (U)S%7/ 592 100.0 1.95 OWS

2 Significant difference within prey taxa between species at P < .05 level.
” No significant difference within prey taxa between species (t test).

VOLUME 84, NUMBER 2 359

Table 5. Dietary composition and characteristics of prey captured for Holcocephala ab-
dominalis in Maryland. Predator lengths (mm) 5.63 + 0.59 6, N = 20; 5.9 + 0.69 2, N = 20.
Means within common taxa are not significantly different (r test).

Females Males
Prey No. % SD No. % 7 SD

Diptera 121 35.6 1.59 + 0.71 114 36.1 (else==0 570
Hymenoptera 118 34.7 2.44 + 0.83 105 3322 2.35 + 0.86
Coleoptera 51 15.0 1.47 + 0.56 55 17.4 S0'=7057
Homoptera, Hemiptera 27 WL 1.52 + 0.47 18 S35 0/ 1.43 + 0.50
Psocoptera 13 3.8 1.67, 0:53 14 4.4 1.587=0535
Misc. 10 229 1.74 + 0.40 10 Bae 1.74 + 0.40

340 99.9 1.74 + 0.53 316 100.0 1-71 20:56

afternoon with only male H. abdominalis showing a decrease in these two
patterns.

Prey selection.—Both species of Holcocephala selected similar prey taxa,
with Diptera and Hymenoptera being taken most frequently (Table 4, see
list of prey). Others (Dennis, 1979; Johnson, 1976; Bromley, 1950b; McAtee
and Banks, 1920) have also reported that Holcocephala feed primarily on
prey within these insect orders. Holcocephala abdominalis captured pro-
portionally more hymenopterous and coleopterous prey and fewer homo-
hemipterous and psocopterous prey than did H. calva. Both sexes of H.
abdominalis captured prey in each taxa in almost equal proportions (Table
5). In contrast, female H. calva captured significantly larger proportions of
hymenopterous, coleopterous, and homo-hemipterous prey and fewer dip-
terous and psocopterous prey than did conspecific males (Table 6).

Table 6. Dietary composition and characteristics of prey captured for Holcocephala calva
in Maryland. Predator lengths (mm) 7.0 + 0.52 6, N = 20; 7.48 + 0.52 2, N = 20.

a

Females Males
Prey No. % x SD No. % < SD

ot SORT RAE Be ee eS ee ee
Diptera 96 26.2 2.08 + 0.083 88 39.1 1. 857210575"
Hymenoptera 119 32.4 252 ee e214 51 22 2.00 + 1.023
Coleoptera 38 10.4 1.86 + 0.62° 21 9.3 1.90 + 0.74
Homoptera, Hemiptera 64 17.4 185. 05708 25 11.1 1.82 + 0.87°
Psocoptera 39 10.6 182) 0542 33 14.7 1.81 + 0.45
Misc. 1] 3.0 1.89 + 0.54 7 Seal 1.86 + 0.31°

367 99.9 Dully==8 0292 225 100.0 1.89 + 0.80

eee ere ee ee eee rere eee Eee

4 Significant difference within prey taxa between sexes at P < .05 level (¢ test).
> No significant difference within prey taxa between sexes at P < .0S level (f test).

360 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Larger asilids usually selected larger prey and vice versa (Table 4). Mean
sizes for all prey taxa for H. abdominalis and H. calva were 1.73 and 1.95
mm, yielding predator-prey length ratios of 3.4 and 3.7, respectively. Most
prey were less than 3 mm in length, with 96.3% and 87.2% for each species,
respectively. Thus H. abdominalis captured significantly more (Z = 4.53,
P < .05) small prey than did H. calva. Within the major prey taxa, only
hymenopterous prey were not significantly different in size for the two
species. The latter is the result of the proportionally large number of winged
reproductive ants (70.4% H. a.; 34.1% H. c.) in each of the samples.

Females of Holcocephala were significantly larger than their conspecific
males, and both sexes of H. calva were larger than those of H. abdominalis
(Tables 5, 6). Sexes of H. calva usually selected larger prey within common
taxa than did H. abdominalis. Furthermore, female H. calva selected larger
dipteran and hymenopteran prey than did its conspecific male. Male H.
calva captured primarily Apocrita Hymenoptera (98.9%), whereas females
captured more formicids (48.1%). Removal of formicids from the females
list reduced the mean hymenopteran size to 2.18 mm, which does not differ
significantly from that of the male. Similar larger hymenopterous prey means
for both sexes of H. abdominalis resulted from the large number of repro-
ductive ants captured, and their removal produced similar results (x = 1.98
mm). Comparisons of prey means within common taxa for the sexes of H.
abdominalis show no significant difference in mean size.

The following are lists of prey taken by H. abdominalis and H. calva at
the study site. In some instances prey are determined only to order or family
level, since some prey were not taken or they were too damaged for iden-
tification. Each notation of prey refers to a single record unless followed by
a number in parentheses. The month and year are recorded only once at the
end of a series for each prey taken.

Prey of H. abdominalis.—ARANEIDA (all immatures): Agelenidae, un-
identified 20 (2) 27. VIII.74; Clubionidae, Liocraninae 23.VIII.74; Thomisi-
dae, Misumeninae 23. VIII.74; Salticidae unidentified (2) 28. VIII.74; Pisaur-
idae, Dolomedes sp. 8.VIII.73. PSOCOPTERA: Unidentified 9.VIII.72;
Ectopsocidae, Ectopsocopsis cryptomeriae (Enderlein) 22 (3) 27, 29. VIII.74,
(9) 16.VIII.77, (6) 1, (2) 3.1X.77; Lepidopsocidae, Echmepteryx hageni
(Packard) 22.VIII.74, 30.VIII.77; Psoquillidae, Rhyopsocus sp. 15.VUI.74.
HOMOPTERA-HEMIPTERA: Aleyrodidae, unidentified 3.1X.77; Antho-
coridae, Orius insidiosus (Say) 9.VIII.72, 30. VIII, (3) 1.1X.77; Aphididae,
unidentified (11) 28.VIII.74, Anoecia corni (F.) 16.VIII.74, A. cornicola
Walsh 22.VIII.74, 16, (2) 30.VHI.77, Aphis forbesi Weed 22.VII1.74, Chai-
tophorus pusillus Hottes and Frison 25.VIII.74, Macrosiphum sp. 9.VIII.72,
M. liriodendri (Monell) 22.VIII.74, (3) 16. VIII, (2) 1.1X.77, Myzocallis sp.
27.VIII.74, Thecabius sp. 22, 29.VIII.74; Cicadellidae, Macrosteles fasci-
formis (Stal) 11.VIII.72, nymphal Cicadellinae 27.VIII.74; Coccoidea, un-

VOLUME 84, NUMBER 2 361

identified 16. VIII, 3.1X.77; Delphacidae, Delphacodes sp. 28.VIII.74; Mi-
ridae, unidentified 25.VIII.74, (2) 28. VIII.75, Halticus intermedius Uhler
(2) 3, 9. VII.72. THYSANOPTERA: Phlaeothripidae, unidentified 9. VIII.72,
22, 23. VIII.74, 1.1X.77: COLEOPTERA: Chrysomelidae, Chaetocnema sp.
9.VIII.72, 23.VIII.74; Hydrophilidae, Cryptopleurum minutum Fabricius
7.VIII.72; Mycetophagidae, Litargus tetraspilotus LeConte 25.VIII.74, (12)
16. VIII, 3.1X.77; Orthoperidae, Orthoperus sp. (3) 16.1X.74; Phalacridae,
Olibrus sp. 28.VIII.74; Ptilidae, Actinopteryx sp. (13) 27, (12) 23, (6) 22, 25,
(5) 27, (3) 28, (2) 29. VIII.74, 16.1X.74; Rhizophagidae, Monotoma longi-
collis Gyllenhal 27. VIII.74; Scaphidiidae, Eubaeocerus sp. 23.VIII.74; Sco-
lytidae, Hypothenemus sp. 29.VIII.74, 30.VUI, 1.1X.77, Pityogenes hop-
kinsi Swaine 20.VIII.74, Pityophthorus pulicarius (Zimmermann) 27.VIII.74,
P. sp. near puberulus LeConte 27.VIII.74, Xyleborus rubricollis Eichhoff
27.VIII.74; Staphylinidae 22, (2) 23, (13) 27, (2) 28, 29, (3) 14, (6) 16, (4)
30. VIII.77. LEPIDOPTERA: Elachistidae, Elachista sp. 27.VIII.74. DIP-
TERA: Anthomyzidae, Mumetopia occipitalis Melander 9.VIII.72,
25.VIII.74, (2) 30. VIII.77, 1.1X.77; Cecidomyiidae, Asteromyia sp. 27, (13)
28.VIII.74, Atrichopogon sp. (2) 22, 25, 28, 29. VIII.74, 16, (2) 30. VIII.77,
(13) 1.1X.77, Cecidomyiidii spp. (22) 22, 23.VIII.74, Contarinia sp.
28. VIII.74 (2) 16. VIII, (3) 3.1X.77, Dasineura sp. (2) 28.VIII.74, Lestodi-
plosis sp. (2) 27.VII1.74, Lestremia sp. (2) 28.VIII.74, Micromya sp. 27,
29.VIII.74, Porricondyla sp. 27.V111.74, Procystiphora sp. 28.VIII.74, Tri-
sopsis sp. 27.VIII.74; Ceratopogonidae, Dasyhelea sp. 29.VIII.74, Forci-
pomyia sp. (7) 28.VIII.74; Chironomidae, Orthocladinae 29.VIII.74, Cri-
cotopus sp. (2) 9.VIII.72, unidentified 22, 27, (16) 28.VIII.74, (13) 16, (4)
30.VIII.77, (3) 1.1X.77; Chloropidae, Oscinella carbonaria (Loew)
29. VIII.74, O. umbrosa (Loew) 9.VIII.72, 25.VII.74, Thaumatomyia bi-
striata (Walker) 28.VIII.74; Dolichopodidae, Chrysotus sp. 9.VIII.72,
1.1X.77; Drosophilidae, Drosophila melanogaster Meigen (2) 9.VIII.72, (2)
30. VIII.77; Empididae, Tachypeza sp. 27.VUI.74; Ephydridae, Discocerina
sp. 9.VIII.72, Hydrellia formosa Loew (2) 22.VII.74, Leptopsilopa nigri-
mana (Williston) 28.VIII.74; Muscidae, Coenosia sp. 9.VIII.72; Phoridae,
Megaselia sp. 20, (12) 28.VIII.74, (3) 16, 17, (2) 30.VIII.77, (3) 1.1X.77,
Puliciphora sp. 20, (2) 28.VIII.74; Pipunculidae, Chalarus spurius (Fallen)
27. VIII.74; Psychodidae, Psychoda sp. 23.VIII.74; Scatopsidae, Scatopse
fuscipes Meigen 28.VIII.74, 30.VIII.77; Sciaridae, Bradysia spp. (17) 22,
B)e23.46)ii 255 (G6) 27,5. Gy’ 28, 16S) 29. VILL.74,. Seiara sp.1'15; VMI. 74, (2)
30. VIII.77, (2) 1.[X.77; Sphaeroceridae, Leptocera sp. 28, (2) 29. VIII.74,
L. palliceps Johnson 25, 28.VIII.74, 16.VIII.77. 3.1X.77. HYMENOP-
TERA: Unidentified (2) 27, (2) 28.VIII.74; Aphelinidae, unidentified
22. VIII.74; Aphidiidae, Praon sp. 23.VIII.74; Braconidae, Aphaereta pal-
lipes (Say) 27.VIII.74, Aspilota sp. 20, 23, 28.VIII.74, Chorebus sp.
28.VIII.74, 12, 16.VIII.77; Ceraphronidae, Ceraphron sp. 20, 23,

362 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

29.VIII.74, Dendrocerus sp. 20, 23, 29.VIII.74; Cynipidae, Hexacola sp.
28.VIII.74, 16.VIII.77; Diapriidae, Trichopria sp. 28.VIII.74; Encyrtidae,
unidentified 28. VIII.74, (4) 1.[X.77; Eulophidae, unidentified 22, 28. VIII.74,
12, (3) 16. VIII.77; Formicidae, Ponera pennsylvanica Buckley 28.VIII.72,
22013) 230(22) 27; 142): 283°(17) 29. VEN 4G) 8 12 EX 4e CL) torent
30. VIII.77, (16) 1, (2) 3.1X.77, Myrmecina americana Emery (13) 22, 27,
(3) 28.VIII.74, (6) 12, (2) 30.VIII.77; Mymaridae, unidentified 22,
28.VIII.74; Pteromalidae, 7.VIII.72, 20, 22, 28, 29. VIII.74, 30. VIII.77, 1,
3.1X.77; Scelionidae, Gryon sp. 27.VIII.74, Telenomus sp. 25, (3)
28. VIII.74.

Prey of H. calva.—ARANEIDA: Unidentified immatures 1.VIII.75, 9,
10, 12.VII.76, 1, 20, (3) 23.VII.77, 16.VIII.77; Linyphiidae, Meioneta
unimaculata (Banks) 17.VII.74. PPOCOPTERA: Unidentified (2) 19. VIII.72,
12.VII.76; Caecilidae, Caecilius aurantiacua (Hagen) 30.VII.74, (2)
17. VII.75, (2) 10. VII.76; Ectopsocidae, Ectopsocopsis cryptomeriae (Ender-
lein) 1, (2) 16, (2) 22, (3) 29. VII.74, 20. VII.75, (2) 9, 10, (8) 12, (4) 13. VII.76,
1, (2) 20, 232VIL.77, 5; Gye 11, 15,912). 16, (4) 1223, 4)225-Vile77; eepidope
socidae, Echmepteryx hageni (Packard) (3) 30. VII.74, 9. VII. 76, 23. VII.77,
(2) 16, 22.VHII.77; Philotarsidae, Aaroniella sp. 29, 30.VII.74, 12.VII.76.
HOMOPTERA-HEMIPTERA: Aleyrodidae, unidentified 25.VIII.77; An-
thocoridae, Orius insidiosus (Say) 22, (3) 25.VIII.77; Aphididae, Anoecia
graminis G. and P. 1, Aphis sp. 30.VIII.74, Chaitophorus pusillus H. and
F. 15.VIII.74, Drepanosiphinae (5) 9.VIII.72, Myzocallis tiliae (L.)
30. VIII.74, Myzus sp. 30. VIII.74, 25. VIII.77, Prociphilus fraxinifolii (Riley)
1. VIII.74, Rhopalosiphum rufiabdominalis (Sasaki) 30.VII.74, Tinocalis ul-
mifolii (Monell) (5) 30. VII.74, (2) 12, 15, (4) 30. VII.75, 1, (2) 5. VHI.75, (2)
1 (3)19326): 15; °C) 16, 6)" 20; 23. VIL 7, Se GS) LPS yO) 16, 42a
25.VIII.77, unidentified 9, 10, 12. VIII.76; Cicadellidae, Dikraneura sp. (2)
27.VIL.74, 9.VII.76, (2) 23; 11. VIII.77, Scaphytopius: sp; 12-Vil.76; Grae
idae, Myndus pictifrons Stal 16.VII.74; Delphacidae, Delphacodes sp.
16.VII.76, 25.VIII.77; Miridae, MHalticus intermedius Uhler 9.11.72,
30.VII.74, 13. VII.76, 8.VII.75, unidentified 20.VII.77, 16.VIII.77; Tingi-
dae, Corythuca ciliata (Say) 12.V1I.76. THYSANOPTERA: Unidentified 1,
(2) 12. VII.74, 5, 20. VII.77, 16, 22, 25. VHI.77. COLEOPTERA: Anobiidae,
Petalium sp. 19.VIII.74; Chrysomelidae, Chaetocnema sp. 3.VIII.72,
30.VII.75, 13. VII.76, (2) 25. VIII.77; Ciidae, unidentified 29. VII.74, 23, (2)
16. VIII.77; Hydrophilidae, Cercyon sp. 6.VUI.75; Lathridiidae, Aridius no-
difer (Westwool) 9.VIII.72, Corticaria sp. 30.VII.74, 9.VII.77, 11, (2) 16,
25.VIII.77; Mycetophagidae, Litargus tetraspilotus LeConte (2) 16, 22, (3)
25.VIII.77; Orthoperidae, unidentified 10. VII.76; Ptilidae, Actinopteryx sp.
10. VII.76; Staphylinidae, unidentified 29, (2) 30.VII.74, 1.VII.74, 10, (4)
Lae 22523; 29. VI1.75, (5) 3.VIIL-75, 10; @yi2, 13s VIL 76.) 20 ili
23.VII.77, 5, (4) 16, (3) 22. VII.77. LEPIDOPTERA: Tineidae, Homosetia

VOLUME 84, NUMBER 2 363

sp. 25.VIII.74. DIPTERA: Agromyzidae, Phytomyza sp. 16.VII.74,
13.VII.76, 16, (2) 22.VIII.77, Cerodontha dorsalis (Loew) 16.VII.74: Chi-
ronomidae, unidentified 15.VII.75, (12) 10. VII.76, (2) 5, 23. VII.77, 5, (4) 16,
(3) 22, 25.VIII.77; Cecidomyiidae, Lestremia sp. 30.VII.75, 12.VII.76, 5,
13, (2) 20, 23. VII.77, 16, 22. VIII.77, Lasioptera sp. 9.VII.77, (2) 25. VIII.77;
Ceratopogonidae, Atrichopogon sp. 12.VII.76, 5, 16, 22.VIII.77, Forcipo-
myia sp. 12.VII.76, (6) 16.VHI.77; Chloropidae, Elachiptera erythropleura
Sabrosky (2) 22.VII.74, Hippelates bishoppi Sabrosky 22.V1I.74, 1. VIII.74,
15. VIII.75, Oscinella frit (L.) 29. V11.74, (2) 22. VIII.77, O. soror (Macquart)
29.VII.74, 10. VII.76; Empididae, unidentified 22.VIII.77; Dolichopodidae,
unidentified 13, 29. VII.75, 3. VIII.76, Chrysotus sp. 20, 23, 16. VIII.77; My-
cetophilidae, unidentified 25.VIII.75; Phoridae, Dohrniphora sp. 30, (2) 9,
(3) 12, 13. VII.76, 11, 15, (4) 16, 22.VIII.77, Megaselia sp. 30.VII.74, (3)
PVT S (4) a1, 1S VEG, “QUIS 15986, 9(4)020;4) 232V 177, Sone,
22.VIII.77, Puliciphora sp. (2) 30.VII.74, 9, 12.VII.76; Pipunculidae, To-
mosvaryella sp. 25.V1I1.77; Psychodidae, Psychoda sp. 25. VIII.77; Scatop-
sidae, Scatopse fuscipes Meigen 31.VII.75, 10, (2) 13.VII.76, 25. VIII.77;
Sciaridae, Bradysia sp. 17.VII.74, (2) 29, (3) 30.VII.74, (3) 1. VIII.74, (2)
POO VILEIS | 2) 9310;, (213. VInTG;-1; (2) 572) 99 @)3¢ 1591694) 202Vi1 LT r
5, (3) 11, G) 15, () 16, (2) 23.VIII.77; Sphaeroceridae, Leptocera sp.
SVAN 2. °O) TL 2 VAL F415 2163 205 (3): 230Vin77; G) 25NVIN77; Sirate
mylidae, Microchrysa polita (L.) 19.VII.72. HYMENOPTERA: Unidenti-
fied (3) 17; (3) 22, (3) 29. VII.74, 1. VIII.74, (8) 26, (12) 29. VII.75, (5) 3, ©)
15.VIII.75; Aphidiidae, Praon sp. 30.VII.74; Braconidae, Aphaereta sp. 12,
Bev. 4..(5) 50. Villy 5,12 Vil-76, 15,(2) 20. VIl.77,.G) 5; 11 VIEL77, Cree
ebus sp. 30.VII.74, 9. VIII.74, 2.VII.75, Euphoriella sp. 30.V11.74; Cera-
phronidae, Ceraphron sp. 22.V11.74, Dendrocerus sp. 30.V11.74; Cynipidae,
unidentified (3) 15. VIII.75, 12, 13.VII.76; Encyrtidae, unidentified 15, (2)
20, 23.VII.77, 5.VIII.77, (2) 16, 22, 23. VIII.77; Eupelmidae, unidentified
(2) 20. VII.77; Eulophidae, unidentified 5, 16, 20, (5) 23. VII.77, 5. VIII.77,
ii; 15;,.-G) 16, 228° 25:N 1.77; Formicidac,. Lasits’sps (426. Vi. 72,13)
SU Vil: 7/4513. Vili7 7, Myrmecina sp: 5; 20.V1L77, G) 11, 1576) 10 VuE77,
Ponera pennsylvanica Buckley (3) 30. VII.74, (10) 12, (7) 14, (8) 15. VII.75,
9.VII.76, 13, 20, (3) 11, (4) 16, (3) 22.VIII.77; Perilampidae, unidentified
20.VII.77, 16, (7) 25.VIII.77; Pteromalidae, prob. Habrocytus sp. 9, (5)
12.VII.76, unidentified 15.VIII.77; Scelionidae, Telenomus sp. 30.VII.74,
unidentified (2) 5. VIII.75.

ACKNOWLEDGMENTS

J. L. Herring (Anthocoridae, Miridae), L. M. Russell and M. B. Stoetzel
(Aphididae), J. P. Kramer (Cicadellidae, Delphacidae, Cixiidae), K. O'Neill
(Thysanoptera), R. E. White (Anobiidae, Chrysomelidae), J. M. Kingsolver
(Ciidae, Ptilidae, Phalacridae, Mycetophagidae, Scaphidiidae, Lathridiidae,

364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Rhizophagidae), R. D. Gordon (Orthoperidae), D. M. Anderson (Scolyti-
dae), R. W. Hodges (Elachistidae), G. Steyskel (Agromyzidae, Sphaero-
ceridae, Muscidae), C. W. Sabrosky (Chloropidae, Anthomyzidae), W. W.
Wirth (Drosophilidae, Stratiomyiidae, Pipunculidae, Chironomidae, Cera-
topogonidae, Phoridae, Ephydridae), R. J. Gagné (Sciaridae, Cecidomyi-
idae, Scatopsidae), F. C. Thompson (Psychodidae, Dolichopodidae), L.
Knutson (Empididae), G. Gordh (Pteromalidae, Mymaridae, Eulophidae,
Aphelinidae, Encyrtidae), P. M. Marsh (Diapriidae, Scelionidae, Cera-
phronidae, Aphidiidae, Braconidae), A. S. Menke (Cynipidae), D. R. Smith
(Formicidae) of the Systematic Entomology Laboratory, IIBIII, Agric. Res.
Service, USDA, Washington, D.C. and Beltsville, Maryland; D. R. Davis
(Tineidae), P. J. Spangler (Hydrophilidae), R. C. Froeschner (Tingidae) of
the National Museum of Natural History, Smithsonian Institution, Wash-
ington, D.C.; E. L. Mockford (Psocoptera) of the Illinois State University,
Normal, and W. P. Peck (Araneida) of Central Missouri State University,
Warrensburg, are thanked for their prompt identification of their respective
groups. Erik Scully, Department of Biological Sciences, Towson State Uni-
versity, provided valuable assistance with the statistical analysis. D. S. Den-
nis, Environmental Sciences Division, Stearns-Roger Incorporated, P.O.
Box 5888, Denver, Colorado, and Robert J. Lavigne, Department of Ento-
mology, University of Wyoming, Laramie, reviewed and made valuable
comments on the manuscript. This study was supported by a grant received
from the Faculty Research Committee at Towson State University.

LITERATURE CITED

Baker, N. T. and R. L. Fischer. 1975. A taxonomic and ecological study of the Asilidae of
Michigan. Great Lakes Entomol. 8: 31-91.

Brimley, C. S. 1922. List of the robber flies (Asilidae, Diptera) of North Carolina. Entomol.
News 33: 294-298.

Bromley, S. W. 1931. A preliminary annotated list of the robber flies of Ohio (Diptera:
Asilidae). Ohio State Univ. Mus. Sci. Bull. 1: 3-19.

—. 1946. Honeybee predators. J. N.Y. Entomol. Soc. 36: 195-199.

. 1950a. Florida Asilidae (Diptera) with description of one new species. Ann. Entomol.

Soc. Am. 43: 227-239.

. 1950b. Ohio robber flies. V. Ohio J. Sci. 50: 229-234.

DeBach, P. 1966. The competitive displacement and coexistence principles. Annu. Rev. Ento-
mol. 11: 183-213.

Dennis, D. S. 1979. Ethology of Holcocephala fusca in Virginia (Diptera: Asilidae). Proc.
Entomol. Soc. Wash. 81: 366-378.

Hull, F. M. 1962. Robber flies of the world. The genera of the family Asilidae. U.S. Natl.
Mus. Bull. 224 (pt. 1), 430 pp.

Johnson, P. E. 1976. A preliminary investigation of the behavior of adult Holcocephala ab-
dominalis (Say) (Diptera: Asilidae) in Putnam County, Indiana, with a description of the
larvae and eggs. Master’s Thesis. DePauw University, Greencastle, Indiana.

Martin, C. H. and J. Wilcox. 1965. Family Asilidae, pp. 360-401. Jn Stone, A. et al., eds., A

VOLUME 84, NUMBER 2 365

Catalog of the Diptera of America North of Mexico. U.S. Dep. Agric., Agric. Handb.
No. 276.

MacArthur, R. H. 1958. Population ecology of some warblers of northeastern coniferous
forest. Ecology 39: 599-619.

McAtee, W. W. and N. Banks. 1920. District of Columbia Diptera: Asilidae. Proc. Entomol.
Soc. Wash. 22: 19.

Scarbrough, A. G. 1974. A faunistic study of Asilidae (Diptera) at three locations in northern
Baltimore County, Maryland: Incidence, relative abundance and seasonal distribution.
Proc. Entomol. Soc. Wash. 76: 385-396.

Scarbrough, A. G. and A. Norden. 1977. Ethology of Cerotainia albipilosa Curran (Asilidae:
Diptera) in Maryland: Diurnal activity rhythm and seasonal distribution. Proc. Entomol.
Soc. Wash. 79: 538-554.

Scarbrough, A. G. and B. Sraver. 1979. Predatory behavior and prey of Atomosia puella
(Diptera: Asilidae). Proc. Entomol. Soc. Wash. 81: 630-639.

Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Inc. 619 pp.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 366-368

AN ADDITION TO THE GENUS ANISOPS OF AUSTRALIA
(HEMIPTERA: NOTONECTIDAE)

CHARLES V. REICHART

Biology Department, Providence College, Providence, Rhode Island
02918.

Abstract.—Anisops ayersi, a new species from Ayers Rock, Northern
Territory, Australia, is described and the male diagnostic features figured.

During a three month collecting tour of Australia for notonectids in the
fall of 1980, I had the opportunity to visit Ayers Rock. This giant sandstone
monolith (4 miles long x 144 miles wide x 1143 feet high) is located about
280 road miles southwest of Alice Springs, Northern Territory.

The annual rainfall at Ayers Rock averages only about six inches a year,
but, at the base of the Rock, two shallow pools are sustained by the ground-
water that results from the runoff of these occasional rains. The first of the
two pools, Initi, on the north side, has a circumference of about 25 feet; the
second, called Mutidjula, is on the south side and is about 10 feet larger.
Vegetation is sparse in each pool but a population of tadpoles and small
Anisops seem to thrive. The Anisops turned out to be an undescribed
species herein described as Anisops ayersi.

Anisops ayersi Reichart, NEw SPECIES

Size: Male, 5.9 mm long, greatest body width 1.6 mm; female, 5.7 mm
long, greatest body width 1.58 mm. Shape: Short, robust species. Color:
Eyes yellowish grey, mottled with red. Vertex greyish red. Pronotum hya-
line permitting color beneath to show through. Scutellum yellowish brown
except apex which is clear hyaline. Hemelytra hyaline. Abdomen yellowish
brown, ventrally dark reddish brown with keel and segmental margins of
connexivum brownish yellow. Legs yellowish brown.

Male structural characteristics —Viewed from above, head rounded with
vertex slightly indented. Greatest width of head slightly less than pronotal
humeral width, 8% x anterior width of vertex and slightly less than 3x
median head length. Synthlipsis 3/; as wide as anterior width of vertex.
Median head length equal to median pronotal length. Humeral width of
pronotum slightly more than 2 the longitudinal length of pronotum; lateral

VOLUME 84, NUMBER 2 367

a =
3
y {
ane
es

Figs. 1-3. Anisops ayersi. 1, Left foreleg. 2, Stridulatory comb of tibia. 3, Rostral prong.

margins of pronotum divergent and 2 median longitudinal length of prono-
tum; posterior margin of pronotum subtly sinuous in the conventional con-
vex-concave anisopine pattern; median shallow depression extending almost
full length of pronotal disk. Scutellum ‘4 wider than long. Facial tubercle
slightly swollen, truncate above labrum with a few scattered white hairs.
Frons depressed between eyes from tubercle to vertex. Labrum barely wid-
er than long, apex broadly rounded with 3 long, white hairs on midline.
Rostral prong (Fig. 3) bluntly acuminate and shorter than 3rd rostral seg-
ment. Stridulatory comb (Fig. 2) with 14-15 teeth which increase in length
from outer and inner margins to center. Chaetotaxy of left foreleg as shown
in Fig. 1.

Female structural characteristics —Viewed from above head rounded
with anterior margin fairly straight; vertex slightly indented; greatest head
width slightly less than pronotal humeral width; 6%4x anterior width of

368 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

vertex. Synthlipsis ’% anterior width of vertex. Median length of head %/6
median length of pronotum. Humeral width of pronotum 24%2x median
length; lateral margins divergent and % median longitudinal length of prono-
tum; posterior margin weakly convex and medianly indented. Scutellum
length ''/i3 of width. Frons above facial tubercle slightly swollen; truncate
above labrum. Labrum with base slightly wider than long; apex bluntly
rounded.

Types and type-locality.—d (holotype) and 2 @ (paratypes), Initi water-
hole, Ayers Rock, Northern Territory, Australia, 8-X-80. Mark E. Pion and
C. V. Reichart.

Deposition of types.—Holotype and one paratype desposited in the Aus-
tralian National Insect Collection at Canberra, A.C.T.; one paratype re-
tained in author’s collection.

Comparative notes.—Anisops ayersi is similar in appearance and mea-
surements to A. nabilla Lansbury (1969) but can be easily distinguished
from the latter by the arrangement of teeth in the stridulatory comb and the
structure of the third rostral segment. In A. ayersi the teeth increase in
length from inner and outer margins toward the center while those of A.
nabilla increase in length from inner to outer margin. The apex of the third
rostral segment is wider than the base of the fourth segment in A. nabilla,
as in A. deanei Brooks (1951), whereas in A. ayersi the two segments are
the same width at the junction.

I did not attempt to place this species in Brooks’ (1951) key. Since the
revision by Brooks, 27 species have been added to Anisops, and a new key
is badly needed.

LITERATURE CITED

Brooks, G. T. 1951. A revision of the genus Anisops (Notonectidae, Hemiptera). Univ. Kans.
Sci. Bull. 34: 304-519.
Lansbury, I. 1969. The genus Anisops in Australia. J. Nat. Hist. 3(3): 433-458.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 369-375

ECTOPARASITES (MALLOPHAGA, ACARINA) FROM
THE DOUBLE-CRESTED CORMORANT
(PHALACROCORAX AURITUS)

IN FLORIDA

WILLIAM THRELFALL

Department of Biology, Memorial University, St. John’s, Newfoundland,
Canada A1B 3X9.

Abstract.—Seventy-seven double-crested cormorants, Phalacrocorox au-
ritus (Lesson), taken between 1973 and 1977 in Florida, were examined for
ectoparasites. Six species of ectoparasites were recovered (3 mallophagans,
1 tick, 2 mites). Quantitative data are given, with significant differences
being noted in the distribution of the parasites on the hosts, and in the
proportion of various age classes of parasites recovered.

The literature on the Mallophaga of Pelecaniformes is quite extensive with
Emerson (1972a) having gathered together North American records. Fur-
ther, several species of ticks and mites have been described and/or re-
covered from these hosts, e.g. Dinalloptes chelionatus Atyeo and Peterson
from the double-crested cormorant (Phalacrocorax auritus (Lesson)) in
Florida. The majority of the works, however, tend to be qualitative in nature
and not quantitative, e.g. Marshall and Nelson (1967), Price (1970), Ryan
and Price (1969) and von Timmermann (1967). A study of ectoparasites of
the double-crested cormorant in Florida was initiated in 1977 to determine
the parasite species present, density of infestation, frequency of occurrence
of each parasite species within the host population, and their distribution
on the host. It was also hoped that differences in density of infestation
between males and females, and between nestlings and adults, would be
revealed.

MATERIALS AND METHODS

Seventy-seven double-crested cormorants from several locations in Flor-
ida were examined for ectoparasites. Forty-four birds (22 adult males,
mean wt 1468 + 391 g, range 960-2100 g; 22 adult females, 1236 + 329 g,
750-2080 g) were taken on the west coast of the peninsula (2 on Sanibel

370 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Island, 24 in the Tampa Bay area, 18 on the Withlacoochee River) in the
period October 1973 to February 1976. An additional 33 birds (17 nestlings,
mean wt 895 + 503 g, range 159-1730 g; 8 adult males, 1953 + 124 g,
1690-2060 g; 8 adult females, 1574 + 146 g, 1310-1720 g) were collected
near the Merritt Island National Wildlife Refuge, on the east coast of the
Florida Peninsula in June and July of 1977. No attempt was made to sys-
tematically examine the former sample, thus only a qualitative picture is
presented for the sample. In the latter case, however, detailed necropsies
were undertaken and quantitative data were obtained. The population sam-
pled in 1973-1976 was composed of two subspecies (P. a. auritus (Lesson)
and P. a. floridanus (Audubon)) while that of 1977 contained only P. a.
floridanus. The former is migratory, the latter a permanent resident.

The techniques for obtaining and examining birds in this study were the
same as those outlined in Eveleigh and Threlfall (1976), as was the treatment
of any parasites found. Mites were sampled by examining 10 feathers from
each region. The classification of the Mallophaga listed herein follows the
scheme of Emerson (1972b, c). Data were analysed using the ¢ statistic for
two means, the Chi-squared evaluation and 2 x K contingency tables. Spec-
imens of the Mallophaga found have been deposited in the National Museum
of Natural History, Washington, D.C. (Nos. 75898-75900), while represen-
tative ticks and an Ornithonyssus sp. are in the collection of Harry Hoog-
straal, NAMRU 3, Cairo, UAR (Nos. HH22,071—HH22,074).

RESULTS AND DISCUSSION

Six species of ectoparasites were taken from the 77 birds examined (34
(44%) infested). Birds from the west coast bore the Mallophaga Eidman-
niella pellucida (Rudow) (11% birds infested), Piagetiella incomposita (Kel-
logg and Chapman) (27% infested), and Pectinopygus farallonii (Kellogg)
(43% infested). The first two species are amblycerans (Menoponidae), while
the latter is an ischnoceran (Philopteridae). East coast birds were host to 1
mallophagan, | tick, and 2 mites (Table 1). The difference noted in the
species composition of the ectoparasite fauna of the two groups may be
attributed to differences in sampling time (month and/or year), to geograph-
ical factors, or to migratory vs. permanent resident subspecies of host.

Table 2 gives details of the distribution of the most common mallophagan
P. farallonii, on the 33 nestling and adult birds. Statistically significant dif-
ferences (P < .005) were noted in the distribution of adults and nymphs,
and in the infestations with adults and nymphs, on both nestling and adult
birds. No differences were detected in the infestations of nestlings as com-
pared to adult birds, other than the fact that no Mallophaga were located
on the crown, auricular, and gular region of nestlings. Far greater numbers
of mallophagan nymphs were found than adults, this perhaps being related
to an increasing population and transfer of organisms from adult birds to

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VOLUME 84, NUMBER 2

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374 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

nestlings with reproduction occurring to rebuild the populations. The sex
ratio of the adult mallophagans was | male: 1.5 females, once again indi-
cating that the population is most probably an expanding one.

Argas radiatus (Railliet) (Argasidae) larvae were found only on chicks,
and in greatest numbers (up to 271 individuals) on birds which were only a
few days old and lacking all but a thin coat of down. No adults of this
species were recovered. A statistically significant difference (P < .001) was
noted in the distribution of these organisms on the host (Table 3). No ticks
or mites were located on the crown, auricular or gular regions of the birds.
The mites did, however, show a distinct preference for various regions of
the body as evidenced by the Michaelichus sp. which was located only on
the primaries and secondaries of the wings.

While the foregoing account shows that differences in the ectoparasite
burden may vary from region to region on a host, between different ages of
host, and from one geographic location to another, we are not yet able to
explain the reasons underlying these differences and much work remains to
be done to elucidate these anomalies.

ACKNOWLEDGMENTS

Thanks are extended to the Natural Sciences and Engineering Research
Council of Canada for the grant (NSERC-A3500) that funded the field work.
The study was supported in part by Research Grant No. 1270-G from the
Florida Game and Freshwater Fish Commission’s Federal Aid to Wildlife
Restoration Program, Florida Pittman-Robertson Project W-4. Thanks are
also due to D. J. Forrester of the College of Veterinary Medicine, University
of Florida, Gainesville, in whose laboratory the work was undertaken by
the author during the tenure of an Adjunct Professorship. H. Hoogstraal,
C. M. Clifford, H. Y. Wassef, W. T. Atyeo, and C. E. Bourgeois aided in the
identification of the parasites, and R. A. Anderson, G. W. Foster, P. P.
Humphrey, and S. A. Nesbitt provided field and laboratory assistance.

LITERATURE CITED

Emerson, K. C. 1972a. Checklist of the Mallophaga of North America (North of Mexico). Part
IV Bird host list. Deseret Test Center, Dugway, Utah, 216 pp.

——. 1972b. Checklist of the Mallophaga of North America (North of Mexico). Part I
Suborder Ischnocera. Deseret Test Center, Dugway, Utah, 200 pp.

—.. 1972c. Checklist of the Mallophaga of North America (North of Mexico). Part II
Suborder Amblycera. Deseret Test Center, Dugway, Utah, 118 pp.

Eveleigh, E. S. and W. Threlfall. 1976. Population dynamics of lice (Mallophaga) on auks
(Alcidae) from Newfoundland. Can. J. Zool. 54: 1694-1711.

Marshall, A. G. and B. C. Nelson. 1967. Bird ectoparasites from South Farallon Island,
California. J. Med. Entomol. 4: 335-338.

Price, R. D. 1970. The Piagetiella (Mallophaga: Menoponidae) of the Pelecaniformes. Can.
Entomol. 102: 389-404.

VOLUME 84, NUMBER 2 375

Ryan, S. O. and R. D. Price. 1969. A review of the genus Eidmanniella (Mallophaga: Meno-
ponidae) from the Pelecaniformes. Ann. Entomol. Soc. Am. 62: 815-823.

von Timmermann, G. 1967. Gruppen Revisionen bei Mallophaga VII. Die Pectinopygus-Arten
der Groskormorane (Gen. Phalacrocorax Brisson, 1760 s. st.). Mitt. Hamb. Zool. Mus.
Inst. 64: 83-85.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 376-390

DENSITY, FECUNDITY, HOMOGENEITY, AND EMBRYONIC
DEVELOPMENT OF GERMAN COCKROACH
(BLATTELLA GERMANICA (L.)) POPULATIONS IN
KITCHENS OF VARYING DEGREES OF SANITATION
(DICTYOPTERA: BLATTELLIDAE)'

D. A. SHERRON, C. G. WRIGHT, M. H. Ross, AND M. H. FARRIER

(DAS) Department of Biology, University of Notre Dame, Notre Dame,
Indiana 46556; (CGW, MHF) Department of Entomology, North Carolina
State University, Raleigh, North Carolina 27650; (MHR) Department of
Entomology, Virginia Polytechnic Institute and State University, Blacks-
burg, Virginia 24060.

Abstract.—Populations of the German cockroach, Blattella germanica
(L.), were mass-collected on a seasonal basis in low-income apartments in
Raleigh, North Carolina. Numbers of oothecae and each sex and instar were
enumerated. Oothecae collected in June were examined for embryonic de-
velopment.

Density showed a significant correlation with sanitation. However, certain
kitchens had fewer cockroaches than expected at sanitation rating #6 (poor-
est sanitation) and others with good sanitation (rating #1) had more than
expected. Possible explanations of these occurrences were based on anal-
ysis of age class frequencies. A structure apparently indicative of a stabi-
lized population occurred at high density in sanitation 6 and low density at
sanitation 1. Variability of age class structure stemmed from the individual
kitchen infestation, nevertheless, there were several general characteristics.
Adults and first-instars were usually the largest groups and nymphal mor-
tality was highest among first-instars. The latter was a secondary factor in
regulation of population growth, with primary control occurring at the level
of oothecal production.

The German cockroach, Blattella germanica (L.), has been the subject of
numerous biological studies (Cornwell, 1968). For example, B. germanica

"Contribution from the North Carolina Agricultural Research Service. Published with ap-
proval of the Director of Research as paper no. 6789 in the Journal Series. Send reprint requests
to C. G. Wright.

VOLUME 84, NUMBER 2 377

is known to shun light (Gunn, 1940), to prefer temperatures of 24-33°C
(Gunn, 1935), and, indeed, to avoid unfavorable temperatures in choosing
its habitat (Ogata, 1976). Water and humidity also affect habitat selection,
as seen in a preference for both warm and steamy conditions (Gunn and
Cosway, 1938; Cornwell, 1968). Less is known concerning factors that affect
the growth of natural populations. Laboratory studies showed an increase
of nymphal mortality with density (Chauvin, 1946). Under uncrowded con-
ditions, Ross (1976) and Willis et al. (1958) found 83-85% of nymphs that
hatched matured, although Ross (1929) reported mortalities of 5% for early
molts and 40% for the last molt. An analysis of age class frequencies of
collections from single family homes showed little decrease in numbers be-
tween first and last stage nymphs, suggesting low mortality (Ross and
Wright, 1977). These collections were characterized by high proportions of
non-productive females. It was suggested that failure to mate or produce
egg cases, known results of insufficient food/water (Kunkel, 1966; Roth and
Stay, 1962), were major factors in controlling growth of these populations.
Moreover, it appeared that an adequate supply of water might be the most
critical factor in the survival of early instars. High cockroach populations
were correlated with poor sanitation (Wright, 1979).

This paper reports on a year long study of German cockroach populations
in apartments characterized by six sanitation ratings. Selected data on cock-
roach age composition, density, fecundity, homogeneity and embryonic de-
velopment, and temperature and relative humidity in apartments are pre-
sented. The data increase our understanding of free populations and provide
information which can be used in the design of cockroach pest management
systems and the evaluation of their success.

MATERIALS AND METHODS

Four low-income apartment complexes in Raleigh, North Carolina, were
used as research sites. Two factors determined apartment selection within
each complex: 1) Presence, verified by visual inspection during the day, of
numerous cockroaches in an apartment, and 2) permission of the tenants,
since the apartments were sampled while the tenants occupied them. In-
spection and response of tenants indicated that commercial pest control
technicians had not recently disturbed the cockroach populations.

Three apartment complexes, designated A, B, and C, had almost identical
construction and floor plans. Each complex was composed of brick and
concrete, and contained a number of apartments with tiled concrete floors
in the kitchen and bathroom and concrete floors elsewhere. Some apart-
ments were interconnected by holes around stove fixtures, and some tenants
spoke of spaces behind cabinets and sink areas. The fourth complex (D)
had fewer apartments in each building and more distance between buildings.
The floor plan was slightly different from complexes A, B, and C, with much

378 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

less wall space between kitchens and living rooms and long, wall-sized glass
doors on one side of the kitchen. All floors had tile on concrete. Apartments
in all complexes had small bathrooms, two floor levels, and continuous
stove and cabinet areas along three walls of the kitchen.

Collections in September and December of 1977 were made in apartments
in one complex, while those in March and June of 1978 were from apart-
ments in all four complexes where no previous collecting had occurred.
Collecting during four months (Sept., Dec., Mar., June) provided seasonal
information on German cockroach populations. Six apartments per complex
were sites for September and December collections, while three apartments
in each of the four complexes were sites for March and June collections.
All collecting occurred during the early afternoon and on different days in
the month.

Collections immediately followed crack-and-crevice treatments with a
synthetic pyrethroid flushing agent, resmethrin. A modified vacuum cleaner
(Wright, 1966) was used to collect the cockroaches. Extensive flushing and
vacuuming were done where an initial, visual examination revealed abun-
dant cockroaches which appeared evenly distributed throughout a room. In
areas of spotty infestation, such as occurred in most bedrooms, crevices in
furniture and closets were flushed. Cockroaches were separated immedi-
ately according to size of sieves of the USA Standard Sieve Series, numbers
7, 8, and 14. Specimens and oothecae were preserved in 70% ethanol for
later classification.

Terminal sternum characters (Ross and Cochran, 1960) and the number
of antennal doublets (Campbell and Priestly, 1970) were used for instar and
sex determinations with a Wild-Heerbrugg M3 microscope at 40 magnifi-
cation. With older nymphs, instar determinations were made by a judgement
of their relative size after sexing, as breakage of antennae usually precluded
use of doublets for instar determination. By combining fourth- and fifth-
instars, the problem of classifying damaged, middle-sized nymphs was
avoided.

At the time collections were made, an Atkins Thermistor Psychrometer,
Model Number 3702B, was used to record ambient humidities and temper-
atures in each kitchen, living room, bathroom, and bedroom. A sanitation
scale (Bennett, 1978; Table 1) was utilized to classify rooms in apartments
according to sanitation.

The methods of Barr et al. (1976) were used to analyze information rel-
ative to the effects of humidity, temperature, and sanitation level on pop-
ulation density and age structure among German cockroaches. F-values
were compared at the 0.05 level of significance.

Numbers of first-instar nymphs, oothecae, and adult females, and in the
case of June collections, the numbers and progress of developing embryos,
permitted fecundity comparisons among German cockroach populations.

VOLUME 84, NUMBER 2 379

Table 1. Sanitary rating scale (Bennett, 1978).

Scale Sanitary Condition

1. Fairly clean, not cluttered Floors not very dirty, shelves and cupboards not clut-
tered, except for normal amount of content, no obvious
piles of trash.

ie)

. Fairly clean and cluttered Floors fairly clean but may be cluttered with trash,
clothes, etc. (nongarbage); cupboard filled with non-
garbage articles.

Ww

. Generally dirty, not cluttered Floors generally dirty and/or greasy; cupboards dirty and
not washed out for some time.

a=

. Generally dirty and cluttered Floors generally dirty and piled with nongarbage trash,
clothing, or small amounts of garbage; cupboards not
washed out, dirty, and filled with nongarbage articles.

an

. Severely dirty, not cluttered Very dirty with garbage obvious; area very greasy and
uncleaned in a long time; dead cockroaches obvious
and not swept up.

an

. Severely dirty and cluttered Very dirty with garbage obvious and piled around, greasy
areas filled with articles, trash, papers, etc. (which
make counting difficult); dead cockroaches obvious and
not swept up.

The presence or absence of eye pigment was used to determine the extent
of embryo development in oothecal chambers (Tanaka, 1976). In addition,
various estimates of first-instar nymph production during selected months
for each apartment complex and season aided productivity comparisons. It
was assumed throughout that productivity did not vary within a complex
during a month’s time.

Estimates of monthly nymphal production were calculated three ways.
The first method was based on the numbers of first-instar nymphs collected.
The first nymphal stadium lasts about five days (Woodruff, 1938). Monthly
hatch was considered to be composed of six equal ‘‘waves’’ of first-instar
nymphs (30 days per month + 5 days per first-instar = 6), giving rise to
the following formula: No. of first-instar nymphs per month = (No. of first-
instar nymphs counted in one wave) X (6 waves per month).

The second method was based on numbers of oothecae collected. Using
a hatch estimate of 76% (Willis et al., 1958) and the mean of 37.6 embryos
per ootheca (Table 2) the formula was: No. of first-instar nymphs per
month = (No. of oothecae per month) x (28.6 nymphs emerging per ooth-
eca).

The third method was based on the reproductive potential of all adult
females collected. Assuming that a female German cockroach produces one
egg case per month (Cornwell, 1968), and applying the hatch estimate of

380 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Estimates of numbers of first-instar nymphs hatching in a month per apartment
complex, based on numbers of first-instar nymphs, oothecae, and adult females collected during

each season.

Method of Calculation

First-Instar Adult

Nymphs?* Oothecae” Females
Month Complex Estimated Number

September A 9708 9667 22,451
December B 16,728 19,191 40,869
March A 5610 4919 11,268
B 3246 5834 12,899

€ 8644 9381 22222

D 6258 16,645 34,063

X 5940 9195 20,113

June A Sil 2889 7550
B 3072 4004 10,096

C 4512 9095 21,078

D 11,094 7808 18,847

¥ 5948 5949 14,392

4 Based on the formula: No. of first-instar nymphs per month = (No. of first-instar nymphs
counted in one wave) X (6 waves per month).

> Based on the formula: No. of first-instar nymphs per month = (No. of oothecae) x (28.6
nymphs emerging per ootheca).

© Based on the formula: No. of first-instar nymphs per month = (No. of adult 2) x (28.6
nymphs emerging per ootheca).

Willis et al. (1958), and the mean number of embryos per ootheca (Table 2),
the formula was: No. of first-instar nymphs per month = (No. of adult
2’s per month) Xx (28.6 nymphs emerging per ootheca).

RESULTS AND DISCUSSION

Ambient relative humidity of rooms was variable, ranging from 25-83%,
with an average of 54.5%. Means were lower in December (52.5) and March
(42.6) and higher in June (64.7) and September (63.3). The humidity in cracks
and crevices is assumed to have varied similarly on a seasonal basis. Pre-
liminary measurements under the kitchen sinks in September were nearly
identical to the ambient values for kitchens. Neither relative humidity nor
the related factor of temperature showed a correlation with either the den-
sity or age structure of the populations at the 0.05 level of significance. It
is doubtful if either of these factors exerted much effect on the populations.
The mean temperature of 26.1°C and range of 21-30° were close to the
optimal temperature of about 25° (Gould, 1941) and the optimal range of
24—33° (Gunn, 1935), respectively. Providing the cockroach has water to
drink, its response to humidity is weak (Gunn and Cosway, 1938).

|

VOLUME 84, NUMBER 2 381

@) 2s oO O)

ie)

Sanitary rating a

O 400 800 1200 1600 2000 2400 2800 3200 3600 4000 4400

Upper limits of intervals for numbers of cockroaches collected®

Fig. 1. Numbers of German cockroach samples collected in kitchens in 36 low-income
apartments arranged by class size and sanitary rating. The numbers 1, 2, 3, and 4 refer to the
total number of kitchens where samples were collected with a specific sanitary rating and a
designated range of cockroaches. For example, there were 3 kitchens where samples were
collected which had a sanitary rating of 3 and the number of observed cockroaches was over
400, but less than 800. A, Based on a sanitation scale developed by Bennett (1978). B, For
example, an upper limit of 400 includes an interval of 1-400 cockroaches.

Sanitation had a significant effect (0.05) on population density in kitchens.
In apartments rated poor in sanitation, greater populations of cockroaches
were present (Fig. 1). One-third of the kitchens had the poorest possible
sanitation rating (rating of 6). Three of these kitchens with poor sanitation
contained cockroach populations exceeding 2400 cockroaches. The German
cockroach is typically active in darkness (Gunn, 1940), yet kitchens with
large cockroach populations had many individuals active in the day. Crowd-
ing in shelter areas could make this unusual behavior necessary. Productiv-
ity in kitchen populations was not affected by sanitation as rated on Ben-
nett’s sanitary scale (see Table 1). Cockroach populations in kitchens with
sanitary ratings of 4-6 produced an estimated average of 22.9 first-instar
nymphs emerging per ootheca, while populations in kitchens with sanitary
ratings of 1-3 produced an estimated average of 23.5 nymphs emerging per
ootheca, based on the number of oothecae and first-instar nymphs collected
in the kitchens. Populations in apartments with unsanitary conditions (4-6
on the sanitary rating scale) averaged 36.7 developing embryos per ootheca,
while those in apartments with better sanitation (1-3 on the sanitary rating
scale) averaged 38.2 embryos per ootheca.

The data have implications for population dynamics in addition to those

382 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Development of German cockroach oothecae collected in kitchens in low-income
apartments in June 1978.

No. Oothecae with Embryos

Mean No. Live Having Eye Pigment”
No. Oothecae Embryos
Complex Examined Per Oothecae* Visible Not Visible
A 26 SY/ {Uae Sis! 16 10
B 35 SB) ye) aed i 28 7
€ 55 Sy/ Sy ae D2: Si 18
D 66 39.7 + 0.7 47 19
x S40) Be PL) 32 14

a Based on the shape and pigment of oothecal compartments. Mean and standard deviation.
> Presence of eye pigment indicates less than 10 days until hatch.

noted above related to environmental factors. Comparison of the three types
of monthly nymphal production in Table 2 gives an indication of the relative
importance of hatch-survival of first-instars vs. egg case production as
mechanisms that limit population growth. Estimates of monthly hatch based
on observed numbers of first-instars are shown in column 3. These estimates
tended to be lower than those based on the total number of oothecae x the
average number of nymphs emerging from an ootheca in the laboratory
(column 4). However, the magnitude of the difference was frequently small,
especially in lower density apartments in complexes A and B. Apparently
neither hatch nor immediate post-hatch survival were drastically reduced in
these apartments. In contrast, March data from complex D and June data
from complex C suggest a greater hatch reduction and/or post-hatch mor-
tality. In these instances monthly hatch as estimated from observed numbers
of first-instars (column 3) was less than half that expected from estimates
based on the number of oothecae (column 4). Surprisingly, June results from
complex D showed the reverse situation. In complex D, it appears that
populations were growing very little in March, but did increase in growth
in June. If collections were made at a time of first egg case hatch of a large
proportion of the females, overall monthly hatch would be over-estimated.
The one ‘‘wave’’ would be larger than the others. Alternatively, it would
be difficult to account for a higher hatch from estimates based on observed
numbers of first-instars than those from laboratory hatch. The large number
and low variability of egg cases of June collections (Table 3, complex D)
support this hypothesis of high input from first egg case hatch. Whether
these differences between population structure-growth in complex D and
the other three complexes are related to structural differences noted earlier
is unknown.

The hatch estimates shown in Table 2 (column 5) are based on an as-
sumption that all adult females were reproductively active. The disparity
between these data and the estimates based on numbers of oothecae (column

VOLUME 84, NUMBER 2 383

Complex C Complex D
9 of
March
(3,962)
10 (8) 10

Adults : J

June
(5,203)

10 0 10

% of cockroaches captured

Fig. 2. Instar-sex pyramids of German cockroaches collected in kitchens during four sea-
sons of 1977-1978. Numbers in parentheses (A) next to the pyramids are the total numbers of
captured cockroaches.

4) indicates that less than one-half of the adult females were producing
viable oothecae. In this species, limited availability of food causes failure
to mate (Roth and Stay, 1962) and failure of mated females to produce
oothecae (Kunkel, 1966). Either limited food and/or water in relatively san-
itary conditions or limited access to these resources due to crowding in
unsanitary conditions could account for the heavy proportion of reproduc-
tively inactive females. A few may have been past the productive part of
their life span (Willis and Lewis, 1957; Willis et al., 1958), although death,
rather than failure to produce viable oothecae, is the major age-related cause
of reproductive failure among females of this species (Cochran and Ross,
unpublished data). The present data provide strong evidence that major

384 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 4.
kitchens by sanitation grade.

Number of

Nymphs by Instar*

Apts. Oothecae* Ist 2nd Apts.

Sanitation Grade |

Oothecae and first- and second-instar German cockroach nymphs collected in

Number of
Nymphs by instar®

Oothecae* Ist 2nd

Sanitation Grade 4

9 33 151 83 3 95 240 139
(10-67) (25-269) (11-149) (88-109) (102-364) (73-173)
Sanitation Grade 2 Sanitation Grade 5
6 38 89 48 l 230 499 239
(9-79) (18-219) (15-101) (0) (0) (0)
Sanitation Grade 3 Sanitation Grade 6
4 39 195 99 13 114 410 192
(27-59) (124-239) (52-209) (33-369) (104-1077) (38-531)
xX per Sanitation Grades 1-3 xX per Sanitation Grades 4-6
19 36 141 76 17 101 385 185
(9-79) (18-269) (11-209) (33-369) (102-1077) (38-531)

@ Mean with range in parentheses.

control of population density was at the level of oothecal production. Sec-
ondary factors were reduced hatch and/or post-hatch mortality (Table 2,
columns, 3 vs. 4) and nymphal mortality causing a drop between numbers
of first- and second-instars (Fig. 2). Keil (1981) also noted a tendency
towards high proportions of non-productive females among shipboard pop-
ulations.

Mortality between the first- and second-nymphal instars ranged from
45-53% when the apartments were grouped together by sanitation grade
(Table 4). The 5% mortality reported by Ross (1920) for early instars is
much less than that observed in these complexes. There was no significant
difference in the percent of nymphal mortality between the first- and second-
nymphal instars when compared by kitchens grouped into the sanitary
grades 1-3 and 4-6. Mortality was 45.2% and 47.9% respectively. However,
there was extensive variation among individual collections (see below). The
number of oothecae collected in apartments grouped by sanitation grade
also varied widely, but more oothecae were collected in apartments with
sanitation grades of 4-6 than in those with grades of 1-3 (Table 4).

Examination of oothecae collected in June from apartments in the four
complexes showed consistency in the number of live embryos per ootheca,
with an average of 37.6 live embryos per ootheca (Table 3), which is less
than the 45 embryos reported by Tanaka (1976). Eye pigment, indicating
nymphal emergence from the oothecae would occur in 10 or fewer days,
was visible in 70% of the oothecae examined.

VOLUME 84, NUMBER 2 385

Table 5. Comparative densities in collections at the two extremes of the sanitation ratings,
i.e., best sanitation (#1) vs. poorest sanitation (#6).

Number of Apartments With Ratings of:

Cockroaches* #1 #6

>2000 3
1500-2000 | |

4 Rounded to nearest 100.

Continuous maturation of new adults and egg case hatch indicated com-
plete overlap of generations. In this situation, there are general patterns of
age class composition that characterize populations at or near balance for
their particular environmental situation. Patterns of age class frequency
should give an indication of the state of the population, i.e., whether sta-
bilized, rapidly growing, or, perhaps, affected by extrinsic influences (in the
present case, tenant’s efforts at control). Such patterns would only be re-
vealed through analysis of individual collections, since these were the
source of significant differences in age composition. In order to explore this
possibility, collections from contrasting sanitation ratings (1 vs. 6) were
compared. It was assumed those with maximum numbers for the particular
environment would give the best indication of the age composition of a
stabilized population.

Collections ranging from 300-900 predominated at sanitation | (Table 5).
In contrast, at sanitation 6 most fell within a range of 900-4000, with 900
apparently on the low side for poor sanitation. Age composition of the higher
density collections differed from that of the smaller collections. Among the
former, at least 25% were first-instars (Fig. 3A). A drop of over 12% oc-
curred between first- and second-instars and, in all except one collection,
first-instars outnumbered adults (difference ranging from 6—16%). In con-
trast, first-instars formed less than 25% of sanitation 6, low density collec-
tions. The drop between first- and second-instars was smaller (<12%), ex-
cept in one intermediate density collection (Fig. 3A—952), and adults
outnumbered first-instars consistently. The unifying feature found at sani-

386 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

40 1168 x
35
997x x 1460
3573 x x 2583
Sa 1254x A
25 xX 4129 SANITATION # 6
x )950
20 670 591 X 952 e (950
15 393 °710
25 °779

INSTARS

% FIRST

B

SANITATION # 1
x )950
e (950

5 10 15 20" (25°30

% DROP BETWEEN 1st AND 2nd INSTARS

Fig. 3. Comparison of the proportion of first-instar German cockroaches and first-instar
mortality at sanitation 6 vs. 1 for low and high density groups. A, Differences between high
and low density groups at sanitation 6; characteristics of high density (x) presumably are those
of a stabilized population. B, Characteristics of presumed stabilized populations which occur
among low density groups at sanitation | (first-instars >25%).

VOLUME 84, NUMBER 2 387

Apt. 1 Apt.2
9 Adults o 0) Adults o
6 6
4-5 Sanitation 1 4-5
3 3
2 2
“eye
20 10 O 10 O
Apt. 3
Q | Adults | o
6
4-5 Sanitation 6
3
2
10 oO 10

% of cockroaches captured

Fig. 4. Examples of German cockroach age class frequencies from single apartments. Apts.
1 and 3 are typical of presumed stabilized type as seen at sanitation | and 6, respectively. Apt.
2 shows a composition that may be indicative of a recent immigration. Apt. 4 illustrates the
structure typical of low density, sanitation 6 groups, as well as some at sanitation |, in which
reproduction was apparently more limited than in the type shown in apts. | and 3.

tation 6 was that high density apparently characterizes populations that are
essentially stabilized for their particular environmental situation. Therefore
the age composition of the low density collections was examined for clues
as to why they were smaller than expected and whether they might provide
insight into situations where sanitation and density do not show a close
correlation. Recent introductions would represent expanding populations,
with higher proportions of immatures than in stabilized groups. This pos-
sibility was ruled out since none of the low density groups had higher pro-
portions of nymphs than the high density groups. Rather, age class fre-
quencies suggested a more limited reproduction. The proportion of first-
instars was smaller, as in the example shown in Fig. 4, Apt. 4. Possible

388 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

explanations are that (1) tenant’s control efforts decreased hatch or caused
high mortality among first-instars or (2) age composition reflected a second
type of stabilized population. The latter might occur if there was a difference
in the relative impact of limiting factors, as suggested earlier in connection
with differences in age class frequencies of ‘‘field’’ and laboratory popula-
tions (Ross and Wright, 1977). At high density, harborage space may have
been an important limiting factor. Among the sanitation 6, low density
groups, the rating might have reflected abundant harborage space (clutter),
obscuring limits on the supply of resources affecting reproduction (food and/
or water). If so, reliable estimates of density for particular environmental
situations may require refining sanitation ratings in terms of relative avail-
ability of the three resources, i.e., food, water, and harborage.

Figure 4, Apts. 1 and 3, shows examples of the general age class com-
position that apparently reflects at least one type of stabilized population.
As expected, such occurred among several of low density (600) at sanitation
1 (Fig. 3B, first-instars >25%) as well as sanitation 6, high density groups
(Fig. 3A). Another small collection (620) at sanitation 1 was similar to the
sanitation 6, low density type noted above and illustrated in Fig. 4, Apt. 4,
in which reproduction was apparently more limited. The smallest collection
showed the pattern illustrated in Fig. 4, Apt. 2. The extremely high pro-
portions of late instars may be indicative of a recent invasion, possibly
dispersal from an adjacent apartment. Data from a shipboard experiment
suggested more widespread dispersal of middle to late instars than other age
groups following an insecticide treatment (Ross et al., 1981). The largest
collection (1947) was close in number and structure to that stabilized at high
density, sanitation 6. Other large collections for sanitation | tended to be
intermediate in that first-instars were <25% but generally higher than the
11-20% range typical of those with apparent limited reproduction. A pos-
sible explanation of both unusually high densities for sanitation | and incip-
ient limits on reproduction is a recent change for the better in house-keeping
habits.

The data presented here and those published earlier (Ross and Wright,
1977) provide some of the first evaluations of the effects of density-inde-
pendent factors (temperature and humidity) and density-dependent factors
(food/water) on density and age class structure. We suggest that the latter,
when considered in conjunction with an evaluation of the environmental
resources, may prove to be a useful tool in evaluating the state of the pop-
ulation (stable, growing, new introduction, decreasing) and, possibly, as-
sessing the effects of control measures. The evidence of population control
at the level of oothecal production is primarily important in respect to re-
covery potential. Lowered density would stimulate reproduction among an
increased proportion of the adult females. This would engender far more
rapid growth than if expansion depended on decreased nymphal mortality,

VOLUME 84, NUMBER 2 389

with the necessity for nymphs to mature and reproduce before the popula-
tion could undergo significant increases in size.

We note an absence of morphological deviants that might have been in-
dicative of mutant phenotypes as this could be helpful if future studies lead
into areas of population genetics.

LITERATURE CITED

Barr, A. J., J. H. Goodnight, J. P. Sall, and J. T. Helwig. 1976. A user’s guide to SAS
[Statistical Analysis System]. Sparks Press, Raleigh, N.C. 329 pp.

Bennett, G. W. 1978. Evaluating pesticides in urban environments. Chem. Times and Trends
2: 55-61.

Campbell, F. L. and J. D. Priestly. 1970. Flagellar annuli of Blattella germanica. Ann. Ento-
mol. Soc. Am. 63: 81-88.

Chauvin, R. 1946. Notes sur la physiologie comparée des Orthopteres. Bull. Soc. Zool. Fr.
71: 39-48.

Cornwell, P. B. 1968. The Cockroach, Vol. 1. Hutchinson and Co., London. 391 pp.

Gould, C. E. 1941. Effect of temperature on the development of cockroaches. Proc. Indiana
Acad. Sci. 50: 242-250.

Gould, C. E. and H. O. Deay. 1940. The biology of six species of cockroaches which inhabit
buildings. Purdue Univ. Agric. Exp. Stn. Bull. No. 451.

Gunn, D. L. 1935. The temperature and humidity relations of the cockroach. III. A comparison

of temperature preference, rates of dessication and respiration of Periplaneta ameri-

cana, Blatta orientalis and Blattella germanica. J. Exp. Biol. 12: 185-190.

. 1940. Daily activity rhythm of the cockroach. J. Exp. Biol. 17: 267-277.

Gunn, D. L. and C. A. Cosway. 1938. The temperature and humidity relations of the cock-
roach. V. Humidity preference. J. Exp. Biol. 15: 555-563.

Keil, C. B. 1981. Structure and estimation of shipboard German cockroach (Blattella
germanica) populations. Environ. Entomol. 10: 534-542.

Kunkel, J. G. 1966. Development and the availability of food in the German cockroach,
Blattella germanica (L.). J. Insect Physiol. 12: 227-235.

Ogata, K. 1976. Studies on establishing factors of domiciliary cockroaches. 2. Experimental
observations of development of German cockroaches in various environments. Jpn. J.
Sanit. Zool. 27: 411-421. In Japanese, English Summary.

Ross, H. H. 1929. The life-history of the German cockroach. Trans. Ill. State Acad. Sci. 21:
84-93.

Ross, M.H. 1976. Laboratory population studies of the German cockroach using a two-chromo-
some and a three-chromosome reciprocal translocation. Ann. Entomol. Soc. Am. 69:
1973-1981.

Ross, M. H. and D. G. Cochran. 1960. A simple method for sexing nymphal German cock-
roaches. Ann. Entomol. Soc. Am. 53: 550-551.

Ross, M. H. and C. G. Wright. 1977. Characteristics of field-collected populations of the
German cockroach. Proc. Entomol. Soc. Wash. 79: 411-416.

Ross, M. H., C. B. Keil, and D. G. Cochran. 1981. The release of sterile males in natural
populations of the German cockroach. Entomol. Exp. Appl. 30: 241-253.

Roth, L. M. and B. Stay. 1962. Oocyte development in Blattella germanica (L.) and Blattella
vaga Hebard (Blattaria). Ann. Entomol. Soc. Am. 55: 633-642.

Tanaka, A. 1976. Stages in the embryonic development of the German cockroach Blattella
germanica (Blattaria, Blattellidae). Kontyd 44: 512-525.

Willis, E. R. and N. Lewis. 1957. The longevity of starved cockroaches. J. Econ. Entomol.
50: 438-440.

390 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Willis, E. R., G. R. Riser, and L. M. Roth. 1958. Observations on reproduction and devel-
opment of cockroaches. Ann. Entomol. Soc. Am. 51: 53-69.

Woodruff, L. C. 1938. The normal growth rate of Blattella germanica. J. Exp. Zool. 79:
145-165.

Wright, C. G. 1966. Modification of a vacuum cleaner for capturing German (Blattella ger-

manica) and brown-banded (Supella supellectilum) cockroaches. J. Econ. Entomol. 59:

759-760.

. 1979. Survey confirms correlation between sanitation and cockroach populations. Pest

Control 47: 28.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, p. 390

NOTE

A 2000+-Year-Old Beetle (Coleoptera: Dermestidae)

Robert Brier, Philosophy Department, Long Island University, Green-
vale, N.Y., recently submitted for identification nine adult specimens of a
dermestid beetle found inside a wrapped, mummified cat. I identified them
as Dermestes frischii Kugelann, a widely distributed species in Europe,
Asia, Africa, and North America. The larvae of this species feed on car-
casses, bones, dried fish, and a number of other substrates, usually dead
animal matter, and have been found in mummies (Hope. 1834. Proc. Ento-
mol. Soc. Lond. 1834: 11-13; Lesne. 1930. Bull. Soc. Entomol. Egypte 1930:
21-24).

The exact origin of the mummified cat is not known, but Brier placed the
date at 332 BC to 30 BC based on the wrapping which he regards as char-
acteristic of the Ptolemaic dynasty in Egypt. The beetle specimens would
thus be 2011 to 2313 years old.

John M. Kingsolver, Systematic Entomology Laboratory, HBII, Agri-
cultural Research Service, USDA, c/o National Museum of Natural His-
tory, Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 391-396

NEW SPECIES OF XESTOCEPHALINAE
(HOMOPTERA: CICADELLIDAE) FROM
MEXICO, PANAMA, PERU, AND BRAZIL

DwiGHT M. DELONG

Department of Entomology, The Ohio State University, Columbus, Ohio
43210.

Abstract.—Three new species of Portanus, P. bilineatus (Peru), P. spi-
nosus (Bolivia), and P. caudatus (Panama), and three new species of Xes-
tocephalus, X. variabilis (Mexico), X. dubius (Bolivia), and X. similis (Bra-
zil), are described.

Most species of both Portanus and Xestocephalus have rather definite
color patterns. Miguel Estribi, working in the tropical laboratory of the
Smithsonian Institution in Panama, has been able to separate the 25, or
more, species of Xestocephalus known to occur in Panama by their color
patterns.

Species of Portanus have previously been treated by Linnavuori (1959),
Kramer (1964), and DeLong (1980). Species of Xestocephalus have previ-
ously been treated by Linnavuori (1959), DeLong and Linnavuori (1978),
and DeLong et al. (1980).

Three species of Portanus and three species of Xestocephalus are de-
scribed in this paper. All types are in the DeLong collection.

Portanus bilineatus DeLong, NEw SPECIES
Figs. 1-4

Description.—Length of male 6 mm, female unknown. Crown bluntly
angled, apex rounded, a little longer at middle than wide at base between
eyes. Crown except marginal gray area, mostly white with a black squarish
spot, each side, just above and mesad of ocelli and a broad longitudinal dark
brown stripe extending from brown spot to base along median line. A narrow
lateral band extends from broad stripe, just basad of ocelli, then basally
along each eye. Pronotum white with 4 dark brown, broad, longitudinal
stripes: 2 at middle, widening near posterior margin, and | each side behind
eye, along lateral margin to apex, then curved to merge with median stripe
along posterior margin of pronotum. Scutellum pale gray with darker basal

392 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

angles. Forewing brownish subhyaline, marked with white and darker
brown spots.

Male genitalia, style elongate with main rod-shaped portion narrowed
apically and finger-like, apex bluntly pointed. A short process arises at half
length of blade, extending laterocaudally, apex bluntly pointed. Aedeagus
short and broad, open anteroventrally, bearing a dorsal spine 4 length of
aedeagus which extends ventrally, along caudal margin.

Holotype.—d, Sinchono, Peru, November 1934, J. G. Sanders, coll.

Comments.—Portanus bilineata resembles P. minor Kramer in general
form and appearance and can be separated by the brown longitudinal stripes
on the crown and pronotum and by the aedeagus which bears a spine at the
apex.

Portanus spinosus DeLong, NEw SPECIES
Figs. 5-8

Description.—Length of male 4.5 mm, female 5 mm. Crown produced
and bluntly angled. Crown brownish yellow. Pronotum dark brown with
numerous small whitish spots. Scutellum yellowish with basal and apical
angles brown. Forewing brownish subhyaline with dark brownish spots or
areas, especially on costa and basal crossveins of apical cells. Most veins
with alternating dashes of brown and white.

Female posterior margin of 7th sternum broadly convexly rounded.

Male genitalia, style narrow, elongate, apical portion pincer-like with the
2 portions of about equal length. Aedeagus slender, curved and elongate
with apex bearing 4 slender apical processes. In apical view the median
portion appears serrate just before slender, pointed, apical tip.

Types.—Holotype: 6, Santa Cruz, Bolivia, 21-I[X-1980, D. Foster coll.
Paratypes: 5 d, 1 2, same data as holotype; | 6, same data except 17-VIII-
1980.

Comments.—Portanus spinosus resembles P. lex Kramer in form and
appearance, but P. spinosus can be separated by the long slender curved
aedeagus.

Portanus caudatus DeLong, NEW SPECIES
Figs. 9-11

Description.—Length of male 6 mm, female unknown. Crown produced,
apex rounded, a little longer at middle than wide at base, between eyes.
Crown white with a pair of small round black spots between ocelli at apex,
just above margin, and an orange area just behind each black spot. Pronotum
white with faint orange markings. Scutellum white. Forewing whitish sub-
hyaline with faint orange markings on clavus, corium, and apical portion.

Male ganitalia, style pincer-like, with 2 processes about equal in length.
Aedeagus elongate, apical portion curved and narrowed, apex bluntly point-

|
|

VOLUME 84, NUMBER 2 393

|
4h

Figs. 1-11. 1-4, Portanus bilineatus. 5-8, P. spinosus. 9-11, P. caudatus. 1, Head, prono-
tum, and scutellum. 2, 6, 10, Style, laterally. 3, 5, 9, Aedeagus, laterally. 4, 7, 11, Aedeagus,
ventrally. 8, Aedeagus, dorso-apical portion enlarged.

ed. Basal end bearing a rather slender finger-like process which is more than
Ys length of aedeagus and is rounded apically.

Holotype.—¢d, Panama, Chiriqui, Fortuna 1058 m, 8°44'N, 82°15'W, 24-
IX-1977, H. Wolda coll.

Comments.—Portanus caudatus is related to P. eburatus Kramer and
can be separated by the whitish predominant color and the absence of apical
processes.

Xestocephalus variabilis DeLong, NEW SPECIES
Figs. 12-15

Description.—Length of male 5.5 mm, female unknown. Crown produced
and bluntly angled. Crown dull yellow with a brownish circular line extend-

394 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ing from near apex to eye, curving around ocellus. This line joins a circular
line arising near eye, extends to base each side of median 4, and encloses
a brownish spot. Pronotum pale yellow with a dark brown spot behind each
eye enclosing 2 small yellowish spots. Anterior portion at middle and disc
pale brownish. Scutellum yellowish with basal angles dark brown. Forewing
whitish or pale yellow with some veins and numerous spots dark brown.

Male genitalia, style with apical footlike portion with a definite “‘heel”’
which is formed by curvature of blade. Aedeagus V-shaped with the basal
portion of ‘‘V’’ broadened. Ventral portion of shaft elongate, slender, ta-
pered to a pointed curved apex. Dorsal portion of shaft 74 length of ventral
portion and robust.

Types.—Holotype: 6, Vergel, Chiapas, Mexico, 19-V-1935, A. Dampf
coll. Paratypes: 13 ¢, same data as holotype; 14 ¢, Finca Vergel, Chiapas,
Mexico, 16,29-V-1935; 1 6, Mont. Grande, Chiapas, Mexico, 28-III-1931,
A. Dampf coll.

Comments.—Xestocephalus variabilis has been collected only in the
southeastern portion of Mexico, in the state of Chiapas. It is closely related
to X. dedecus DeLong, Wolda, and Estribi from Panama and X. similis
DeLong from Brazil but the basal portion of the aedeagal shaft is thicker
and shorter, and the apical ‘‘foot’’ portion of the style is less angled with
the blade. The aedeagus is the general shape of the aedeagus of the X.
tessellatus—desertorum complex but bears no aedeagal processes.

Xestocephalus dubius DeLong, NEw SPECIES
Figs. 16-18

Description.—Length of male 3.5 mm, female 4 mm. Crown produced
and rounded apically. Crown yellowish with brownish irrorations. Pronotum
yellowish, rather uniformly mottled with brownish areas. Scutellum same
as pronotum with dark brown basal angles. Forewing pale brownish with
darker brown spots and areas. Costa broadly white near middle.

Female posterior margin of 7th sternum broadly, slightly, concavely
rounded.

Male genitalia, style bearing a spine on ventral margin at more than % its
length, apical portion broadened just before pointed apex. Aedeagus broadly
U-shaped, with 2 portions of shaft about equal in length. Dorsal portion
narrowed to form a bluntly pointed apex. Ventral portion of shaft broader,
pointed apically.

Types.—Holotype: ¢, Teguipana, Bolivia, 12-X-1980, Don Foster coll.
Paratypes: 13 ¢, same data as holotype; 5 ¢, Santa Cruz, Bolivia, 1-VI-
1980, D. Foster coll.; 9 5, same except 21-X-1980; 4 6, Portachuela, Bo-
livia, 21-X-1980; | 2, same as holotype except 21-VII-1980.

Comments.—Xestocephalus dubius is related to X. albometus DeLong,

VOLUME 84, NUMBER 2 395

14

13

15
18 :
20
16 21
17 19

Figs. 12-21. 12-15, Xestocephalus variabilis. 16-18, X. dubius. 19-21, X. similis. 12, Head,
pronotum, and scutellum. 13, 16, 21, Style, laterally. 14, 17, 20, Aedeagus, laterally. 15, 18,
19, Aedeagus, ventrally.

Wolda, and Estribi which has a similar style. They can be separated by
comparing the bifid portion of the aedeagus which is longer in albometus.

Xestocephalus similus DeLong, NEw SPECIES
Figs. 19-21

Description.—Length of male 4 mm, female unknown. Head produced
and bluntly pointed. Crown, pronotum, and scutellum dark brownish. Dark-
er brown markings behind each eye and dark brown spots in basal angles
of scutellum. Forewing rather uniformly pale brown with a few darker
brown spots.

Male genitalia, style with apex elongate, footlike, “‘heel’’ scarcely visible.
Apical footlike portion elongate, slender. Aedeagus V-shaped with the 2
portions slightly separated at base. Main shaft almost twice as long as ven-
tral portion, tapered apically and with a slender sharp pointed apex, basal
portion slender.

396 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Holotype.—¢, Viscosa, Amazonas, Brazil, 14-X-1929.

Comments.—Xestocephalus similis is related to X. superbus VanDuzee
and can be separated by the absence of the aedeagal processes and the
distinct heel on the apical footlike portion of the style.

LITERATURE CITED

DeLong, D. M. 1980. New South American xestocephaline leafhoppers (Homoptera: Cica-
dellidae). Entomol. News 91(3): 79-84.

DeLong, D. M. and R. E. Linnavuori. 1978. New tropical Xestocephalus (Homoptera: Cica-
dellidae) and illustrations of little known species. J. Kans. Entomol. Soc. 51(1): 35-41.

DeLong, D. M., H. Wolda, and M. Estribi. 1980. The xestocephaline leafhoppers (Homoptera:
Cicadellidae) known to occur in Panama. Brenesia 17: 251-280.

Kramer, J. P. 1964. A key for Portanus with new records and descriptions of new species
(Homoptera: Cicadellidae: Xestocephalinae). Proc. Entomol. Soc. Wash. 66(1): 5-11.

Linnavuori, R. E. 1959. Revision of the Neotropical Deltocephalinae and some related families
(Homoptera). Ann. Zool. Soc. ‘Wanamo’ 20(1): 1-370 (pp. 35-45).

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 397-401

AXILLARY STRUCTURE OF THE TORTRICIDAE (LEPIDOPTERA)
D. ADAMSKI AND T. M. PETERS

Department of Entomology, University of Massachusetts, Amherst, Mas-
sachusetts 01003.

Abstract.—A fourth axillary sclerite (axillary,,) is reported within the
hindwing base of the Tortricidae. Pivot points of coupled forewings and
hindwings in tortricids are in the same plane, parallel to the median longi-
tudinal body axis. These observations contradict previous interpretations
that imply pivotal points must be contiguous to their corresponding body
segment. Supportive evidence regarding our findings is provided by pho-
tographs of whole mount preparations and a diagrammatic illustration.

The axillary area of Lepidoptera has been studied by several early in-
vestigators (Comstock and Needham, 1898-99; Berlese, 1909; Crampton,
1909, 1920; Snodgrass, 1909, 1935). Recent work by Sharplin (1963a, 1963b,
1964) on lepidopteran wing articulation has provided nomenclatural stabil-
ity, has more clearly defined functional organization, and has traced an
evolutionary series from monotrysian to ditrysian forms.

Wing folding of Lepidoptera, with respect to the axillary sclerites, has
been reviewed by the aforementioned investigators, however, emphasis was
restricted to the forewing. This paper describes a new axillary sclerite in the
base of the hindwing of Tortricidae and discusses the morphological rela-
tionships between the anterior hinge of the forewing and hindwing as to
their function in wing folding.

The present morphological work was conducted as a part of a taxonomic
revision of Nearctic species of the moth genus Apotomis Hubner (Olethreu-
tinae). All tortricid species examined exhibit pivot points of the hindwing
in the same plane as the forewing, parallel to the median longitudinal body
axis. These findings contradict previous interpretations that imply pivotal
points are contiguous to their corresponding body segment. A study of this
discrepancy has led to our discovery of a fourth axillary sclerite within the
hindwing base of Tortricidae.

MATERIALS AND METHODS

Eight species of Tortricidae were examined; these represented two of the
three subfamilies of North American Tortricidae, as recognized by Powell

398 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

DORSAL AXILLARY,

VENTRAL AXILLARY,

PROXIMAL MEDIAN PLATE

TEGULA

HUMERAL PLATE
BASISUBCOSTALE

ANTERIOR NOTAL WING PROCESS

MEDIAN NOTAL WING PROCESS
POSTERIOR NOTAL WING PROCESS

<<.
AXILLARY CORD
PLICA BASALIS

_ erennun
|

BASISUBCOSTALE ___
AXILLARY, .
RADIAL BRIDGE

ANTERIOR NOTAL PROCESS
AXILLARY. \

AXILLARY,
AXILLARY,
MEDIAN NOTAL PROCESS

DISTAL CUBITAL PLATE
PROXIMAL CUBITAL PLATE

?
Y
/.
asad Ce
°
i—

ANAL PLATE

L
~
>
+
N
>

POSTERIOR NOTAL PROCESS

li
w
>

4
W/. YY
iv)
c
v

— MEDIAN PLATE
AXILLARY CORD

2

Figs. 1, 2. Wing bases of Apotomis albeolana. 1, Forewing. 2, Hindwing.

(in press). Species of Tortricidae examined are: Tortricinae: Acleris minuta
(Robinson), Choristoneura fumiferana (Clemens), Archips cerasivoranus
(Fitch), Tortrix viridana (L.); Olethreutinae: Apotomis albeolana (Zeller),
Hedia ochroleucana (Hiibner), Cydia pomonella (L.), Grapholita molesta
(Busck). Specimens of the Chlidanotinae were not available for study.
Specimens were heated in 10% KOH until cleared. Pterothoracic seg-
ments were dissected along the median longitudinal body axis and treated

VOLUME 84, NUMBER 2 399

Table 1. Structural differences between forewing and hindwing bases of Tortricidae.

Structure Forewing Hindwing
Median arm of axillary, present (well developed) absent
Cubital plates absent present
Median plates present (well developed) present (reduced)
Radial bridge absent present
Axillary, complex simple
Axillary, , absent present

with Mallory’s triple stain (Barbosa, 1974). Right half-sections were slide
mounted in balsam. Left half-sections were studied in temporary glycerine
mounts using depression slides. Phase contrast and interference contrast
microscopes were used to examine both sections of the pterothorax. ‘‘True”’
pivot points were determined from glycerine by securing the thorax with
forceps and flexing the forewing and hindwing.

RESULTS AND DISCUSSION

Forewing and hindwing bases of examined specimens are similar to de-
scriptions by earlier workers. Differences between forewing and hindwing
bases are illustrated (Figs. 1, 2) and are summarized (Table 1). Except for
axillary,, (which Sharplin, 1963b, misidentified as the basisubcostale in the
hindwing of Tortrix viridana) our findings agree with Sharplin (1963a,
1963b).

The small axillary,, (Fig. 3) is received by a cuplike invagination of the
anterior apex of axillary,. Movement of axillary,, is restricted by a flat strip
of thin cuticle attached to its base. Axillary,, articulates distally with the
basisubcostale. This juncture marks the ‘“‘true’’ pivot point of the hindwing.

Figure 4 is a diagrammatic representation of the tortricid pterothorax with
the large triangle representing the mesothorax, followed by the smaller
metathorax. According to the workers previously cited, the blackened apex-
es would depict the approximate location of axillary,. In their interpretation
the pivot point axis represented by a broken line is contiguous to the re-
spective body segment. However, our study reveals the observed pivot
point axis to be along the solid line, axillary,, of the hindwing, accounting
for the difference in angle.

We have not conducted an extensive survey throughout the order to de-
termine if the axillary sclerite we describe is present only in the tortricids;
the literature does not indicate its presence in other lepidopteran families.
The structure may have originated along the axis (solid axis of Fig. 4) where
the wing buckled due to frenular wing coupling and the mechanical stress

400 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 3, 4. 3, Hindwing base of Apotomis albeolana; axillary,, indicated by arrow (100%).
4, Diagrammatic representation of the tortricid pterothroax. Large triangle represents meso-
thorax, small triangle represents metathorax. Blackened apexes depict approximate location
of axillary, demarking the pivot point axis represented by a broken line. Observed pivot point
axis is along the solid line. Note that the frenulum is along observed pivot point axis.

at that point. We suggest that this hypothesis could be tested outside the
tortricidae. Further studies are needed to (1) define structural and functional
homologies of axillary,, and (2) determine to what degree, if any, wing
coupling (jugal lobe, frenulum, or expanded costa of the hindwing) is cor-
related with the evolution of axillary,,.

ACKNOWLEDGMENTS

We thank William E. Miller, North Central Forest Experiment Station,
St. Paul, Minnesota; Richard L. Brown, Mississippi State University; and
Ronald W. Hodges, Systematic Entomology Laboratory, USDA, Washing-
ton, D.C., for their constructive advice.

We also express thanks to Chih-Ming Yin, Department of Entomology,
University of Massachusetts, for his encouragement during the preparation
of this paper.

This work was supported by project MS-27 of the Massachusetts Agri-
cultural Experiment Station. Published with the aid of a grant from the
GUY CHESTER CRAMPTON RESEARCH FUND of the University of
Massachusetts.

VOLUME 84, NUMBER 2 40]

LITERATURE CITED

Barbosa, P. 1974. Manual of Basic Techniques in Insect Histology. Autumn Publishers. Am-
herst, Mass. 253 pp.

Berlese, A. 1909. Gli Insetti. Milan. 1004 pp.

Comstock, J. H. and J. G. Needham. 1898-99. The wings of insects. Am. Nat. 32: 43-48,
81-89, 231-257, 413-422, 560-565, 769-777, 903-911; 33: 117-126, 573-582, 845-860.

Crampton, G. C. 1909. A contribution to the comparative morphology of thoracic sclerites of

insects. Proc. Acad. Nat. Sci. Phila. 61: 3-54.

. 1920. A comparison of the external anatomy of the lower Lepidoptera and Trichoptera

from the standpoint of phylogeny. Psyche (Camb., Mass.) 27: 23-34.

Powell, J. A. In press. Tortricoidea. In Hodges, R. W., ed., A Checklist of the Lepidoptera
of America, North of Mexico. London.

Sharplin, J. 1963a. Wing base structure in Lepidoptera. I: Forewing base. Can. Entomol. 95:

1025-1050.

. 1963b. Wing base structure in Lepidoptera. II: Hindwing base. Can. Entomol. 95:

1121-1145.

. 1964. Wing folding in Lepidoptera. Can Entomol. 96: 148-149.

Snodgrass, R. E. 1909. The thorax of insects and articulation of the wings. Proc. U.S. Natl.

Mus. 36: 511-595.

. 1935. Principles of Insect Morphology. McGraw-Hill, New York. 667 pp.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 402-408

NEW SYNONYMIES, LECTOTYPE DESIGNATIONS, AND
OTHER NOTES ON NORTH AMERICAN EPICAUTA
(COLEOPTERA: MELOIDAE)

JOHN D. PINTO

Department of Entomology, University of California, Riverside, Califor-
nia 92521.

Abstract.—Eight nominal species of North American Epicauta are re-
duced to junior synonyms. Epicauta mixta Duges, long considered a junior
synonym of E. neglecta, is given renewed status. The taxonomic position
of E. atricolor Champion and E. unicalcarata Champion is discussed. Both
are herein placed in the subgenus Macrobasis. Also, lectotypes for 11 nom-
inal species are designated.

A revision of the nominate subgenus of Epicauta of North America in-
cluding Mexico and Central America is now in progress. The establishment
of certain synonyms and lectotypes is necessary now, however, to allow
correct name usage in impending biological studies. Brief justifications for
synonymies are presented here. More detailed treatment of the intraspecific
variation relating to these as well as complete species synonymies will fol-
low in the revision. The subgeneric assignment of two species, E. atricolor
Champion and E. uwnicalcarata Champion, also is discussed.

SYNONYMIES
Epicauta brunnea Werner, 1944: 67
Epicauta innomina Dillon, 1952: 401. New Synonymy.

Dillon (1952) differentiated E. innomina from E. brunnea by its lighter
body color, more flattened hindtibial spurs and the single foretibial spur in
the male. Comparison of numerous E. brunnea with the type of E. innomina
indicates that these traits are not significant. The presence of one rather
than two foretibial spurs would appear to be important, but this turns out
to be variable throughout the range of E. brunnea. In fact, the anterior spur
is absent from the foretibia of the holotype of E. innomina but present in
the male paratype from the same locale. In some cases the anterior spur is
absent from one leg but not the other. It is not clear if the missing anterior

VOLUME 84, NUMBER 2 403

spur is always due to breakage. In certain individuals the socket of the
missing spur is visible suggesting breakage. In others, including the holotype
of E. innomina, a distinct socket cannot be seen.

Coloration of E. brunnea varies geographically. Representatives from the
eastern portion of the range (central Texas) are considerably lighter than
those from Arizona. Those from western Texas are somewhat intermediate.

The hindtibial spurs vary in width only slightly in E. brunnea. Those of
the type of E. innomina are well within the range of variation.

Epicauta cinctipennis (Chevrolat, 1834: 59)
Epicauta ruidosana Fall, 1907: 258. New Synonymy.

Fall (1907) apparently was unaware of E. cinctipennis at the time E.
ruidosana was described. Material from Arizona, New Mexico, and Texas
has been historically identified as E. ruidosana (e.g. Dillon, 1952; Werner,
1945), that from Mexico as E. cinctipennis (Werner et al., 1966). However,
no appreciable differences exist between these populations. Some of the
material from Arizona has black rather than white setae on much of the
mid- and hindlegs, but this trait is variable even within Arizona.

Epicauta nigritarsis (LeConte, 1853: 340)
Epicauta hesitata Dillon, 1952: 398. New Synonymy.

Dillon (1952) considered E. hesitata closest to E. crassitarsis. Examina-
tion of the types of E. hesitata shows them to be merely a minor color
variant of E. nigritarsis. The head, pronotum, and elytra are uniformly tan
in most E. nigritarsis. Several specimens throughout its range, however,
have a dark head and pronotum. At an extreme, as in the types of E.
hesitata, the head and anterior half of the pronotum are almost entirely
black. The entire range of variation commonly occurs within the same se-
ries. In fact, a specimen from the same series as the holotype of E. hesitata
(but not designated as a paratype) is of a lighter color.

Epicauta obesa (Chevrolat, 1835: 81)

Epicauta auricomans Champion, 1892: 424. New Synonymy.
Epicauta ficta Werner, 1949: 100. New Synonymy.
Epicauta pseudosolani Dillon, 1952: 395. NEw SyNonyMy.

Epicauta obesa, as now defined, ranges from south central United States
to Oaxaca, Mexico. Several traits vary geographically. Specimens from cen-
tral and southern Mexico are generally less densely setate and have finer
setae than those to the north. Mexican populations have somewhat more
slender hindtibial spurs. The structure of the foretarsus in the male also
varies. The first two segments are elongate, broadened, heavily padded, and

404 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lack cinereous setae on the dorsum. This characterizes all populations but
is better developed in Mexican material. All variation that I am aware of is
clinal and not taxonomically significant.

Epicauta ficta and E. pseudosolani, described from Oklahoma and Texas,
respectively, represent variants typical of the northern portion of the range.
Type-material of E. auricomans, from Coahuila and Oaxaca, have longer
and denser body setation more typical of populations from south central
United States rather than from Mexico.

Epicauta rufipedes (Duges, 1870: 163)
Epicauta insignis Horn, 1885: 110. New Synonymy.

Epicauta rufipedes is one of the most widespread and common species of
Epicauta in Mexico. The species is characterized by the strongly ensiform
antenna and single foretibial spur in the male. Variation in color pattern is
extensive. In the commonest pattern the cuticle of the head and pronotum
is black; the elytra also are black except for a narrow band of brown on the
lateral, medial, and apical margins. These structures are generally covered
with short, relatively sparse, cinereous pubescence which is denser at the
elytral margins and along a narrow line down the center of the elytral disc.

Epicauta insignis, from southern Arizona and northwestern Mexico, ap-
pears to be nothing more than a variant of E. rufipedes. Unlike most E.
rufipedes, specimens conforming to E. insignis have entirely brown elytra,
which contrasts with the black head and pronotum. The elytral vitta is either
absent or obsolescent. This color pattern, however, is not entirely unknown
in central and southern Mexico, and, similarly, some individuals from Ari-
zona have almost entirely black elytra as in typical E. rufipedes. The sim-
ilarity of courtship behavior and first-instar larvae in a population from
southeastern Arizona and one of typical rufipedes from the state of Puebla
further supports synonymy (Pinto, unpublished data).

Epicauta stigmata (Duges, 1870: 159)
Lytta neglecta Haag-Rutenberg, 1880: 54. New SyNonymy.

The type-series of E. neglecta is mixed. The lectotype designated below
for E. neglecta belongs to E. stigmata, a species of Macrobasis. One para-
lectotype belongs to the nominate E. mixta (Duges). See discussion of E.
mixta below.

RENEWED STATUS

Epicauta mixta Duges, 1889: 83

Lytta neglecta: Haag-Rutenberg, 1880: 54 (in part).
Epicauta neglecta: Champion, 1892: 423 (in major part).

VOLUME 84, NUMBER 2 405

Epicauta mixta was treated as a junior synonym of E. neglecta (Haag-
Rutenberg) by Champion (1892). However, of the three syntypes of E. ne-
glecta examined, two belong to Epicauta (Macrobasis) stigmata (Dugés).
Only the third represents the nominate species recognized as E. neglecta
by Champion. Since Haag-Rutenberg’s description of E. neglecta clearly
correlates with the two misidentified types, I have chosen one of these as
lectotype (see below). Epicauta neglecta now is a junior synonym of E.
stigmata, and E. mixta Duges is the valid name for the E. neglecta of
Champion (1892).

The source of confusion in this case stems from the superficially similar
variation in E. stigmata and E. mixta. Pubescence color varies similarly,
ranging from entirely cinereous to entirely black. Both have a form that is
black with intermixed cinereous hairs. The three types of E. neglecta are
of this form. Also, a smooth glabrous callus on either side of the pronotal
midline, a common feature in FE. mixta, is present in some E. stigmata,
including the two misidentified types of E. neglecta.

As indicated, Haag-Rutenberg’s original description of E. neglecta clearly
refers to E. stigmata. He states that the first antennal segment of the male
is thick, three times as long as broad, and that the second segment is half
as long as the third. Both traits characterize Macrobasis. He also states
that the pronotum is somewhat longer or as long as broad. Again, this agrees
with E. stigmata, but not E. mixta where the pronotum is noticeably broader
than long.

Duges’ (1889) description of E. mixta was based on material from Oaxaca.
It is not clear how many specimens were before him, but the lack of a range
in his length measurement suggests only one. I have examined two speci-
mens in what remains of the Duges Collection at the Universidad Nacional
Autonoma de Mexico (UNAM) labeled as types of FE. mixta. One is labeled
‘‘Oaxaca,”’ the other, ‘“‘Moro Leon.’’ Both fit Duges’ rather generalized
description. However, the latter is almost certainly not a type since Mo-
roleon is a Guanajuato locality.

TRANSFERS TO MACROBASIS
Epicauta atricolor Champion, 1892: 419

Epicauta atricolor was removed from Macrobasis (as Gnathospasta) and
placed in the nominate subgenus by Werner (1958). Its transfer was justified
primarily by the absence of the hindtibial comb in one of the types. This
trait is characteristic of most Macrobasis but is absent in some species
(Selander and Mathieu, 1969). Examination of the type-series convinces me
that E. atricolor should be reassigned to Macrobasis. The male lectotype
(designated below) actually does have a poorly developed comb of three

406 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

teeth. Also, the first antennal segment of the male is slightly enlarged and
curved. The antennal dimorphism is not as marked as in most Macrobasis
but is no less developed than in certain species of the subgenus such as E.
evanescens Champion. Epicauta atricolor should tentatively be placed in
the Uniforma Group as originally indicated by Werner (1954).

Epicauta unicalcarata Champion, 1892: 412

Epicauta unicalcarata has never been formally assigned to subgenus,
but its omission from Werner’s (1954) list of species in Macrobasis (as
Gnathospasta) and from other discussions of the limits of that subgenus
(Werner, 1958; Selander and Mathieu, 1969) suggest its placement in the
nominate subgenus. Indeed, Champion made no mention of antennal modi-
fications typical of Macrobasis and considered this species closest to E.
horni Champion, a nominate species.

I have examined the male holotype of E. unicalcarata in the British Mu-
seum (from Amula, Guerrero). It is badly damaged with the head and prono-
tum entirely missing. The hindlegs, however, are intact and the presence of
a well-developed hindtibial comb indicates that the species belongs to Mac-
robasis.

I am not aware of additional material of this species. Features not men-
tioned by Champion that should help identify it are a small patch of brown
pubescence on the apex of each elytron, a distinctly concave metasternum,
flattened hindtrochanters, and the glabrous, subconcave ventral surface of
the hindfemora.

It is not possible to assign E. unicalcarata to species group at this time.

LECTOTYPE DESIGNATIONS

Syntypes of several North American species described by Champion
(1892), Haag-Rutenberg (1880), and Chevrolat (1834) were examined, and
lectotypes are designated below. Syntypes were primarily identified by the
correspondence of specimens and attached labels with original descriptions.
All lectotypes are labeled as such and deposited in their appropriate mu-
seums. Additional syntypes are labeled paralectotypes.

Specific names are listed below in alphabetical order. The original genus
follows the specific name in parentheses. Locales and other data are given
as they appear on labels. Data from individual labels are delimited by a
slash (/).

Abbreviations for museums are as follows: BMNH (British Museum of
Natural History, London); HNHM (Hungarian Natural History Museum,
Budapest); UZMH (Universitets Zoologiska Museum, Helsinki); ZSM
(Zoologische Staatssammlung, Munich).

VOLUME 84, NUMBER 2 407

atricolor Champion (Epicauta). Lectotype: ¢, ‘‘Oaxaca, Mexico/Salle
Coll./’?; (BMNH). Paralectotypes: 5 (BMNH).

auricomans Champion (Epicauta). Lectotype: ¢, ‘‘Oaxaca, Mexico/
Hoege/’’; (BMNH). Paralectotypes: 4d, 22 (BMNH). The specimen cho-
sen as lectotype was figured by Champion (1892: Table 19, Fig. 24). This
individual is abnormal in that the hindtibiae are abruptly and strongly
bowed.

basimacula Haag-Rutenberg (Lytta). Lectotype: ¢, ‘*Mexico/Samml. Haag-
Rutenberg/’’; (ZSM). Paralectotypes: 12 (ZSM), 1¢ (UZMH).

carmelita Haag-Rutenberg (Lytta). Lectotype: 6, ‘‘N. Grenada/F. Bates/
typ. Haag/’’; (BMNH). Paralectotypes: 16 (HNHM), 32 (ZSM), 12
(UZMH).

cinctipennis Chevrolat (Lytta). Lectotype: 6, ‘“‘Mexico, Mineral del Zi-
mapan/Sommer/’’; (UZMH). Paralectotype: 12 (UZMH).

curvicornis Haag-Rutenberg (Lytta). Lectotype: ¢, ‘‘Mexico/F. Bates/ typ.
Haag./’’; (BMNH). Paralectotype: 12 (BMNH). I have examined 13 ad-
ditional specimens from UZMH which are questionably from the original
series. At least 4 of these represent species other than curvicornis.

emarginata Champion (Epicauta). Lectotype: ¢, ‘San Isidro, Mexico,
Hoge/’’; (BMNH). Paralectotypes: 32 (BMNH); 12° (HNHM).

leucocoma Champion (Epicauta). Lectotype: 2, ‘‘Tepanistalabuaca, Mex-
ico, Salle Coll./’’; (BMNH). Paralectotypes: 22 (BMNH). The lectotype
was figured by Champion (1892: Table 19, Fig. 23); it is labeled a male in
error).

neglecta Haag-Rutenberg (Lyfta). Lectotype: 9, ‘“‘Mex./F. Bates/Typ.
Haag/’’; (BMNH). Paralectotypes: |1¢ (HNHM); 12 (ZSM). As indicated,
the original series of E. neglecta is mixed. The lectotype and male para-
lectotype are assignable to E. stigmata. The female paralectotype rep-
resents E. mixta.

singularis Champion (Epicauta). Lectotype: d, “‘Monterey, Nuevo Leon,
Hoge/’’; (BMNH). Paralectotypes: 46, 12 (BMNH); 1d (HNHM).

subvittata Haag-Rutenberg (Lytta). Lectotype: 6, ‘“‘Mexiko/Coll. Blny./”’;
(UZMH). Paralectotypes: 1¢ (UZMH); 22 (BMNH). Five other speci-
mens in UZMH also may represent part of the original series. Epicauta
subvittata has long been considered a junior synonym of E. rufipedes
(Champion, 1892).

ACKNOWLEDGMENTS

I thank the following individuals for the loan of type material: M. H. D.
Brendell (BMNH); H. R. Burke (Texas A&M University, College Station);
Z. Kaszab (HNHM); G. Scherer (ZSM); H. Silfverberg (UZMH). The re-

408 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

visionary studies of Epicauta are being supported by grant DEB-7915307
from the National Science Foundation.

LITERATURE CITED

Champion, G. C. 1891-93. Family Meloidae. Jn Godman, F. D. and O. Salvin, Biologia Cen-
trali-Americana, Coleoptera, 4(2): 364-450, 462-464.

Chevrolat, L. A. 1834-35. Coléopteres du Mexique, Fasc. 3, 4. Strasbourg.

Dillon, L. 1952. The Meloidae (Coleoptera) of Texas. Am. Midl. Nat. 48: 330-420.

Duges, E. 1870. Descripcion de algunos Meloideos indigenas. Naturaleza |: 125-128, 157-171.

. 1889. Sinopsis de los Meloideos de la Republica mexicana. An. Mus. Michoacano 2:

34-40, 49-114.

Fall, H. C. 1907. Descriptions of new species, pp. 218-272. In Fall, H. C. and T. D. A.
Cockerell, The Coleoptera of New Mexico. Trans. Am. Entomol. Soc. 33: 145-272.

Haag-Rutenberg, J.G. 1880. Beitrage zur Kenntniss der Canthariden. Dtsch. Entomol. Z. 24:
17-90.

Horn, G. H. 1885. Studies among The Meloidae. Trans. Am. Entomol. Soc. 12: 107-116.

LeConte, J. L. 1853. Synopsis of The Meloides of the United States. Proc. Acad. Nat. Sci.
Philad. 6: 328-350.

Selander, R. B. and J. M. Mathieu. 1969. Ecology, behavior and adult anatomy of The Albida
Group of the genus Epicauta (Coleoptera: Meloidae). Ill. Biol. Monogr. 41: 1-168.

Werner, F. G. 1944. Some new North American species of Epicauta (Coleoptera: Meloidae).

Psyche (Camb. Mass.) 50(1943): 65-73.

. 1945. A revision of the genus Epicauta in America north of Mexico. Bull. Mus. Comp.

Zool. 95: 421-517.

. 1949. Additions to Epicauta, with new synonymy and a change of names (Coleoptera:

Meloidae). Psyche (Camb. Mass.) 56: 93-111.

—. 1954. A review of the subgenus Gnathospasta of the genus Epicauta (Meloidae).
Coleopt. Bull. 8: 25-27.

—. 1958. Studies in the genus Epicauta of the North American Continent (Meloidae) II.
The Uniforma-Group. Coleopt. Bull. 12: 1-19.

Werner, F. G., W. R. Enns, and F. H. Parker. 1966. The Meloidae of Arizona. Univ. Ariz.
Agric. Exp. Stn. Tech. Bull. 175, 96 pp.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 409-410

NoTE

Silphids Attracted to Mammal Carrion at Cheltenham,
Maryland (Coleoptera: Silphidae)

Several field experiments dealing with carrion beetles (Silphidae) were
conducted from 2 June 1981 to 3 August 1981 at Cheltenham, Prince Georges
Co., Maryland. The work was done on the reservation where the Naval
Communication Unit is located. The reservation’s 220 ha contain a variety
of natural habitats such as marsh, old field, mature deciduous woodland,
mature coniferous woodland, and young mixed woodland. The most exten-
sive habitat is the latter.

Four traps (4.21 cans covered by 1.27 cm mesh hardware cloth and a rain
cover 5 cm above the can opening) were suspended 45 cm above the ground.
Each of these cans was suspended from a crosspiece between 2 stakes in the
young woodland habitat. Two similar traps were situated in each of the
following habitats: Mature coniferous woodland, marsh, and old field. No
trap was closer than 75 m to a neighboring trap. Each trap was baited with
a whole fetal pig weighing about 380 g and the carrion was left to decompose
for 7 days. Traps were usually examined on days 1, 2, 3, 4, and 7 after
having been set. Carrion beetles were removed, identified, and the data
recorded for each trap.

A total of 300 trap collections was made during the study and a total of
1873 carrion beetles, representing 8 species, was taken. The species, the
number of each species collected, the percentage of all silphids this repre-
sents, and rank order for each species are in Table 1.

It is clear from this data that Silpha americana is the dominant silphid
species collected since it accounted for well over half of all Silphidae taken.

Table 1.
_ ee eee eee ee ee
Number % of
Species Collected Total Rank
Silpha americana L. 1027 54.83 l
S. noveboracensis Forster 318 16.98 2
S. inaequalis F. 253 ISvou 3
Nicrophorus tomentosus Weber 139 7.42 4
N. orbicollis Say 109 5.82 5
Necrodes surinamensis F. 24 1.28 6
Nicrophorus pustulatus Herschel 2 0.11 4
N. marginatus F. l 0.05 8

Total 1873 100.00

ee ——————————————————n

410 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Silpha noveboracensis is a very poor second. In previous studies in New
Jersey, Shubeck has found that Silpha noveboracensis is clearly the domi-
nant species through June and July, and S$. americana is second or third in
abundance (Shubeck et al. 1977. The Wm. L. Hutcheson Memorial Forest
Bull. 4(1): 12-17; Shubeck et al. 1981. Entomol. News 92(1): 7-16).

The 8 species of Silphidae collected in Maryland during this study are the
same 8 silphid species that have been taken by Shubeck during the last 21
years of carrion beetle studies in New Jersey. Continued field studies may
reveal that additional ‘‘common’”’ silphid species (according to previous lit-
erature), are, in fact, no longer common in New Jersey and Maryland.

P. P. Shubeck and D. L. Blank, Code 4357, Naval Research Laboratory,
Washington, D.C. 20375. P. P. Shubeck, permanent address: Biology De-
partment, Montclair State College, Upper Montclair, New Jersey 07043.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 410-411

NOTE

The Habits and Appearance of a Rare Mealybug, Eurycoccus blanchardii
(King and Cockerell) (Homoptera: Coccoidea: Pseudococcidae)

Between 1897 when Eurycoccus blanchardii (King and Cockerell) was
described and the present time, there have been only two recorded collec-
tions of this species and only a brief mention of a host or the feeding habits
and appearance of the insect in life. It therefore seems desirable to record
its rediscovery in what may have been an unusual location.

On 23 June 1968 while digging up daffodil bulbs in Silver Spring, Mary-
land, I found a husked hickory nut (Carya sp.) approximately 6” below the
surface of the soil. The nut was cracked open about !/16”, and it had a root
sprout about 6” long that extended deeper into the soil. There were 20
mealybugs inside the nut next to the shell, a most unusual location for
mealybugs as far as known, and there were also 2 mealybugs on the root
about 2” below the nut. Some of the mealybugs inside the nut appeared to
be feeding on the sprouting kernel. The mealybugs were attended by ants
which escaped before I could get a container to place them in. Although I
have dug in the same plot annually since 1968, I have not found the mealy-
bug again.

The mealybugs were a dull, but almost glistening white owing to the waxy
pulverulence that covered them. Underneath the waxy secretion the bodies
were pinkish red instead of reddish purple as described by King and Cock-
erell for the original example of blanchardii.

The specimens were identified by the late H. L. McKenzie of the Uni-
versity of California at Davis.

VOLUME 84, NUMBER 2 411

An annotated list of all previous articles known to me that have mentioned
blanchardii is given as an aid to persons who may encounter this mealybug.

King and Cockerell. 1897. Can. Entomol. 29: 92-93. Ripersia blanchardii
described from ‘‘Haverhill, Mass., October 4th, 1896, in a nest of Lasius
claviger, Rog., under a stone with a small herd of another species; only
one found, not feeding.”

Tinsley. 1899. Can. Entomol. 31: 46. Noted that the type-specimen had
malformed antennae.

King. 1899. Can. Entomol. 31: 111. Stated *‘Large and not often met with

Lindinger. 1908 (1907). Berl. Entomol. Z. 52: 90. Changed the spelling to
blanchardi.

MacGillivray. 1921. The Coccidae, p. 141. Listed as a species described in
Ripersia.

Britton. 1923. Conn. State Geol. and Nat. Hist. Surv. Bull. 34: 382. Indi-
cated that the species might be found in Connecticut because it was
known in Massachusetts.

Trimble. 1928. Entomol. News 39: 44. Recorded the species ‘‘on grass roots
in ants’ nest’’ in Pennsylvania. Through the courtesy of G. B. Sleesman,
formerly of the Department of Agriculture, Harrisburg, Pennsylvania, I
have been able to examine specimens that may be the ones Trimble listed.
The data with these specimens are ‘‘Rip. blanchardii?, F 108, under a
stone attended by ants, Rockville, Pa., 4-16-22, A. B. Champlain.’’ Un-
fortunately the specimens are in such poor condition that I cannot deter-
mine whether they are blanchardii. Other Trimble specimens labeled
‘‘Rip. minima T & K? on grass roots, in ant nest, Rockville, Pa., 2-15-21,
coll. F. M. Trimble’’ are also very poor mounts, but these insects definitely
are not blanchardii. These two lots are the only ones found in the Trimble
collection that might be the one he recorded.

Ferris. 1953. Atlas of the Scale Insects of North America 6: 349-350. Ferris
redescribed and illustrated blanchardii and transferred it to Eurycoccus
Ferris 1950 though stating that it was not typical of the genus.

Williams. 1958. Proc. R. Entomol. Soc. Lond., Ser. B., Taxonomy 27(1-2):
22-24. Mentioned similarity of his new species to blanchardii.

Sailer. 1968. USDA, Coop. Econ. Insect Rep. 18(36): 866. Recorded my
collection as new for Maryland.

Balachowsky and Ferrero. 1969. Bull. Inst. Fondam. Afr. Noire, Ser. A,
Sci. Nat. 31: 138. Listed blanchardii as a species of Eurycoccus.

Brooks. 1972. J. Aust. Entomol. Soc. 11: 129. Listed blanchardii as a
species of Eurycoccus.

Louise M. Russell, Cooperating Scientist, Systematic Entomology Lab-
oratory, IIBIII, Agricultural Research Service, USDA, Beltsville Agricul-
tural Research Center, Beltsville, Maryland 20705.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, p. 412

Book REVIEW

The Plant Protection Discipline, Problems and Possible Developmental
Strategies. By W. H. Sill, Jr. Allanheld, Osmun and Co.; distr. John
Wiley and Sons, New York. 190 pp. 1979. Cost: $25.00.

This hardcover book provides an interesting, wide-ranging discussion of
the practical applications and philosophical foundations of plant protection.
Topics include world food losses due to pests; complexity of agro-
ecosystems; interdisciplinary research and coordination needed; integrated
pest management; comparison of plant protection in developed and devel-
oping nations; economics of crop production; viewpoints of administrators
and research scientists; public relations problems; low prestige of agricul-
ture as a profession; training of extension agents and other crop protection
and crop production experts; academic training; possible organizational
strategies; legislation; and regulation of pesticides with certification of ap-
plicators, resembling the regulation of pharmaceuticals and pharmacists.
Useful and explicit definitions of terms commonly in the literature are pro-
vided, the concepts and nomenclature defining plant protection are dis-
cussed, and numerous authorities are extensively quoted. The daunting
complexity of understanding and managing the many interacting and chang-
ing components that comprise agroecosystems are correctly stressed. The
author emphasizes that, contrary to the viewpoint of many city dwellers,
agriculture cannot be performed by the uneducated peasants, but instead is
rapidly evolving into what must be the most complex and challenging of all
human endeavors, if world food production is to be adequate.

Although this book is addressed primarily to those concerned with plant
protection, there is much of interest regarding crop production, general
ecology, and sociology as well. It is well written, thoughtful, interesting to
read, and should be available to agricultural planners, research scientists,
and extension workers.

S. W. T. Batra, Research Entomologist, Beneficial Insect Introduction
Laboratory, IBIII, Agricultural Research Service, USDA, Beltsville, Mary-
land 20705.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, p. 413

Book REVIEW

Systematics of the Colletidae Based on Mature Larvae with Phenetic Anal-
ysis of Apoid Larvae (Hymenoptera: Apoidea). By R. J. McGinley. Uni-
versity of California Publications in Entomology, Vol. 91, 307 pp. 1981.
Cost: $14.50.

The colletid bees are generally regarded to be the ancestors of the bees
(Apoidea), based on the morphological similarity of adults to certain sphe-
coid wasps and the relict distribution of colletids, with speciation principally
in South America and Australia. As there are over 20,000 species of bees
worldwide, on which most higher plants depend for pollination (including
about 20 billion dollars’ worth of crops), the origin and evolution of the bees
are of considerable basic scientific and agricultural significance.

This volume takes a novel and difficult approach to assess the evolution-
ary relationships among colletid subfamilies and genera, and their possible
relationships to other bee families, because it is based on the relatively scant
and indistinct morphological characters of the larvae. Numerical taxonomy
was used to generate a cladistic analysis, and larval information did not
support colletid monophyly. This conclusion is supported by the recent
findings of this reviewer and colleagues (unpublished) that the secreted nest
membrane in Colletidae consists of two chemically unrelated types; in Col-
letinae, it is a polyester derived from lactones in the Dufour’s gland (similar
to Halictidae), and, in Hylaeinae, it is a nest membrane composed of silk
secreted by salivary glands (similar to silk of pemphredonine wasps). The
bifid or blunt glossa of Colletidae may thus be only the result of convergent
evolution in at least two groups that use the glossa for applying a secreted
nest membrane, and therefore should not be considered as the definitive
family characteristic.

This well written and amply illustrated paperback publication includes
descriptions of, and a key to, known mature colletid larvae; a phenogram;
partial taxometric maps; similarity matrix; and ranked nearest-neighbor list-
ing. A phenogram and nearest-neighbor diagram for 96 apoid taxa indicates
11 major phenetic groupings based on larvae, not all of them corresponding
to existing taxonomic groups based on external morphology of the adults.
I recommend this book to institutional libraries, hymenopterists, numerical
taxonomists, and those interested in the evolution of the bees.

S. W. T. Batra, Research Entomologist, Beneficial Insect Introduction
Laboratory, IIBIII, Agricultural Research Service, USDA, Beltsville, Mary-
land 20705.

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, p. 414
SUMMARY REPORTS OF SOCIETY OFFICERS FOR 1981

TREASURER
(1 November 1980 to 31 October 1981)

Special

General Publication
Summary Fund Fund Totals
On hand, | November 1980 21,462.57 33,133.73 54,596.30
Total receipts 38,434.76 5,857.00 44,291.76
Total disbursements 42,381.66 8,145.05 50,526.71
On hand, 31 October 1981 17,515.67 30,845.68 48 361.35

EDITOR
(Calendar Year 1981)

A total of 139 manuscripts were received and processed from November
1, 1980 to October 31, 1981. Four numbers of the Proceedings were pub-
lished in 1981. The 824 pages represented 81 scientific articles, 13 scientific
notes, 8 book reviews, 2 obituaries, 5 announcements, and minutes for 12
Society meetings. Reports for officers for 1980, information for contributors,
table of contents for volume 83, index to new taxa for volume 83, and PS
Form 3526 were also published. This is the largest volume published by the
Society.

Editorial charges were waived for 8 articles totaling 54 pages. Full edi-
torial charges were paid for immediate publication of 6 articles totaling 65
pages and one article of 4 pages by non-members. Ten lengthy articles
included full editorial charges for 58 pages.

In September, the Society published Memoir Number 9, The Flower Flies
of the West Indies by F. Christian Thompson. The 200 page Memoir is
available from the Society at a cost of $10.00.

The Society covered expenses for the Editor to attend the Editorial Work-
shop held at Allen Press, Inc., Lawrence, Kansas, October 15-16, 1981.
Representatives from 20 scientific journals attended. The session included
a tour of the Allen Press plant and seminars on costs of production, scientific
illustrations, style of publications, new developments from the U.S. Postal
Service, marketing of scholarly publications, and the librarian’s view of
scholarly journals.

Publications Committee: E. Eric Grissell, John M. Kingsolver, Wayne N.
Mathis, George C. Steyskal, Thomas E. Wallenmaier, and David R. Smith
(Editor).

PROC. ENTOMOL. SOC. WASH.
84(2), 1982, pp. 415-420

SOcIETY MEETINGS
880th Regular Meeting—October 1, 1981

The 880th Regular Meeting of the Entomological Society of Washington
was called to order by President Jack E. Lipes at 8:00 PM, October 1, 1981,
in the Naturalist Center, National Museum of Natural History. Twenty-one
members and two guests attended.

Minutes of both the 878th and 879th meetings were read and approved.

Membership Chairman Joyce Utmar read the names of fifteen new mem-
bers:

Nancy J. Agafitei, St. Charles, Illinois.

Wesley Bicha, Logansport, Indiana.

D. Christopher Darling, Department of Entomology, Cornell University,
Ithaca, New York.

Andrew G. Gerberich, Arlington, Virginia.

Lester Paul Gibson, Plain City, Ohio.

Roger H. Grothaus, Columbia, Maryland.

Akey C. F. Hung, Insect Identification and Beneficial Insect Introduction
Institute, USDA, Beltsville, Maryland

Adriean J. Mayor, Riverside, California.

Gary L. Miller, Department of Entomology and Plant Pathology, Uni-
versity of Tennessee, Knoxville, Tennessee.

D. Muraleedharan, Department of Zoology, University of Kerala, Kerala,
India.

Terry Prewitt Nuhn, Arlington, Virginia.

Jim Pakaluk, Department of Entomology, Cornell University, Ithaca,
New York.

Sharon Rose, Department of Biology, Montana State University, Boze-
man, Montana.

Akihiko Shinohara, College of Agriculture, University of Osaka Prefec-
ture, Sakai, Osaka, Japan.

M. A. Tidwell, Department of Entomology, The Citadel, Charleston,
South Carolina.

President Elect Margaret Collins announced that there was an omission
of acknowledgment of the contribution of the consulting company of Mr.
John W. Kennedy on the program of the annual banquet, and she stated
that a letter of apology for the oversight had been sent to him. President
Lipes announced the formation of an Auditing Committee and a Nominating
Committee to propose names as candidates for Society officers for 1982.

416 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The speaker for the evening was Dr. Jeffrey Aldrich of the Insect Phys-
iology Laboratory, USDA, in Beltsville, who presented a talk with slides,
entitled ‘‘Chemical Ecology of the Hemiptera: Exocrinology and Endocri-
nology.’’ His talk discussed the phylogeny of the Hemiptera in the light of
the chemical structures of allomones and pheromones and described some
of the research being conducted to evaluate the behavioral roles of these
chemicals.

NOTES AND EXHIBITIONS

Editor David R. Smith displayed the latest issue of the Society’s Memoir
series, Flower Flies of the West Indies by F. Christian Thompson.

President Lipes passed around correspondence of historical interest to
the Society, consisting of letters between the Department of Entomology,
The University of Connecticut, and Dr. E. A. Back regarding the Back
Collection.

Special acknowledgment was given to Dr. Curtis Sabrosky, recently re-
tired from the Systematic Entomology Laboratory, USDA, as recipient of
the L. O. Howard Award at the Eastern Branch meetings of the Entomo-
logical Society of America.

President Lipes reported on the deaths of Dr. Bennet Porter, 89, former
head of the Bureau of Entomology and Plant Quarantine at Beltsville, and
Dr. Lowd E. Goodhue, inventor of the insecticide aerosol.

The meeting was adjourned at 9:25 PM for cookies and cider prepared by
Mrs. Helen Sollers-Riedel.

David A. Nickle, Recording Secretary

88Ist Regular Meeting—November 5, 1981

The 88st Regular Meeting of the Entomological Society of Washington
was Called to order by President Jack E. Lipes at 8:00 PM on November 5,
1981 in the Naturalist Center of the National Museum of Natural History.
Thirty-three members and seven guests attended.

Minutes of the previous meeting were read and approved.

Membership Chairman Joyce Utmar read the names of three new mem-
bers:

William C. Scharf, Northwestern Michigan College, Division of Science
and Mathematics, Traverse City, Michigan.

Kenneth W. Stewart, Department of Biological Sciences, North Texas
State University, Denton, Texas.

Robert Wharton, Department of Entomology, Texas A&M University,
College Station, Texas.

VOLUME 84, NUMBER 2 417

President Lipes announced the nominations of the following individuals
for officers of the coming year. These names were submitted for consider-
ation by the Nominating Committee, consisting of Drs. Raymond Gagné,
Neal Morgan, and Ivan Rainwater.

President-elect, Dr. Manya B. Stoetzel
Recording Secretary, Dr. Thomas E. Wallenmaier
Corresponding Secretary, Dr. Ralph E. Harbach

Treasurer, Dr. F. Christian Thompson

Editor, Dr. David R. Smith
Membership Chairman, Ms. Joyce A. Utmar
Program Chairman, Dr. Jay C. Shaffer
Custodian, Dr. John F. Carroll

Hospitality Chairman, Mrs. Helen Sollers-Riedel

President Lipes opened the floor for other names to be submitted, but no
entries were made.

The speaker for the evening was Dr. Ron Goor, National Heart, Lung
and Blood Institute, NIH, who presented a colorful slide talk entitled
‘Shooting Insects in the Wild.’ The talk focused on insect photography,
and suggestions were presented for successful close-up photographs and
slides of insects.

NOTES AND EXHIBITIONS

Editor David R. Smith passed around the Entomological Society of Wash-
ington Memoir No. 9, Flower Flies of the West Indies, by F. Christian
Thompson. He also described his trip to Allen Press, Inc., Lawrence, Kan-
sas, to attend an editor’s workshop presented by Allen Press, the printer of
the Society’s publications.

Mr. Ed Saugstad presented a book published in 1851, Episodes of Insect
Life by Acheta domestica, M. E. S., dedicated to William Kirby, William
Spence, and Edward Forbes. It consists of essays on the habits and life
cycles of common insects.

Dr. J. H. Fales passed around the program by Sigma Xi of Towson State
University of the Symposium of Threatened and Endangered Plants and
Animals of Maryland.

Dr. Dee Wilder showed slides of dolichopodid flies collected in California.
She was especially interested in a fly, Tachytrechus flabellifer Osten Sack-
en, in which males have a conspicuous silvery band extending from an
elongated clypeus across the thorax. Females lack this band, and it is pre-
sumed to attract females as part of a pair formation strategy.

Dr. Manya Stoetzel showed slides of northern Arizona and the retired
Systematic Entomology Laboratory staff members Dr. Barney Burks and
Mrs. Kellie O’ Neill Burks.

418 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Dr. Ashley B. Gurney mentioned that Dr. Frederick Poos celebrated his
90th birthday and that Mr. Oscar Cartwright was in the hospital with heart
problems.

Visitors were introduced, and the meeting was adjourned at 9:35 PM for
refreshments.

David A. Nickle, Recording Secretary
882nd Regular Meeting—December 10, 1981

The 882nd Regular Meeting of the Entomological Society of Washington
was called to order by President Jack E. Lipes at 8:00 PM on December 10,
1981 in the Naturalist Center, National Museum of Natural History. Thirty-
four members and 11 guests attended.

Minutes of the previous meeting were read and approved.

Membership Chairman Joyce Utmar read the names of the following new
members:

Bozidar Curcic, Institute of Zoology, Faculty of Science, Studentski trg
16, 11000 Beograd, Yugoslavia.

Steve Marshall, Department of Environmental Biology, University of
Guelph, Guelph, Ontario.

John, W. Neal, Jr., USDA, Beltsville Agricultural Research Center,
Beltsville, Maryland.

Paul P. Shubeck, Department of Biology, Montclair State College, Upper
Montclair, New Jersey.

Donald W. Webb, Illinois Natural History Survey, 607 E. Peabody,
Champaign, Illinois.

Corresponding Secretary Mignon Davis reported that the current mem-
bership of the Society was 532 with subscriptions totalling 330.

A motion by Dr. Raymond Gagné to vote in toto for the nominees for
offices in 1982 was passed. The attending members voted unanimously in
favor of electing all nominees.

Treasurer F. C. Thompson presented a report of the financial condition
of the Society. He pointed out that although the assets as of October 31,
1981 were $17,515.67 (General Fund) and $30,845.68 (Special Publications
Fund), both Funds are down from last year. He recommended a membership
dues increase of $5.00 (from $8 to $13) and a change in Article 4, Section
| of the Constitution to read ‘‘thirteen’’ instead of ‘‘eight.”’

Editor Smith reported that 104 articles totalling 824 pages had been pub-
lished in the Proceedings in 1981, making this volume the largest in the
Society’s history.

The speakers for the evening were Drs. Lloyd Knutson and Robert D.

VOLUME 84, NUMBER 2

Fig. |. Zebra butterfly, the correct resting position.

420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Gordon, IIBII, USDA, who together presented a talk entitled, ‘“‘Entomol-
ogy in China—from Taxonomic and Biological Control Viewpoints with
Comments on Opportunities for and Difficulties in Cooperation.’ Dr. Knut-
son outlined the recent history of the broadening cooperative research ac-
tivities between the United States and the People’s Republic of China in
systematics and biological control. Dr. Gordon presented slides of the recent
trip to China which he and Dr. Knutson had made to explore research
potentials in China.

NOTES AND EXHIBITIONS

Dr. Ed Barrows passed around a box of instant rice containing a dead
specimen of the Southern Brown Cockroach, Periplaneta brunnea Bur-
meister, which had been packaged with the rice.

To inspire the spirit of Christmas through a presentation of the traditional
colors of red and green, Dr. David Nickle displayed a living bright pink
katydid, Amblycorypha floridana Rehn and Hebard, on a branch of Amer-
ican holly.

Mrs. Mignon Davis circulated the October issue of The Naturalist (for-
merly The Trinidad Naturalist) with a cover photograph of several speci-
mens of Heliconius charitonius simulator Roeber. Commonly called the
zebra butterfly, it is the only species with parallel black and white stripes.
The same individuals return nightly to the same branch of certain trees for
several weeks. Zebra butterflies do not rest standing upright but hang from
the branch (Fig. 1). The cover photograph was printed upside down. Mrs.
Davis discovered and assisted in photographing these butterflies just after
dusk on a road in a Jamaican plantation near Runaway Bay.

Dr. Thompson showed volume 2 of Myia, Charles Howard Curran, a
catalog of the entomological taxa of Curran and his bibliography. Later, Dr.
Thompson announced the death of Dr. Charles P. Alexander, dipterist and
author of more than 1000 papers, primarily on crane flies of the world. His
announcement was accompanied by slides of Dr. Alexander at his home in
Amherst, Massachusetts.

President Lipes presented letters of appreciation to officers M. Davis, S.
Nakahara, and D. Nickle, and to Mrs. Helen Sollers-Riedel and Dr. Donald
Anderson for their special services to the Society. He then read a letter he
had written to the Selection Committee for Outstanding Scientists, ARPE,
in support of the Award in Category 6 (Systematics and Morphology) for
Dr. Richard H. Foote, Systematic Entomology Laboratory, USDA.

Visitors were introduced, and the meeting was adjourned at 9:35 PM for
cookies and Holiday punch.

David A. Nickle, Recording Secretary

PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

MISCELLANEOUS PUBLICATIONS

Cynipid Galls of the Eastern United States, by Lewis H. Weld

Cynipid Galls of the Southwest, by Lewis H. Weld

BMrEpaperssOmGYMplyPalls} 2... e. 5 see enna ose

Identification of Aiaskan Black Fly Larvae, by Kathryn M. Sommerman

Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides ,
TVET AVENEL AVC ers cent I 2 eee ey, Si os oe et eee ae Be

A Short History of the Entomological Society of Washington, by Ashley B.
(CHITIE)®, a8 Sod 2 ei et re a io ai ae ae oe MOORE OS Tad

Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by
OTHE Ge SHES EN oes Me RE oe Bee Re oP ee ee ee ee =

Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephriti-
MAG OVETCOLRS Casey Shale cee 22.226 42st ee

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse.
TR ae ee ee cs See, Ee ae nn Sr

No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by
PACs ERVIN Sy ML O42 cs ie EL een ee ne SE ae ee

No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by
PAPA NV SOTO ATI on 1940. ete 2 a5 oe ee eee he

No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller.

No. 5. A Classification of the Siphonaptera of South America, by Phyllis T.
Trey parr ihe Me Gi Pk ee ee RR eee ee ee ee ree SS ee Set Ss

No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P.
: Murdoch and Hirosi Takahasi. 1969 .......--__-_--_---------------

No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette
Mates Felt 7 ( Se ne ae See eee errors See ae a ote

No. 8. The North American Predaceous Midges of the Genus Palpomyia Mei-
gen (Diptera: Ceratopogonidae), by W. L. Grogan, Jr. and W. W.

No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F.
musta baonipson. 1961 o.oo seks canker

$15.00
(out of
print)
15.00

15.00

15.00

Back issues of the Proceedings of the Entomological Society of Washington are available
at $18.00 per volume to non-members and $9 per volume to members of the Society.

Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are
allowed a discount of 10 per cent on all items, including annual subscriptions, that are
paid in advance. All orders should be placed with the Custodian, Entomological Society
of Washington, % Department of Entomology, Smithsonian Institution, Washington,

D.C. 20560.

CONTENTS
(Continued from front cover)
KONDRATIEFF, B. C. and R. E. KIRCHNER—Notes on the winter stonefly genus
Allocapnia (Plecoptera: Capniidae)
O'BRIEN, M. F. and F. E. KURCZEWSKI—Further observations on the ethology of
Alysson conicus Provancher (Hymenoptera: Sphecidae)

PINTO, J. D.—New synonymies, lectotype designations, and other notes on North
American Epicauta (Coleoptera: Meloidae)

REICHART, C. V.—An addition to the genus Anisops of Australia (Hemiptera:
Notonectidae)

ROLSTON, L. H.—A revision of Euschistus Dallas subgenus Lycipta Stal (Hemip-
tera: Pentatomidae)

ROTH, L. M.—Ovoviviparity in the blattellid cockroach, Symploce bimaculata (Ger-
staecker) (Dictyoptera: Blattaria: Blattellidae)

SCARBROUGH, A. G.—Coexistence in two species of Holcocephala (Diptera: Asilidae)
in a Maryland habitat: Predatory behavior

SHERRON, D. A., C. G. WRIGHT, M. H. ROSS, and M. H. FARRIER—Density,
fecundity, homogeneity, and embryonic development of German cockroach (Blat-
tella germanica (L.)) populations in kitchens of varying degrees of sanitation
(Dictyoptera: Blattellidae)

SMITH, C. F.—A key to the species in Hyalomyzus (Homoptera: Aphididae) in North
America, with the description of a new species

STEINER, W. E., JR.—Poecilocrypticus formicophilus Gebien, a South American
beetle established in the United States (Coleoptera: Tenebrionidae)

THRELFALL, W.—Ectoparasites (Mallophaga, Acarina) from the double-crested cor-
morant (Phalacrocorax auritus) in Florida

TODD, E. L.—The noctuoid moths of the Antilles—part II (Lepidoptera: Arctiidae:
Pericopinae)

WHEELER, A. G., JR.—Clanoneurum americanum (Diptera: Ephydridae), a leafminer
of the littoral chenopod Suaeda linearis

WOJTOWICZ, J. A.—Description of the larva and female of Pycnopsyche flavata
(Banks) with comparative notes on the ecology of P. flavata and P. gentilis (Mc-
Lachlan) (Trichoptera: Limnephilidae)

NOTES:
KINGSOLVER, J. M.—A 2000+-year-old beetle (Coleoptera: Dermestidae)

RUSSELL, L. M.—The habits and appearance of a rare mealybug, Eurycoccus blanch-
ardii (King and Cockerell) (Homoptera: Coccoidea: Pseudococcidae)

SHUBECK, P. P. and D. L. BLANK—Silphids attracted to mammal carrion at Chelten-
ham, Maryland (Coleoptera: Silphidae)

BOOK REVIEWS
SUMMARY REPORTS OF SOCIETY OFFICERS FOR 1981
SOCIETY MEETINGS

315

297

JULY 1982 NO

"VOL. 84
5 5, 706 io (ISSN ay
fent

PROCEEDINGS

of the

gNTOMOLOGICAL SOCIETY
ot WASHINGTON

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

a
s

7

CONTENTS 4

| ALM, S. R. and F. E. KURCZEWSKI—Antennal sensilla aitd~setae of Anoplius
tenebrosus (Cresson) (Hymenoptera: Pompilidae) .............20.e0cccncsesccen 586

BLANCHARD, A. and E. C. KNUDSON—A new species of Symmetrischema Povolny
oeenidoptera: Gelechiidac) from Texas <2... 6500 .cnsccnse es case caereusesacsinens 628

BROWN, R. L.—Notes on Gretchena: A new species and the synonymy of Gwen-
RBC Me DIdOpterds) LOLmCIGAG), << xincisnmciaints Gs ta scare caste © ide vale sos by scviee Green 594

BURNS, J. M.—Lychnuchoides frappenda from central Mexico joins /unus and zwei-
feli in a lunus group of Atrytonopsis (Lepidoptera: Hesperiidae: Hesperiinae) ... 547
CALABRESE, D. M. and P. TALLERICO—Chromosome study in males of Nearctic
species of Gerris Fabricius and Limnoporus Stal (Hemiptera: Heteroptera: Ger-
CWIKLA, P. S. and P. H. FREYTAG—Three new leafhoppers (Homoptera: Cica-
ICS CUNT CSOCOR ARIAIND — cx vce Bik ec kne tt cans posh Amen es ahinw ans eRe een 632
DARLING, D. C. and N. F. JOHNSON—Egg mortality in the eastern tent caterpillar,
Malacosoma americanum (Lepidoptera: Lasiocampidae): The role of accessory
Mana SECretiONS ANd fe MASS SHAPE ....... ces ccnrcsseeovsassndesesaenesym wae 448
DELONG, D. M.—Some new Neotropical leafhoppers of the subfamilies lassinae and
Deltocephalinae (Homoptera: Cicadellidae) ......... 6. cece cere rere eer eeeeees 610
GILES, F. E. and W. W. WIRTH—Three new species of Serromyia (Diptera:
_ Ceratopogonidae) from Sri Lanka, with new records and a key to the Oriental and
IRIAN SHCCION og cides eva k Sees ots vada s dons e¥ Ob Ua Sa pwRs AN bees eke 440
GLICK, J. I. and G. R. MULLEN—A new species of biting midge of the genus
Alluaudomyia Kieffer (Diptera: Ceratopogonidae) from the southeastern United
OE ES a rer er rer ere Pewee tre ere fie 539

(Continued on back cover)

THE

ENTOMOLOGICAL SOCIETY

OF WASHINGTON

ORGANIZED MARCH 12, 1884

OFFICERS FOR 1982

MARGARET S. COLLINS, President HELEN SOLLERS-RIEDEL, Hospitality Chairm
MAnyYA B. STOETZEL, President-Elect Jay C. SHAFFER, Program Chairn
THOMAS E. WALLENMAIER, Recording Secretary Joyce A. UTMAR, Membership Chair
RALPH E. HARBACH, Corresponding Secretary JOHN F. CARROLL, Custod
F. CHRISTIAN THOMPSON, Treasurer MarGaretT S. Cotiins, Delegate, Wash. Acad.

Davip R. SmitnH, Editor

Publications Committee
E. Eric GRISSELL GEORGE C. STEYSKAL
JOHN M. KINGSOLVER WAYNE N. MATHIS THOMAS E. WALLENMAIER

Honorary President
C. F. W. MUESEBECK

Honorary Members
FREDERICK W. Poos ASHLEY B. GURNEY RAYMOND A. St. GEORGI

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PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 421-425

DESCRIPTION OF A NEW SPECIES OF NOCTICANACE MALLOCH
(DIPTERA: CANACIDAE) FROM SRI LANKA WITH
NOTES ON TWO RELATED SPECIES

WAYNE N. MATHIS

Department of Entomology, NHB 169, Smithsonian Institution, Wash-
ington, D.C. 20560.

Abstract.—Nocticanace taprobane from Sri Lanka is described and com-
pared with two closely related species, N. mahensis (Lamb) and N. sinensis
Delfinado. Appropriate illustrations are provided.

Based on specimens collected by the Lund University Ceylon Expedition
in 1962, Delfinado (1975) reviewed the beach fly fauna of Sri Lanka (Ceylon).
Delfinado described two new species, each in a separate genus, Chaeto-
canace brincki and Xanthocanace zelanica, and included notes on a third
genus and species, Procanace grisescens Hendel. During a collecting trip
to Sri Lanka in 1980, I collected specimens of a fourth genus, Nocticanace
Malloch, all of which are apparently conspecific and represent a new
species. This species is described herein. Notes and illustrations of two
closely related species, N. mahensis (Lamb) and N. sinensis Delfinado, are
included for comparative purposes.

Nocticanace taprobane Mathis, NEw SPEcIES
Figs. 1-3

Diagnostic description.—This species belongs to a species-group of Noc-
ticanace which lacks an anterior notopleural bristle and has the apical scu-
tellar bristles anaclinate. Within that group, it is distinguished from related
congeners, particularly N. mahensis and N. sinensis, by the following char-
acters, mostly from the male terminalia: Small to moderately small beach
flies, length 1.65 to 2.31 mm.

Male terminalia (Figs. I-3) as follows: Surstylus a single, ventrally pro-
jected lobe, moderately wide, tapered abruptly ventrally to form a moder-
ately long, slender, ventral process, posterior margin with approximately 6
short, stout setae.

422

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

hy,

Wrectatet

hw

Figs. 1-3. Nocticanace taprobane. |, External male terminalia, lateral view. 2, Surstylus,

posterior view. 3, Surstylus, lateral view. Figs. 4-6. N. mahensis. 4, External male terminalia,
lateral view. 5, Surstylus, posterior view. 6, Surstylus, lateral view.

VOLUME 84, NUMBER 3 423

Type-material.—Holotype ¢ is labeled ‘SRI LANKA: Galfle]. Dist[rict].
Mirigama 26 April 1980/Collectors: W. N. Mathis[,] T. Wijesinhe[,] L.
Jayawickrema.”’ Allotype and 28 paratypes (11 ¢, 17 2: USNM) bear the
same label data as the holotype. Other paratypes are as follows: SRI LAN-
KA. Trincomalee District: Nilaveli (5 km N) 3 May 1980, W. N. Mathis, T.
Wijesinhe, L. Jayawickrema (2 ¢, | 2; USNM). Hambamtota District: Kir-
inda, 25 Apr 1980, W. N. Mathis, T. Wijesinhe, L. Jayawickrema (9 6, 18
2; USNM). The holotype is double mounted (minute nadel in plastic block),
is in excellent condition, and is in the Smithsonian Institution, Washington,
D.C., USNM 100224.

Remarks.—This species and N. mahensis are very closely related, as
evidenced by the similar conformation of the male terminalia of both
species, and I can distinguish between them by the shape of the apical
process of the surstylus, which is more slender, and by the number and size
of the stout setae along the posterior margin of the surstylus.

The specific epithet, taprobane, is the classical Greek name for Sri Lanka
and is a noun in apposition.

Nocticanace mahensis (Lamb)
Figs. 4-6

Canace mahensis Lamb, 1912: 328.
Nocticanace mahensis: Wirth, 1951: 274.

Type-material.—Holotype 2 is labeled “‘Type H. T. [disc with red
border]/Seychelles Is[land]. Prof. J. S. Gardiner. 1914-537/Seaweed [hand-
written in pencil]/TYPE Canace mahensis. Lamb __ [??, I cannot decipher
what is written after ‘“‘Lamb’’; ‘“‘TYPE” black on blue, printed; otherwise
label black on white, handwritten].”’ The holotype is double mounted (mi-
nute nadel in cork block; block has ‘*107’’ handwritten on top), is in good
condition, and is in the British Museum (Natural History), London.

Remarks.—The illustrations of the male terminalia are from a topotypical
specimen that was compared with the female holotype.

This species is apparently closely related to N. sinensis and N. taprobane,
as evidenced by the similar shape of the surstylus in all three species. When
viewed from slightly different angles, their surstyli can be positioned to
appear to be virtually identical. From a direct lateral view, however, there
are consistent differences, and I can only conclude that three species are
represented. Externally all three species are inseparable.

Male terminalia (Figs. 4-6) are as follows: Surstylus generally compara-
tively wide, narrowed abruptly ventrally to form rather narrow, short ven-
tral process; ventral process moderately tapered, especially in posterior
View; posterior margin with 3—5 short, stout setae on dorsal half of surstylus.

424 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

=

Figs. 7, 8. Nocticanace sinensis. 7, External male terminalia, lateral view. 8, Surstylus,
lateral view.

Nocticanace sinensis Delfinado
Figs. 7, 8

Nocticanace sinensis Delfinado, 1971: 120.

Type-material.—Holotype 2 is labeled ‘“‘HONG KONG: N. T. Sai Kung
Station 27. I. 1965/W. J. Voss & Hui Wai Ming Light Trap BISHOP MUS./
° [handwritten in pencil] HOLOTYPE @ Nocticanace sinensis Md.
[Mercedes Delfinado; handwritten, black on pink].’’ The holotype is double
mounted (glued to a paper point), is in poor condition (the body is covered
with lepidopteran scales, several legs and setae are missing), and the ab-
domen has been removed (the terminalia and other structures of the abdo-

» are slide mounted). The holotype is deposited in the B. P. Bishop
Museum, Honolulu, Hawaii.

Remarks.—This species is evidently closely allied to N. mahensis, as the
shape of the surstylus of both species is very similar, although consistently
differs (Figs. 7, 8) as follows: Surstylus generally more slender; ventral
projection long, somewhat triangular, with distinct taper; posterior margin
with several long stout setae, especially on ventral half.

VOLUME 84, NUMBER 3 425

ACKNOWLEDGMENTS

I thank Brian H. Cogan, British Museum (Natural History), and Neil
Evenhuis, B. P. Bishop Museum, for permitting me to examine the holo-
types of N. mahensis and N. sinensis, respectively. I am also grateful to
Willis W. Wirth, Systematic Entomology Laboratory, USDA, for reviewing
a draft of the manuscript, to S. Dillon Ripley, Secretary of the Smithsonian
Institution, for support to conduct fieldwork through a Fluid Research
Grant, and to Karl V. Krombein, Department of Entomology, Smithsonian
Institution, for encouraging my work on the dipteran fauna of Sri Lanka

and for making arrangements for my fieldwork in Sri Lanka.

LITERATURE CITED

Delfinado, M. D. 1971. New species of shore flies from Hong Kong and Taiwan (Diptera:
Canaceidae). Orient. Insects S(1): 117-124.

| ——. 1975. Diptera: Canaceidae from Ceylon. Pp. 221-223. Jn Reports from the Lund
University Ceylon Expedition in 1962, Vol. II. Entomol. Scand. Suppl. 4.

Lamb, C. G. 1912. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the
Leadership of Mr. J. Stanley Gardiner, M. A. Vol. IV. No. XIX. Diptera: Lonchaeidae,
Sapromyzidae, Ephydridae, Chloropidae, Agromyzidae. Trans. Linn. Soc. Lond. (2)

| 15: 303-3458.

— Wirth, W. W. 1951. A revision of the Dipterous Family Canaceidae. Occas. Pap. Bernice

Pauahi Bishop Mus. 20(14): 245-275.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 426-429

TETRASTICHUS CECIDOBROTER (HYMENOPTERA: EULOPHIDAE),
A NEW PHYTOPHAGOUS SPECIES DEVELOPING WITHIN THE GALLS
OF ASPHONDYLIA (DIPTERA: CECIDOMYIDAE) ON ATRIPLEX
(CHENOPODIACEAE) IN SOUTHERN CALIFORNIA

GORDON GORDH AND BRADFORD A. HAWKINS

Division of Biological Control, Department of Entomology, University of
California, Riverside, California 92521.

Abstract.—Tetrastichus cecidobroter, new species, is described and illus-
trated. Its larvae are phytophagous gall-formers within cecidomyliid galls in
Atriplex spp. in the southwestern United States. This species is the third
phytophagous Tetrastichus known.

Ecological studies by one of us (BAH) on the gall fauna of Atriplex spp.
in southern California have revealed an undescribed species of Tetrastichus
which develops on modified plant tissue in galls caused by species of the
Asphonadylia atriplicis Cockerell complex. The novel biology of this wasp
will be presented elsewhere. The purpose of this paper is to make a name
available for this species. Descriptive terminology is principally after Gra-
ham (1961).

Tetrastichus cecidobroter Gordh and Hawkins, NEw SPECIES
Figs. 1-7

Female.—Holotype 2.7 mm long; body black with very faint and limited
bluish tinge in certain plays of light; antennal scape and pedicel tan, flagel-
lum dusky; coxae black, femora similarly dark colored except apices pale;
tibiae dusky mesad, basal and distal portions tan; pretarsi dusky, tarsomeres
I-III nearly white; wings hyaline with bluish tinge in certain plays of light.

Yead: Slightly wider than mesosoma, in dorsal aspect with weak irregu-
lar, \ransverse reticulate sculpture and scattered pale setae; frontovertex
about 0.6 as wide as head; ocelli forming a strongly obtuse triangle sur-
rounded by a superficial line of weakness. Head in dorsal aspect with scrobal
impression broad and shallow, medially with 2 superficial lines of weak-
ness diverging dorsad and terminating near median ocellus; paraocular area
slightly more pubescent than vertex; toruli just dorsad of imaginary trans-

VOLUME 84, NUMBER 3 427

LL td

yidy!

Vv gy!

Figs. 1-7. Tetrastichus cecidobroter. 1, Female forewing. 2, Female propodeum. 3, Female
habitus, dorsal aspect. 4, Male forewing. 5, Male genitalia. 6, Female antenna. 7, Male antenna.

verse line projecting between ventral margins of compound eyes; ventral
margin of clypeus without tooth-like projections; area between ventral mar-
gin of clypeus and toruli moderately setose. Malar space about 0.7 as long
as compound eye; malar sulcus well defined, complete. Antenna as illus-
trated (Fig. 6).

Mesosoma: Rather robust (Fig. 3); pronotum with posterior margin
strongly arched, anterior face weakly striate or transversely reticulate, lat-
eral face more boldly reticulate; posterior margin with a line of inconspic-
uous setae, lateral face with several pale setae. Middle lobe of mesoscutum
boldly reticulate, without median line, with 5-8 inconspicuous adnotaular
setae forming an irregular line (forming 2 irregular lines when 8 setae are
present); lateral lobe of mesoscutum with sculpture less distinct (weakly,
irregularly reticulate), and setal number 3—6. Scutellum weakly and minutely

428 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

longitudinally reticulate, with 2 inconspicuous setae laterad of each sub-
median longitudinal line, the cephalad setae just anterior of imaginary trans-
verse line bisecting scutellum. Dorsellum minutely and weakly reticulate.
Prepectus and mesepisternum boldly reticulate; anterior 0.6 of mesepimeron
smooth and polished, posterior 0.4 boldly reticulate. Propodeum (Fig. 2)
reticulate (usually boldly), with complete median carina and postspiracular
carina; callus with 2 rather long, pale setae, sculpture less conspicuous.
Hindcoxa boldly reticulate. Forewing as illustrated (Fig. 1), submarginal
vein with 2 widely separated dorsal setae. Apex of hindwing not acute.

Metasoma: Conic-ovate, 1.6 as long as mesosoma; Tergum I smooth-
polished, with 2 setae laterad; T II mostly concealed, but similarly smooth-
polished and with 2 setae laterad; T III finely reticulate and with several
setae laterad; T IV—VI finely reticulate and densely setose; epipygium
densely setose; ovipositor and gonostyli slightly exserted.

Male.—Similar to female in habitus, coloration, sculpture, and chaeto-
taxy. Differing in smaller size, antennal segmentation and shape (Fig. 7),
wing shape (Fig. 4), and genitalia (Fig. 5). Also more variation in coloration
of tibiae (frequently very faintly dusky, nearly tan), and a third seta may be
on the propodeal callus.

Material examined.—Holotype: 2, California, Riverside Co., Palm Des-
ert; 4/XI/1979; ex stem gall on Atriplex canescens; B. A. Hawkins, Collec-
tor. Paratypes: Palm Desert, 6 2, 3 6, 10/II/1979; 1 9, 3/III/1979; 4 2, 2
3, 22/VII/1979: 8 2, 5 3, 29/VIT/1979: 1 2; I2Z2/VINI/19792 5 25 2 oa
1979. California, Riverside Co., nr. Valle Vista, | 2, 11/V/1980; 15 9°, 8
6, 1S/VII/1979; 3 2, 2.6, 23/VIN/1979; 4 6, 13/VIL/1979; 29, 20/Viii is:
22,26, 27/VII/1979; 1 2, 21/X/1979. California, Riverside Co., nr. Ther-
mal, 3 2, 11/V/1980. All material collected and reared from cecidomyiid
galls on Atriplex canescens by B. A. Hawkins.

Holotype, 5 9 and 4 ¢ paratypes deposited in the National Museum of
Natural History, Smithsonian Institution, Washington, D.C. Remaining
paratypes deposited in the Division of Biological Control, University of
California, Riverside.

Etymology.—Cecido (Greek = gall), broter (Greek = eater).

Discussion.—In the most recent revision of the Nearctic Tetrastichus
(Burks, 1943), 7. cecidobroter runs to T. cormus Burks. The new species
'iffers from T. cormus in that the head and thorax are not minutely sha-
greened, the toruli are located higher on the head, the submarginal vein
bears only two dorsal setae, part of the thoracic pleuron is polished, and
the propodeum lacks short anterolateral carinae. Burks (1963) subsequently
described ten additional species of Tetrastichus from material taken in
North America. 7etrastichus cecidobroter displays no affinity with any of
these species.

'

VOLUME 84, NUMBER 3 429

Phytophagy has apparently evolved several times in the Chalcidoidea (see
Gahan, 1922), but it is relatively uncommon in the Eulophidae. Only two
species of Tetrastichus other than T. cecidobroter are known to be phy-
tophagous. Ishii (1931) described T. ardisiae from material taken on Ardisia
japonica (Thunb.) (Myrsinaceae) and reported that the portion of the shoot
in which 7. ardisiae developed ‘swells to some extent.’’ Both 7. ardisiae
and 7. cecidobroter are similar in having four ring segments in the female
antenna. They differ in that the median ocellus of 7. cecidobroter is not
within the scrobal impression, the ventral margin of the clypeus is not
toothed, the mesoscutum lacks a median groove, and the submarginal vein
does not bear seven dorsal setae.

A more convincing case of phytophagy in Tetrastichus has been discussed
by Teitelbaum and Black (1957) who report that a species “‘near venustus”’
develops on sweet clover (Melilotus alba Desc. and M. officinalis (L.)
Lam.). This species has remained undescribed. Burks regarded it as mor-
phologically indistinguishable from 7. venustus Gahan, although the phy-
tophagous form is thelytokous. Nikolskaya (1933) regarded venustus as a
junior synonym of the well-known European species 7. brevicornis (Pan-
zer), but this synonymy has not been accepted in more recent catalogs
(Domenichini, 1966; Burks, 1979).

LITERATURE CITED

Burks, B. D. 1943. The North American parasitic wasps of the genus Tefrastichus—A con-

tribution to biological control of insect pests. Proc. U.S. Natl. Mus. 93: 505-608.

. 1963. Ten new reared species of Tetrastichus (Hymenoptera, Eulophidae). Proc. Biol.

Soc. Wash. 76: 47-58.

—. 1979. Family Eulophidae. Pp. 967—1022. Jn Krombein, K.V. et al., Catalog of Hy-
menoptera in America North of Mexico, Vol. 1. Smithsonian Institution Press, Wash-
ington, D.C.

Domenichini, G. 1966. Index of Palearctic Tetrastichinae. Le Francois, Paris, 101 pp.

Gahan, A. B. 1922. A list of phytophagous Chalcidoidea with descriptions of two new species.
Proc. Entomol. Soc. Wash. 24: 33-S8.

Graham, M. W. R. de V. 1961. A new species of Tetrastichus Walker (Hymenoptera: Eu-
lophidae) from Britain. Entomol. Gaz. 25: 106-110.

Ishii, T. 1931. Notes on the phytophagous habit of some chalcidoids, with descriptions of two
new species. Kontyu 5: 132-138.

Nikolskaya, M. N. 1933. A new species of seed-infesting chalcid-fly, Eurytoma onobrychidis,
sp. n. (Chalcididae) on Onobrychis sativa, and its parasites. [In Russian.] Rev. Entomol.
URSS 25: 119-133.

Teitelbaum, S. S. and L. M. Black. 1957. The effect of a phytophagous species of Tetrastichus,
new to the United Staes, on sweet clover infected with tumor-wound virus. Phytopath-
ology 44: 548-550.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 430-439

ABUNDANCE AND DISTRIBUTION OF THE SEVENTEEN- YEAR
PERIODICAL CICADA, MAGICICADA SEPTENDECIM
(LINNAEUS) (HEMIPTERA: CICADIDAE—BROOD ID),

IN CONNECTICUT

CHrRIs T. MAIER

Department of Entomology, The Connecticut Agricultural Experiment
Station, 123 Huntington Street, New Haven, Connecticut 06504.

Abstract.—Mean density of Magicicada septendecim (Linnaeus), based
on number of emergence holes, ranged from 2.3 to 14.3 cicadas/m? under
apple trees and from 2.2 to 8.0 cicadas/m? under forest trees at three sites.
Density along transects through these sites showed a slight or large buildup
near the orchard and forest margin at two sites. Emergence holes occurred
at 70/72 (97.2%) of sampling stations despite variation in forest character-
istics. Connecticut populations of M. septendecim in Brood II occurred only
in the central part of the state. Examination of 75 sites known to have had
cicada populations in 1945 indicated that four (5.3%) of the populations
disappeared due to human disturbance.

Early surveys conducted by Riley (1885) and Marlatt (1907) indicated that
Brood II and XI of the 17-year periodical cicada, Magicicada septendecim
(Linnaeus), occurred in Connecticut. Brood XI last emerged in 1954 and is
now apparently extinct (Manter, 1974). By contrast, adults of Brood II ap-
peared at many sites in central Connecticut during their scheduled emer-
gence in 1962 (Leonard, 1964). These adults successfully reproduced as
Maier (1980) found nymphs in several Connecticut apple orchards and for-
ests in 1978-9.

Marlatt (1907) reported Brood II in five of eight Connecticut counties, but
Britton (1911) found it in only three counties. Accuracy of these surveys is
unknown because both authors relied heavily upon observations of others
and deposited few specimens in collections. The more detailed study of
Britton (1911) showed that M. septendecim was largely restricted to the
Connecticut Valley in central Connecticut. Britton (1911) made casual ob-
servations on density but provided no quantitative data.

This study was conducted (1) to determine density of cicadas in selected
apple orchards and forests in Connecticut, (2) to characterize forests inhab-

|

VOLUME 84, NUMBER 3 431

ited by cicadas, (3) to provide detailed distributional records, and (4) to
report the fate of 75 populations recorded in 1945. Hopefully, this paper will
furnish basic data necessary for future comparative studies.

MATERIALS AND METHODS

Density at Guilford, Middlefield, and Southington, Connecticut was mea-
sured in mid-July by counting nymphal emergence holes at sampling points
along two parallel, east-west transects/site that were separated by 100 m.
Each transect extended from the center of an apple orchard to approxi-
mately the center of an adjacent forest. Samples were taken at approxi-
mately 10 m intervals in orchards and forests for a distance of 110 m into
each habitat. No sampling was done in the 25 m wide treeless and shrubless
zone that separated orchard and forest. At each sampling point (=each apple
tree in a row), two | m? quadrats located 0.25 m to each side of the transect
were inspected for holes (n = 48 quadrats/transect, 24 in each habitat). To
characterize the forest, all trees (=7.5 cm in diameter) within 5 m of each
sampling point were identified, their diameter at a height of 1.3 m was
measured, and total basal area of each species was calculated.

When density/hectare (ha) was estimated, all area under the continuous
forest canopy was assumed to have been inhabited by nymphs. In the or-
chard only soil under the canopy of apple trees was considered to have been
occupied as few or no nymphs emerged between trees. Thus, estimates of
number/ha in orchards were calculated using mean number/m? (given in
Table 1) for the area under the canopies and 0 for the area outside the
canopies.

Distribution of M. septendecim in Brood II was determined by collecting
adults, nymphs, or nymphal exuviae or by hearing adult singing and finding
exuviae. Although inspections were made in every county in Connecticut,
they were concentrated in the Connecticut Valley where M. septendecim
in Brood II was previously reported (Maier, 1980, and references therein).
In addition, all reports of cicada activity filed by fruit growers (i.e. positive
responses to questionnaires), newspersons, and other citizens in 1979 were
investigated; and most sites known to have had cicadas in 1945 (unpublished
data) were examined. Specimens collected in the survey were deposited in
the insect collection housed in the Department of Entomology at the Con-
necticut Agricultural Experiment Station, New Haven.

RESULTS AND DISCUSSION
ABUNDANCE AND DISTRIBUTION

Although density in sampled areas was higher in the orchard than in the
forest at Guilford and Southington, the estimated density/ha was higher in
forests at all three sites (Table 1). Forsythe (1976) and Maier (1980) found

432 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Density of Magicicada septendecim (Brood II) in apple orchards and adjacent
forests in Connecticut during 1979.

Orchard* Forest
Mean Mean
Number/m* Estimated Number/m? Estimated
Sampling Site (n = 48) Range Number/ha” (n = 48) Range Number/ha”
Guilford 5.0 0-16 14,746 DED -11 22,000
Middlefield 23) 0-9 8,335 6.1 0-41 61,000
Southington 14.3 3-57 57,367 8.0 0-31 80,000

@ Variety, age, canopy diameter, and spacing were as follows: Guilford—MclIntosh and Red
Delicious, 45 years, 6.2 m, 10.7 x 12.2 m; Middlefield—Stayman, 35 years, 7.0 m, 9.8 x 8.5
m; Southington—Baldwin, 45 years, 7.0 m, 9.8 x 9.8 m.

> See Materials and Methods for explanation of calculations.

a similar pattern in overall abundance. Densities recorded in this and my
previous study (Maier, 1980) were well below 386 nymphs/yd? (=462
nymphs/m?) reported by Leonard (1964) for one sample in a Connecticut
orchard in 1962. Estimates of abundance at my study sites fell at the lower
end of ranges reported for other Magicicada populations which consisted
of one or more species (e.g. Dybas and Davis, 1962, and references therein;
Forsythe, 1976).

The density profile along transects differed between sites (Fig. 1). No
clear pattern existed at Guilford; however, at Middlefield and Southington
density in the orchard, forest, or both tended to be higher near the edge.
Density showed no consistent relationship with elevation which fluctuated
less than 6 m along any one transect. Soil moisture, previous oviposition
activity of females, and human disturbance (which was highest at Guilford
and Middlefield) probably contributed in forming the density pattern. For
example, at Middlefield density in the forest was low in wet or submerged
soils which were located between 10 and 40 m along transect | and between
10 and 20 m along transect 2. A very distinct pattern, i.e. higher density
near margins, emerged at Southington where disturbances other than insec-
ticide applications between 1962 and 1978 were minimal.

Abundance pattern at Southington (Fig. 1) typifies that in many upland
areas. Females apparently oviposit preferentially in many species of young
or at least sunlit trees (Lloyd and White, 1976; White, 1980), which abound
aiong the forest border at Southington and other locations. At Southington
adulis readily fly from forest to orchard or vice versa (Maier, 1982). Thus, high
density along forest margins should influence the development of density
profiles in adjacent apple orchards. Density in orchards and forests can be
expected to decline as the distance from the edge of the forest increases.

Magicicada septendecim inhabited forests that usually resembled one
another in species composition but varied in the structure of dominance

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VOLUME 84, NUMBER 3

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434 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Characteristics of Connecticut forests inhabited by Magicicada septendecim
(Brood Il). Basal area is the cross-sectional area of trees in all sampling plots along one
transect. Locations and transects are those in Fig. 1.

Guilford—tTransect | Guilford—Transect 2 Middlefield—Transect |

% of % of % of

Total Total Total

% of Basal % of Basal % of Basal

Total Area Total Area Total Area

Number Number (=33,330 Number Number (=37,363 Number Number (=24,667

Tree Species of Trees of Trees cm?) of Trees of Trees cm’) of Trees of Trees cm’)

Acer rubrum L. 6 Voll Ue l ita 0.8 21 16.37 19

A. saccharum Marsh. 3 3.8 6.6 18 19545 e325 10 7.8 55

Betula alleghaniensis Britton 6 4.7 5.6

B. lenta L. 6 Tall Sell 6 6.5 8.2 1 0.8 0.3

Carpinus caroliniana Walt. 4 Dall 0.9 l 0.8 0.2
Carya cordiformis (Wang.) K.

Koch 1 L3} 0.5

C. glabra (Mill.) Sweet 7 9:0, 24:6 5 5.4 55 10 Way 2c
C. ovata (Mill.) K. Koch 6 Well WS) 7) DD, eps 3 28) 1.8
C. tomentosa Nutt. 2 jiep) es!
Cornus florida L. 22282 6.2 19 20.4 4.3 Dy 1.6 0.6
Fagus grandifolia Ehrh. 5 5.4 1.9 9 7.0 8.0
Fraxinus americana L. 10 122 Sipe RO 22 De, Dal 17 6}574 113.3)
Liriodendron tulipifera L. 12 AY Biles) 1 0.8 N57)
Ostrya virginiana (Mill.) K.

Koch 3 3.8 3)55) 9 97 2.6 2 1.6 0.4
Prunus serotina Ehrh. 1 0.8 0.8
Quercus alba L. 2 2.6 37 7 Toss oy! 5 3.9 4.6
Q. prinus L.

QO. rubra L. 14 10.9 17.4
Q. velutina Lam. 3 3.8 les 2 DD, Sil 7/ 5.4 Seo)
Sassafras albidum (Nutt.)

Nees 4 Sel 4.0
Tsuga canadensis (L.) Carr. 1 1.3 4.2 2 Pep 0.6 14 10.9 8.1
Ulmus americana L. 4 Sel 2.0 l ile il 1.1 1 0.8 0.2

Total 78 93 129

hierarchies (Table 2). Tree species at the same site sometimes changed
markedly along a transect and differed in prevalence and size in two tran-
sects. Nonetheless, nymphs emerged at 70/72 (97.2%) of sampling stations

presented in Fig. |. In five of six transects oaks and hickories accounted
for 54.5-54.2% of total basal area (Table 2). In the other (Middlefield, tran-
sect 2), white ash, red and sugar maple, and tuliptree represented 76.8% of
total basal area as compared to only 0.2% for oak and 0.0% for hickory. A
species of oak had the greatest basal area in three transects and pignut
hickory, sugar maple, and tuliptree in one each. Variation in species com-
position and basal area was probably influenced especially by plant disease

VOLUME 84, NUMBER 3 435

Table 2. Continued.

ES

Middlefield—Transect 2. Southington—Transect | Southington—Transect 2
% of % of % of
Total Total Total
% of Basal % of Basal % of Basal
Total Area Total Area Total Area
. Number Number (=22,952 Number Number (=27,208 Number Number (=39,162
Tree Species of Trees of Trees cm*) of Trees of Trees cm?) of Trees of Trees cm?)
Acer rubrum L. 14 [352) e20l8 12 Lile7 5.8 5 5.0 1.5
A. saccharum Marsh. 34), » B22) 072530 9 8.7 10.4 13 12.9 6.2
Betula alleghaniensis Britton
B. lenta L. 2 1.9 Bez 20 19.4 12.3 11 10.9 8.1
Carpinus caroliniana Walt. ) 8.5 as l 1.0 0.2
Carya cordiformis (Wang.) K.

Koch l 1.0 0.1
C. glabra (Mill.) Sweet 2 1.9 Sy 4 4.0 5.0
C. ovata (Mill.) K. Koch
C. tomentosa Nutt. 2 2.0 2.6
Cornus florida L. l 0.9 02 «311 30.1 1-99 226.9 257, 3.9
Fagus grandifolia Ehrh. 2 220 0.6
Fraxinus americana L. 16 [5% 16.2 11 10.9 4.7
Liriodendron tulipifera L. 6 57 14.8 6 Sey Alles 8 79° 19:9
Ostrya virginiana (Mill.) K.

Koch 14 13.2 8.9 5 4.9 1.9 2 PAY 0.3

Prunus serotina Ehrh.
Quercus alba L. l 1.0 4.2 2 2.0 6.6
Q. prinus L. 13 1226, 22:8 4 4.0 3.0
Q. rubra L. 4 3.9 19.9 6 529) 3100
Q. velutina Lam. 0.9 0.2 2, 2.0 5.9
Sassafras albidum (Nutt.)

Nees I 0.9 0.7 l 1.0 0.3
Tsuga canadensis (L.) Carr. 4 3.8 1.6
Ulmus americana L. 4 3.8 3.9

Total 106 103 101

(e.g. Maier, 1980), soil chemistry and moisture, and various human distur-
bances. In the last century, partial clearing of land or selective logging
occurred at all these sites. Nevertheless, cicadas survived probably because
some trees were left standing after every disturbance.

Periodical cicadas in Brood II occurred in orchards and disturbed forests
in central Connecticut (Fig. 2). Most populations inhabited upland forests
that were protected from extensive disturbance by their location on rocky
soil near basalt ridges, policy of owners, or both. Absence of cicadas in
floodplains in Connecticut should not lend support to arguments on inter-
specific competition and host preference of cicadas (Dybas and Lloyd, 1974;

436 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

WINDHAM

°

° TOLLAND

ie)

LITCHFIELD

eo)

°
NEW LONDON

Fig. 2. Distribution of Magicicada septendecim (Brood II) in Connecticut during 1978-9).
Records from 1978 are extracted from Maier (1980). Numbers 1, 2, and 3 represent sites at
Guilford, Middlefield, and Southington, respectively, which were sampled to obtain data in
Tables 1, 2, and Fig. 1. Solid and hollow circles indicate presence and absence, respectively.
Sampling sites separated by <1 km are represented by the same symbol. Lines delimit counties.

White, 1980). Original floodplain forest was destroyed to provide farmland,
and now only small remnants of second-growth floodplain forest remain
(Dowhan and Craig, 1976). Nearly all sizeable second-growth floodplain
forests lie outside the present distributional range of M. septendecim.

Distributional range of Brood II (Fig. 2), the only extant brood in Con-
necticut (Manter, 1974), resembled that outlined in 1911 (Britton, 1911).
~reviously, Marlatt (1907) reported that M. septendecim in Brood II also
occurred in the two western-most counties. Unfortunately, the validity of
Marlatt s (1907) records cannot be determined because they were apparently
based on correspondence and previously published literature (Riley, 1885),
not on specimens.

I visited 75 sites known to have had cicadas in 1945 (unpublished data)
and found them absent at four (5.3%). These populations had been elimi-

VOLUME 84, NUMBER 3 437

nated by suburban housing developments (three sites) and by construction of
an interstate highway (one site). Britton (1911) listed 13 large populations of
M. septendecim, and of these two (15.4%) were extinct and two were rel-
atively small. The smallest population occurred in an oak-hickory forest at
the northernmost positive site represented in Fig. 2. On June 6, I heard only
three singing males and collected one nymphal exuviae during a 2-hour
search at midday. Populations located on protected lands, especially those
located in parks and forests owned by the state and in forests controlled by
water companies, will probably thrive in the foreseeable future. However,
those on lands suitable for development will probably decline unless human
disturbance is curbed.

COLLECTION DATA

Precise distribution records for M. septendecim of Brood II are given
below. These data are for adults unless noted otherwise. Abbreviations for
roads are: C = Connecticut State highway; I = interstate highway; and
US = United States highway.

HARTFORD CO.: Berlin, 1.8 km NW jet. C-71 and C-364, 17 June 1979;
Berlin 3.0 km SW jet. C-71 and C-364, 23 May 1979, nymphs; Berlin, 3.0
km ENE jet. C-66 and C-120, 23 May 1979, nymph; Berlin, Lamentation
Mt., 3.7 km ESE jet. C-71 and C-364, 4 June 1979; Farmington, 3.2 km NE

_ jet. C-4 and C-10, 6 June 1979, nymphal exuviae and adult singing; Farm-

ington, 1.9 km E jet. US-6 and C-10, 12 June 1979, nymphal exuviae and
adults chorusing; Farmington, Rattlesnake Mt., 0.9 km SE jct. US-6 and C-
10, 16 June 1979; Farmington, 1.4 km S jct. US-6 and C-10, 16 June 1979;
New Britain, 2.4 km W jet. C-71A and C-72, 12 June 1979; Plainville, 2.8
km ENE jet. C-10 and C-72, 12 June 1979; Plainville, 2.5 km SE jct. C-10
and C-72, 19 June 1979; Southington, 3.0 km ENE jet. I-84 and C-10, 12
June 1979; Southington, 2.8 km E jet. I-84 and C-10, 17 June 1979; South-
ington, 2.7 km E jet. I-84 and C-10, 20 Nov. 1978, nymphs; Southington,
3.3 km ESE jet. I-84 and C-10, 23 May—22 June 1979, nymphs and adults
in emergence traps; Southington, 2.8 km W jet. C-71 and C-364, 17 June
1979: Southington, 0.8 km N jct. C-66 and C-120, 4 June 1979; Southing-
ton, 0.9 km WNW jet. C-66 and C-120, 4 June 1979; Southington, 1.2 km
SSE jct. C-66 and C-120, 14 June 1979.

MIDDLESEX CO.: Durham, 3.6 km WNW jet. C-17 and C-68, 2 Oct.
1978, nymphs; Durham, 4.4 km W jet. C-17 and C-68, 5 June 1979; Killing-
worth, 4.0 km SW jet. C-80 and C-81, 17 June 1979; Middlefield, 1.4 km SW
jet. C-147 and C-157, 12 Oct. 1978, nymphs; Middlefield, 3.8 km WNW ject.
C-17 and C-68, 5 June 1979; Middletown, 2.5 km W of Westfield, nr. E.
Bradley Brook, 20 Oct. 1978, nymphs; Middletown, 0.6 km N jet. C-66 and
e-21747 June 1979.

| NEW HAVEN CO.:: Branford, 1.6 km ESE jet. C-80 and C-100, 22 May

438 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1979, nymphs; Branford, 2.6 km SE ject. C-80 and C-100, 13 June 1979,
nymphal exuviae; Branford, 0.9 km S jct. US-1 and C-22, 13 June 1979;
Cheshire, 2.0 km S jet. C-66 and C-120, 6 June 1979; Cheshire, 2.2 km SSE
ject. C-10 and C-70, 13 Oct. 1978, nymphs; Cheshire, 2.5 km S jet.
C-10 and C-70, 13 Oct. 1978; East Haven, 1.5 km N ject. C-80 and
C-100, 1 June 1979; Hamden, 5.7 km NNW jet. C-15 and C-22, 6 June
1979: Hamden, 4.8 km NNW jet. C-15 and C-22, 6 June 1979; Hamden,
Sleeping Giant St. Pk., 4.9 km N jet. C-15 and C-22, 6 June 1979; Hamden,
Sleeping Giant St. Pk., 4.9 km NNE jet. C-15 and C-22, 6 June 1979; Ham-
den, Sleeping Giant St. Pk., 4.3 km NNE jet. C-I5 and C-22, 6 June 1979;
Hamden, 3.1 km NW jet. C-15 and C-22, 20 June 1979, nymphal exuviae
and adult singing; Guilford, 6.4 km NNE jet. C-77 and C-80, 13 June 1979;
Guilford, 2.3 km NNW jet. C-77 and C-80, 2 Oct. 1978, nymphs; Guilford,
2.5 km NE jet. US-1 and C-22, 18 Oct. 1978, nymphs; Guilford, 0.8 km SSE
jct. US-1 and C-22, 13 June 1979; Madison, 2.0 km SSE jet. C-79 and C-80,
13 June 1979; Meriden, East Peak of Hanging Hills, 1.4 km E jct. C-66 and
C-120, 3 Oct. 1978, nymphs; Meriden, Hubbard Pk., 1.7 km ESE jct. C-66
and C-120, 3 Oct. 1978, nymphs; N. Branford, 3.6 km ENE jet. C-17 and
C-22, 26 May 1979, nymphs; N. Branford, 2.3 km SSW ject. C-17 and C-22,
25 May 1979; N. Branford, 2.6 km SSW jet. C-17 and C-22, 4 Oct. 1978,
nymphs; N. Branford, 3.3 km SSW ject. C-17 and C-22, 23 May 1979, nymph;
N. Branford, 3.1 WNW jet. C-77 and C-80, 21 May 1979; N. Branford, 1.0
km N jet. C-80 and C-139, 23 May 1979, nymphs; Wallingford, 1.3 km SE
jet. I-91 and C-68, 5 June 1979; Wallingford, 3.1 km E of E. Wallingford, 5
June 1979; Wallingford, 3.2 km E of E. Wallingford, 7 Dec. 1978, nymphs.

ACKNOWLEDGMENTS

David Wagner provided valuable technical assistance throughout this in-
vestigation. David Carlson, Larry Sacramone, fruit growers, and many oth-
er people assisted in mapping the distribution of cicadas. Mark McClure,
Ronald Weseloh, and anonymous reviewers constructively criticized earlier
drafts of this manuscript. Special thanks are due to Leo Baker, Gene Bish-
op, and Stewart Ramsay who allowed me to survey cicadas in apple or-
chards and forests under their supervision. R. G. Cooper surveyed peri-
odical cicada populations in 1945. His unpublished data used in this paper
are filed in the Department of Entomology, The Connecticut Agricultural
Experiment Station.

LITERATURE CITED

Britton, W. ©. 1911. The periodical cicada or seventeen year locust in Connecticut in 1911.
Conn. Agric. Exp. Stn. Ann. Rep. 35: 296-305.
Dowhan, J. J. and R. J. Craig. 1976. Rare and endangered species of Connecticut and their

habitats. Conn. State Geol. Nat. Hist. Surv., Rep. Investigations No. 6, 137 pp.

VOLUME 84, NUMBER 3 439

Dybas, H. S. and D. D. Davis. 1962. A population census of seventeen-year periodical cicadas
(Homoptera: Cicadidae: Magicicada). Ecology 43: 432-444.

Dybas, H. S. and M. Lloyd. 1974. The habitats of 17-year periodical cicadas (Homoptera:
Cicadidae: Magicicada spp.). Ecol. Monogr. 44; 279-324.

Forsythe, H. Y., Jr. 1976. Estimating nymphal populations of 17-year cicadas in eastern Ohio,
1968. Ohio J. Sci. 76: 95—96.

Leonard, D. E. 1964. Biology and ecology of Magicicada septendecim (L.) (Hemiptera: Ci-
cadidae). J. N. Y. Entomol. Soc. 72: 19-23.

Lloyd, M. and J. White. 1976. On the oviposition habits of 13-year versus 17-year periodical
cicadas of the same species. J. N. Y. Entomol. Soc. 84: 148-155.

Maier, C. T. 1980. A mole’s-eye view of seventeen-year periodical cicada nymphs, Magicicada

septendecim (Hemiptera: Homoptera: Cicadidae). Ann. Entomol. Soc. Am. 73: 147-

152.

. 1982. Observations on the seventeen-year-periodical cicada, Magicicada septendecim

(Hemiptera: Homoptera: Cicadidae), in Connecticut. Ann. Entomol. Soc. Am. 75; 14-23.

Manter, J. A. 1974. Brood XI of the periodical cicada seems doomed, pp. 100-101. /n Beard,
R. L., ed., 25th Anniversary Memoirs, Conn. Entomol. Soc., The Conn. Entomol. Soc.,
New Haven.

Marlatt, C. L. 1907. The periodical cicada. U.S. Dep. Agric. Bur. Entomol. Bull. 71: 1-181.

Riley, C. V. 1885. The periodical cicada. U.S. Dep. Agric. Div. Entomol. Bull. 8: 146.

White, J. 1980. Resource partitioning by ovipositing cicadas. Am. Nat. 115: 1-28.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 440-447

THREE NEW SPECIES OF SERROMYIA (DIPTERA:

CERATOPOGONIDAE) FROM SRI LANKA, WITH

NEW RECORDS AND A KEY TO THE ORIENTAL
AND AUSTRALASIAN SPECIES

FRANCIS E. GILES AND WILLIS W. WIRTH

(FEG) Biology Department, Loyola College, Baltimore, Maryland 21210;
(WWW) Systematic Entomology Laboratory, IIBIII, Agricultural Research
Service, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Three new species of Serromyia are described from Sri Lan-
ka: heveli, maculipennis, and punctata. A key is given for separation of the
six known Oriental and Australasian species of the genus. New Oriental
records are S. esakii Tokunaga from the Philippines and Sabah, and S.
pendleburyi Macfie from Vietnam. Notes are given on the hitherto unknown
female of S. pendleburyi.

The genus Serromyia Meigen is well represented in the Northern Hemi-
sphere with about two dozen species known from the temperate Holarctic
Region (Wirth et al., 1974). It extends into the African continent with five
species (de Meillon and Wirth, 1981), but elsewhere it is absent or poorly
represented. No species have been reported from the Neotropical Region
(Wirth, 1974), only two from the Australasian Region (Debenham, 1970),
and only one heretofore described Oriental species (Wirth, 1973). The pur-
pose of this study is to report three new species from Sri Lanka, to describe
the hitherto unknown female of S. pendleburyi Macfie (1934) from Vietnam,
and to report new records of S. esakii Tokunaga (1940) from Southeast Asia.

In this manuscript the first values given are those of the holotype followed
by the range of variation of the paratypes in parenthesis. The methods of
measurements and ratios used are those of Debenham (1970).

‘he authors are indebted to Karl V. Krombein, Director of the Smith-
sonian Ceylonese Insect Project, for specimens used in this study, and to
Molly Ryan for illustrations used in this paper. USNM indicates the Na-
tional Museum of Natural History, Smithsonian Institution, Washington,
DC:

VOLUME 84, NUMBER 3 44]

KEY TO THE ORIENTAL AND AUSTRALASIAN SPECIES OF SERROMYIA

. Hindfemur and hindtibia dark brown, unbanded: mesonotum dark

DR tet Ee or 2 J SEU OS eT ee ve 2
Hindfemur and hindtibia yellowish with brown bands or other mark-
ings; mesonotum yellowish, at least on sides ................... 3

. Large species, wing 2.2—2.6 mm long; female hindfemur armed ven-

trally with about 40 strong black spines in at least 3 series; costa
long, costal ratio 0.75; male 9th tergum long and slender distally;
Balteh IQhUSCAteW m2. 20. eis 2.0) oi IN eee, pendleburyi Macfie
Smaller species, wing 1.0 mm long; female hindfemur armed ven-
trally with 20 strong spines in 2 series; costa shorter, costal ratio
0.69 in female, 0.60 in male; male 9th tergum short and bluntly
rounded Gistally: halter) pales. 2... rsd oes cle evo reyei Debenham

. Wing with small black spot midway in anal cell (mesonotum and legs

yellowish with numerous small brown punctations at the setal bases;
fore- and midfemora and tibiae with narrow brown bases, hindfemur
with narrow brown base and broad brown apical band, hindtibia
brown on bent basal knee portion, with broad subbasal and narrow
apical brown bands; spermathecae pale yellowish; halter slightly in-
fuscated, posterior side of knob whitish) .. maculipennis, new species
Wing without small black spot midway in anal cell on vein IA .... 4

. Legs yellowish, apices of tibiae brownish, hindtibia brown at base,

hind femur brown, only moderately swollen and bearing 20 strong
ventral spines; halter slightly infuscated; spermathecae pale brown;
OSCE AMON Da 2 niche ysieverses <iasire- rae le eshcghaa ee BIE aves esakii Tokunaga
Legs with prominent dark brown bands or punctations; hindfemur
markedly swollen and bearing 20—28 strong ventral spines; costal
PTR AL ONC WO ae ERE ion 4-6, bpevSuaiesiagonensh PRS ALS ex ich aeieuele Sea tetas ene

”n

. Mesonotum and legs yellow, with prominent brown punctations at

setal bases; apices of tibiae narrowly brown; hindfemur with sub-
apical and tibia with subbasal, narrow brown bands; spermathecae
Beet VC OES I nachan shave psns tenn < Fn555, «Bem gxeed age als ane mts punctata, new species
Mesonotum brown in broad midportion; fore- and midlegs pale
brownish; hindfemur dark brown with broad yellow band in mid-
portion; hindtibia dark brown proximally, yellow on distal ¥; sper-
Mate AS dAGks DEOWM: c,s.5' +h daneer® ea eile Seeeoe oe heveli, new species

Serromyia esakii Tokunaga

Serromyia esakii Tokunaga, 1940: 218 (2; Caroline Islands; fig. wing, an-

tenna); Tokunaga and Murachi, 1959: 402 (¢, 2; Caroline Islands, fig.
male palpus, genitalia).

442 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Distribution.—Caroline Islands, Philippines, Sabah.

New records.—PHILIPPINES: Balabac, Dalawan Bay, 9 Oct. 1961, at
mercury light, Noona Dan Exped., 1 2. SABAH: Labuan I., Borneo, Sept.—
Oct., 1948, March 1950, D. H. Colless, at light, | ¢, 1 9; March 1952;
sweeping, | @.

Serromyia pendleburyi Macfie

Serromyia pendleburyi Macfie, 1934: 280 (6; Malaysia; fig. genitalia); De-
benham, 1970: 165 (compared with C. reyei).

Distribution.—Malaysia, Vietnam.

New record.—Vietnam, Dalat, 26-27 Sept. 1960, J. L. Gressitt, 2 2.

Notes.—The female of this species has not been described before. The
Vietnam females have the wing length 2.2 mm; costal ratio 0.75; hindfemur
with 40 ventral spines in at least 3 series, hindclaw 1.28 as long as Sth
tarsomere, the short, slender basal barb only 0.15 as long as claw. This
species resembles the Northern Hemisphere species in its large size and
uniformly dark brown color with only the tarsi and the four anterior legs
yellowish.

Serromyia heveli Giles and Wirth, NEw SpPeEcIEs
Fig. |

Female holotype.—Wing length 1.07 mm, breadth 0.43 mm.

Head: Yellowish brown, setae long and strong. Eyes bare, separated by
distance equal to 3 facets. Antenna (Fig. la) pale, apices of segments 3-10
light brown, segments 11-15 darker brown; verticils well developed on all
segments; lengths of flagellar segments in proportion of 36-2 1-24-26-28-28-
30-30-42-40-50-52-66, antennal ratio 1.12. Palpus (Fig. 1b) pale; 3rd segment
moderately swollen from base with long capitate sensilla irregularly clumped
on distal 4; palpal ratio 2.36. Proboscis brown, moderately long, P/H ratio
0.56; mandibular teeth 9 (8), large and coarse.

Thorax: Yellowish; scutum dark brownish mesally, yellowish on sides;
pleuron yellowish with brown blotches dorsad, ventral areas darker brown;
thorax with strong setae, scutellar setae slightly longer than hindclaw. Legs
(Fig. 1d—f) brownish; fore- and midtrochanters yellowish, hindcoxa yellow-
ish subdorsally, hindtrochanter dark brownish; forefemur and tibia yellow-
ish brown; midfemur with dark brownish base, distal 74 and tibia pale
b-own; hindfemur yellowish brown, basal % dark brown except for small
yellowish area at junction of trochanter, distal '/; dark brown; hindtibia
yellowish brown, basal % dark brown. Hindfemur swollen, measuring 0.63
by 0.14 mm, bearing 20 (21) strong ventral spines, the basal one weaker and
lighter in color; hindtibia arcuate, with comb of 6 (8) spines. Tarsi pale;
hindtarsal ratio 2.03; fore- and midleg with claws small and equal, each with
well-developed teeth, ratio of length of claw to Sth tarsomere on Ist leg

~ VOLUME 84, NUMBER 3 443

a
2 a
an %
N
“yp
NINA e
ae y SF N
y,
Rey a = —<— 3
e
Be ae — So
\< —— — SS SSH % ay
ERR ERE f ) g h JING

Fig. 1. Serromyia heveli, 2 holotype. a, Antenna. b, Palpus. C, Wing. d, Foreleg. e, Midleg.
f, Hindleg. g, Spermathecae. h, Sth tarsomere and claw of hindleg.

0.45, on 2nd leg 0.50, hindleg with single claw (Fig. lh) 1.10 as long as 5th
tarsomere and with fine basal tooth 0.29 the length of claw.

Wing (Fig. Ic): Milky white, with fine microtrichia; base of veins Rs and
1A brownish; Ist and 2nd radial cells slightly yellowish; 22 short setae along
costa to tip of 2nd radial cell, 7 on radius; vein R4+5 obliquely meeting
costa; radial cells with lumens poorly developed; Ist and 2nd radial cells
respectively 0.122 by 0.019 mm and 0.169 by 0.026 mm; vein M2 obsolescent
only a short distance at base; cell M4 extending from level of base of r-rm
crossvein to a level past tip of 2nd radial cell; costal ratio 0.66. Halter
yellowish.

Abdomen: Yellowish with faint segmental dark bands, terminal segments
brownish. Spermathecae (Fig. 1g) 2, dark brown with minute hyaline per-
forations; slightly unequal, measuring 0.094 by 0.055 mm and 0.087 by 0.062
mm; necks arising obliquely, long and tapering; rudimentary spermatheca
present.

Male.—Unknown.

Distribution.—Sri Lanka.

Type.—On slide in phenol balsam. Holotype 2, Sri Lanka, Mannar Dis-
trict, 4 mi NW Mannar, UV light, 100 ft, 3 Nov. 1976, coll. Hevel et al.
(Type no. 76579, USNM).

Discussion.—This species is named for Gary F. Hevel of the Smithsonian
Institution in appreciation of his interest and assistance in the collection
of Sri Lanka Ceratopogonidae.

444 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Te Eee es ee

= =
oe ~
EEE Fae \
an \
/ ZO =
C [ EVA
( VY /, i)
| hee Nf
- we N Wi AS /
/ Cae Wee
[ N ak)
\ / 8
ae : ti S | \ (Gas ~
2) @ f alee an
PS a Ih a
%) , 1
7 | | ~
GE EEE eee SS
r a a a. } Se ———— ho Wika

J
Fig. 2. Serromyia maculipennis, 2 holotype. a, Antenna. b, Palpus. c, Wing. d, Sperma-

thecae. e, Mandible. f, Head, anterior view. g, Foreleg. h, Midleg. i, Hindleg. j, 5th tarsomere
and claw of hindleg.

Serromyia maculipennis Giles and Wirth, New SPEctEs
Fig. 2

Female holotype.—Wing length 1.44 (1.34-1.66, n = 7) mm; breadth 0.58
(0.51-0.60, n = 7) mm.

Head: Yellowish with brown dots at setal bases. Eyes (Fig. 2f) bare,
separated by a distance equal to 2 facets. Antennal segments 3—10 stramin-
eous, 11-15 pale brown, verticils well developed on all segments; flagellar

cements (Fig. 2a) with lengths in proportion of 42-34-34-32-34-32-34-36-59-
62-/0-74-94, antennal ratio 1.38 (1.20-1.38, n = 4) mm. Palpus (Fig. 2b) pale
yellow, ist, base of 2nd, and tip of Sth segments darkest; lengths of segments
in proportion of 20-47-74-40-60; 3rd segment cylindrical with sensilla scat-
tered on surface; palpal ratio 3.70 (3.40-4.44, n = 7). Proboscis brown,
moderately long, P/H ratio 0.57 (0.51-0.65, n = 7); mandibular teeth (Fig.
2e) 11 (10-13, n = 7), large and coarse.

~ VOLUME 84, NUMBER 3 445

Thorax: Yellowish, with brown setal bases; a broad median brown band ex-
tending the length of mesonotum, extending ventrad to meet pleuron between
anterior spiracle and wing base; pleuron with brown splotches. Legs (Fig.
2g-i) yellowish with brown setal bases; fore- and midcoxae and trochanters
with brown streaks, hind ones dark brown; femora and tibiae with setal
punctations tending to form transverse bands; fore- and midfemora with
basal and apical brown bands, hindfemur with basal and subapical brown

_ bands; tibiae with apical brown bands: mid- and hindtibiae with subbasal

brown bands; hindfemur swollen, measuring 0.09 by 0.21 mm, bearing 28
(26-34, n = 7) strong ventral spines; hindtibia arcuate, with comb of 10 (10-
12, n = 7) spines; tarsi yellowish; hindtarsal ratio 2.13; fore- and midlegs
with claws subequal and simple, empodium small; ratio of length of claw to
Sth tarsomere on first leg 0.44, on 2nd leg 0.47; hindleg with single striated
claw (Fig. 2j) 1.55 as long as Sth tarsomere, and with fine basal tooth 0.16
the length of claw.

Wing (Fig. 2c): Hyaline, with fine whitish microtrichia; subcosta, radius,
and M1+2 strong, yellowish, tip of vein R4+5 darker; large setae numerous
the length of costa to end of 2nd radial cell, 7 setae scattered along radius;
Ist radial cell measuring 0.197 by 0.035 mm, 2nd radial cell 0.254 by 0.054
mm with tip quadrate; vein M2 interrupted at base; cell M4 extending from
level of a line % the length of r-m crossvein proximad of the crossvein
distally to the level of end of 2nd radial cell; a spot on vein 1A at midlength
of anal cell composed of densely packed long microtrichia, the spot brown-
ish with a small green area iridescent in some lights; costal ratio 0.70 (0.68—
0.72, n = 7). Halter pale brownish, posterior portion of knob white.

Abdomen: Yellowish, setal bases and terga brown, posterior segments
tending to darken; spermathecae (Fig. 2d) pale brownish, 2, ovoid with
short, slender, oblique necks; unequal, measuring 0.166 by 0.123 mm and
0.129 by 0.104 mm.

Male.—Unknown.

Distribution.—Sri Lanka.

Types.—All on slides in phenol balsam. Holotype 2 Sri Lanka, Jaffna
District, Chundikkulam Sanctuary, 25 ft, UV light, 7 Nov. 1976, coll. G. F.
Hevel, R. E. Deitz IV, S. Karunaratne, and D. W. Balasooriya (Type no.
76580, USNM). Paratypes, 26 2, same data as holotype. Holotype and 24
paratypes deposited in USNM; one paratype will be deposited in the Colom-
bo National Museum, Colombo, Sri Lanka, and another in the National
Museum at the University of Sri Lanka at Peradeniya. Paratype 2, Sri
Lanka, Mannar District, 4 mi NW Mannar, UV light, 100 ft, 3 Nov. 1976,
coll. Hevel et al. (deposited in USNM).

Discussion.—Serromyia maculipennis derives its name from the promi-
nent dark spot in the anal cell. It can readily be identified since all others
in this genus have clear wings.

446 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

—— Se SSS : ee
SS {SS »
a IEEE IIL Ss ae ——~ ‘eS ( “4 b
ESE SE a —
——- re ena a
~ Cx ~
) ber oa
[ t
p> f ; } ye af!
M inna <a ( i? ( ,
DHT TANT TMT AIT S / ul
I G alt 7 0
S. oa = rte aoe ko 255) ——— \
= d . \
eres | \
oS EE SSS e { \
\
> — I eae \
BS ae aa Vaca =i | \
SE NDR ERA ba
so SUTURE f ) g Mp h

Fig. 3. Serromyia punctata, 2 holotype. a, Antenna. b, Palpus. c, Wing. d, Foreleg. e,
Midleg. f, Hindleg. g, Spermathecae. h, Sth tarsomere and claw of hindleg.

Serromyia punctata Giles and Wirth, New Species
Fig. 3

Female holotype.—Wing length 1.20, breadth 0.48 mm.

Head: Yellowish brown. Eyes bare, contiguous for a distance equal to |
facet. Antenna (Fig. 3a) with segments 3-10 pale yellow, 11-15 becoming
progressively darker brown; length of flagellar segments in proportion of 34-
26-26-26-28-26-28-30-46-48-58-60—-71, antennal ratio 1.26. Palpus (Fig. 3b)
pale, segments | and 5 brownish; lengths of segments in proportion of 1I1-
29-50-29-36, 3rd segment slightly swollen from base, with long capitate sen-
silla scattered on surface; palpal ratio 3.1. Proboscis brown, moderately
long, P/H ratio 0.63; mandibular teeth 12 (13), large and coarse.

Thorax: Yellowish with brown setal bases; pleuron with brown blotches.
Legs (Fig. 3d-f) yellowish with brown setal bases; foreleg with coxa brown,
trochanter yellowish, femur with faint brownish basal band, small brown
blotches on flexor surface, tibia with small brown blotches on extensor
surface and narrow apical brown band; midleg with coxa and trochanter
i¢ht brown, femur with narrow base brown, tibia with small brown blotches
on extensor surface and narrow apical brown band; hindleg with coxa brown
on distal '4, trochanter brown, femur with irregular brown basal area, nar-
row brown supapical band and irregular brown splotches throughout, tibia
with irregular brown areas distally, brown subbasal band, and brown blotch-
es on extensor surface; hindfemur swollen, measuring 0.69 by 0.16 mm,
bearing 27 (28) strong ventral spines; hindtibia arcuate, with 9 spines in

VOLUME 84, NUMBER 3 447

tibial comb. Tarsi yellowish; hindtarsal ratio 1.37; fore- and midlegs with
claws small and equal, empodium small; ratio of length of claw to Sth tar-
somere 0.45 on foreleg, 0.40 on midleg; hindleg with single striated claw
(Fig. 3h) 1.72 as long as Sth tarsomere, with fine basal tooth (paratype—
tooth 0.25 as long as claw).

Wing (Fig. 3c): Milky white, with fine microtrichia; base of radius, and

1A, and tip of vein R4+5 dirty yellowish, costa, M1+2, r-m crossvein and
distal % of Rs yellowish; 30 short setae scattered the length of costa to tip
_of 2nd radial cell, only 3 setae on radius; vein R4+5 with beadlike swelling
distally; Ist and 2nd radial cells measuring respectively 0.080 by 0.024 mm
and 0.221 by 0.038 mm; tip of 2nd radial cell quadrate; costal ratio 0.68; cell
M4 extending from level of base of r-m crossvein to a level opposite tip of
2nd radial cell. Halter pale brownish.

Abdomen: Pale yellowish; a pair of brownish spots on each tergum. Sper-
'mathecae (Fig. 3g) 2, pale yellowish scarcely discernible; oval with short
slender necks; unequal, measuring 0.156 by 0.104 mm and 0.115 by 0.071
mm.

Male.—Unknown.

Distribution.—Sri Lanka.

Types.—All on slides in phenol balsam. Holotype 2 , Sri Lanka, Puttalam

District, Wilpattu National Park, 14-15 Feb. 1979, coll. K. V. Krombein,
_T. Wijesinhe, S. Siriwardane, and T. Gunawardane (Type no. 76581,
_USNM). Paratype 2, Sri Lanka, Jaffna District, Chundikkulam Sanctuary,
dt. UV light, 7 Nov. 1976, coll. G. F. Hevel, R. E: Deitz IV, S. Karu-
-naratne, and D. W. Balasooriya (USNM).

LITERATURE CITED

Debenham, M. L. 1970. Australasian Ceratopognidae (Diptera, Nematocera). Part XIV: The
genus Serromyia Meigen. Proc. Linn. Soc. N.S.W. 94: 160-165.

Macfie, J. W. W. 1934. Report on a collection of Ceratopogonidae from Malaya. Ann. Trop.
Med. Parasitol. 28: 279-293.

Meillon, B. de and W. W. Wirth. 1981. Subsaharan Ceratopogonidae (Diptera) VII. The biting
midges of the Kruger National Park, South Africa, exclusive of the genus Culicoides.
| Ann. Natal Mus. 24: 563-601.

Tokunaga, M. 1940. Ceratopogonidae and Chironomidae from the Micronesian Islands. Phil-
ipp. J. Sci. 71: 205-226.

Tokunaga, M. and E. K. Murachi. 1959. Diptera: Ceratopogonidae. Insects Micronesia 12:
103-434.

Wirth, W. W. 1973. Family Ceratopogonidae (Heleidae). Pp. 346-388. Jn Delfinado, M. D.
and D. E. Hardy, eds., A Catalog of the Diptera of the Oriental Region. Vol. 1. Univ.
Press of Hawaii, Honolulu. 618 pp.

| . 1974. Family Ceratopogonidae. A Catalogue of the Diptera of the Americas south of

the United States, Vol. 14. Sao Paulo, Brazil. 89 pp.

Wirth, W. W., N. C. Ratanaworabhan, and F. S. Blanton. 1974. Synopsis of the genera of
Ceratopogonidae (Diptera). Ann. Parasitol. 49: 595-613.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 448-460

EGG MORTALITY IN THE EASTERN TENT CATERPILLAR,
MALACOSOMA AMERICANUM (LEPIDOPTERA:
LASIOCAMPIDAE): THE ROLE OF ACCESSORY

GLAND SECRETIONS AND EGG MASS SHAPE

D. CHRISTOPHER DARLING AND NORMAN F. JOHNSON

Department of Entomology, Cornell University, Ithaca, New York 14853;
NFJ, present address: Department of Entomology, The Ohio State Univer-
sity, 1735 Neil Ave., Columbus, Ohio 43210.

Abstract.—Four species of hymenopteran parasitoids were reared from
egg masses of the eastern tent caterpillar, Malacosoma americanum (F.)
(Lepidoptera: Lasiocampidae) in New York State: Telenomus clisiocampae
Riley (Scelionidae), Tetrastichus malacosomae Girault (Eulophidae), Ooen-
cyrtus clisiocampae (Ashmead) (Encyrtidae), and Ablerus clisiocampae
(Ashmead) (Aphelinidae). Three classes of host eggs were distinguished,
exposed eggs, eggs covered with spumaline on the margin of the mass, and
eggs covered with spumaline in the interior portion of the mass. Parasitism
rates of exposed (27%) and covered marginal eggs (23%) were significantly
greater than covered interior eggs (1%). Inviability was greater in exposed
and covered marginal eggs relative to interior eggs. An analysis of relation-
ships within Malacosoma indicates that frothy spumaline is an evolution-
arily labile trait and has undergone either multiple elaborations or parallel
reductions. The hypothesis that frothy spumaline has evolved convergently
in Malacosoma species as a response to hymenopteran egg parasitoids is
discussed.

Egg parasitoids (primarily of the hymenopteran superfamilies Chalcidoi-
dea and Proctotrupoidea) are one of the most important biological factors
affecting the survival of many species of Lepidoptera and Hemiptera. Their
influence is not as widely appreciated as that of larval, nymphal, and pupal

orasitoids, in large part because of the difficulty in sampling egg popula-
tions. Where data are available, commonly more than 50% of the eggs are
parasitized (Anderson, 1976). Such high rates indicate strong selective pres-
sures on the host to reduce loss, for example, by elimination of the cues
used by parasitoids to locate the eggs, or to provide the eggs with a means
of protection if they are found.

The oviposition behavior of species in the genus Malacosoma Hiibner

VOLUME 84, NUMBER 3 449

(Lepidoptera: Lasiocampidae) is particularly interesting with regard to egg
parasitism. The European pest species M. neustria (L.) is typical of all Old
World species of the genus. It lays its eggs in a single-layered exposed mass
on twigs; a typical mass contains 140-200 eggs (Schwenke, 1978). In these
masses parasitism rates of 40% have been recorded (Romanova and Loz-
inskij, 1958).

In contrast parasitism rates in New World Malacosoma have never been
found to exceed 15% and are usually much lower (Anderson, 1976; Witter

_ and Kulman, 1972). All New World species except M. tigris (Dyar) cover

their egg masses with a foamy, accessory gland secretion known as spu-

- maline. Hodson (1939) noted that egg parasitism of M. disstria Hiibner is

concentrated in eggs along the margin of the mass and suggested that this
was due to the incomplete covering of spumaline in this area. Other authors
have anecdotally referred to prevention of parasitism as the function of
spumaline (e.g. Stehr and Cook, 1968; Wellington, 1965; Witter and Kul-
man, 1979). Prentice (1953) reported that M. disstria eggs covered with
spumaline were less heavily parasitized (2.3%) than those exposed (60.9%
in partially covered masses; 72.5% in completely exposed masses). He cited
only these percentages and gave none of the information necessary to eval-

uate his conclusion (e.g., sample size, variation). The function of spumaline

therefore merits further investigation.

In this paper we present natural history information on the egg parasitoids
associated with Malacosoma americanum (F.). We compare parasitism and
inviability rates of eggs as functions of both position within the mass and
spumaline covering. We discuss the possible role of egg parasitoids in the
evolution of oviposition behavior in the genus Malacosoma.

METHODS AND MATERIALS

Egg masses of Malacosoma americanum were collected in the spring of
1980 prior to the hatching of caterpillars. A total of 90 egg masses were
collected from four localities in central and western New York State (Ithaca,
Beebe Lake, Tompkins Co., n = 17 egg masses; Ithaca, Cascadilla Creek,
Tompkins Co., n = 30; Texas Hollow Wildflower Preserve, Schuyler Co.,
n = 28; and Dunkirk, Chautauqua Co., n = 15). Egg masses were then
scored as either completely covered with spumaline or as having some ex-
posed eggs. All exposed eggs were marked with paint to allow recognition
after caterpillar emergence.

Egg masses were maintained at 21°C, 63% relative humidity, and 12/12
hour photoperiod until after the hatching of caterpillars and parasitoids.
Unhatched eggs were then examined to determine both parasitism and in-
viability rates.

Data collected from each parasitized egg consisted of identity of parasit-
oid, date of emergence, position in the egg mass, and whether or not it was

450 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Composition of egg parasite fauna on Malacosoma americanum from 4 localities
in New York. [% composition and (number of individuals). Total: 937 parasitoids. ]

Locality Tetrastichus Telenomus Ooencyrtus Ablerus

(n = egg masses) malacosomae clisiocampae clisiocampae clisiocampae
Dunkirk

(7 — 13) .541 (92) 159) (27) .294 (50) .006 (1)
Beebe Lake

(a = 159) -900 (226) .068 (17) .028 (7) .004 (1)
Cascadilla Ck.

(n = 26) .761 (124) .147 (24) -080 (13) .012 (2)
Texas Hollow

(ne— 22) .926 (327) .065 (23) .008 (3) —
Totals .821 (769) .097 (91) .078 (73) .004 (4)

covered with spumaline. Four species of hymenopteran egg parasitoids were
encountered in this study: Ablerus clisiocampae (Ashmead) (Aphelinidae)
which appeared first, followed, in order of emergence, by Telenomus cli-
siocampae Riley (Scelionidae), Ooencyrtus clisiocampae (Ashmead) (En-
cyrtidae), and Tetrastichus malacosomae Girault (Eulophidae). Vouchers
of specimens are deposited in the Cornell University Insect Collection,
lot no. 1107. Locations of parasitized host eggs in the mass were scored
as exposed, not covered with a layer of frothy spumaline; covered mar-
ginal, on the perimeter of the egg mass or next to an exposed egg;
or covered interior, and ranked according to the number of eggs by
which it was separated from the margin. Another source of egg mortality
was the failure to produce either a caterpillar or a parasitoid wasp. These
were termed inviable eggs, and their positions within the egg mass were
also recorded. Ten egg masses were then randomly chosen for a detailed
analysis of parasitism rate, inviability rate, and the distribution of parasit-
ized eggs in the egg mass.

A weakness in previous studies on egg parasitoids has been the failure to
separate eggs that do not hatch from those that produce parasitoids. Pre-
vious investigators simply counted and identified the parasitoids and esti-
mated the size of the egg mass (e.g. Hodson, 1941; Romanova and Lozinskij,
1958). This procedure does not allow assessment of the relative contribu-
tions of parasitism and inviability to mortality.

ir procedure determined the fate of each egg along with its position and
the «.stribution of spumaline on the egg mass. Parasitoid identity was es-
tablished by the distinctive shape and color of the meconium of each
species. Telenomus clisiocampae produces a yellowish-green amorphous
meconium; Tetrastichus malacosomae deposits large dark green, ovoid pel-

VOLUME 84, NUMBER 3 451

| Table 2. Distribution of parasitized and inviable eggs in ten egg masses of M. americanum.
(Exp = exposed eggs; Marg = marginal eggs; Int = interior eggs; as defined in text.)
—EeyEe—————————E—————— eee

Parasitism Rate Inviability Rate

Egg No. No. No No.

Mass Egg Exp Marg Int Exp Marg Int Exp Marg Int
ccl9 145 3 43 99 0 .209 0 35 0 0
ccl8 339 2 65 272 0 .123 .004 -500 .062 .004
ccl2 387 3 39 345 7333 .462 .020 .667 .077 .009

/ecll 290 18 77 195 0 .169 0 .278 .208 031
cc4 120 3 43 74 0 0 0 0 .046 027
th29 272 31 81 160 452 Pai fez 0 .258 .062 0
th21 339 36 56 247 361 232 .012 .056 089 .004

thd 216 18 45 153 333 333 0 167 .044 .007
th8 203 8 51 144 .750 OAs .062 .250 AES, .076
bl12 274 = 64 206 500 az. .005 0 0 0
Mean 258.5 12.6 56.4 189.5 ips 225 .010 Beil .073 .016

SD BersQmevl2:6 9514.8. 81,9 r 263%, 195 901d Bue 062 024

lets; Ooencyrtus clisiocampae and Ablerus clisiocampae both produce small

black pellets. The latter two species were distinguishable because their
‘emergence periods did not overlap. Successful development of the cater-
pillar was determined by a completely empty opened egg (chorion), with
neither yolk nor meconium, or by the presence of the caterpillar within
_unhatched eggs. The ten egg masses contained at least one viable exposed
-egg. We used f-tests to compare the mean differences in parasitism and
inviability rates in the ten egg masses for all three pairwise combinations of
-egg location (exposed versus interior, exposed versus marginal, marginal
versus interior).

RESULTS

| Malacosoma americanum populations have been relatively large in the
study sites in the three years preceding this study (1976-1979), and a high
proportion of the egg masses were parasitized (89%). Interlocality variation
in the proportion of masses parasitized varied from 100% at Texas Hollow
to 76% at Cascadilla Creek. Exposed eggs occurred in 64% of the 76 egg
masses, with the number of exposed eggs usually quite low (mean = 13,
SD = 10).

Table | documents the composition of the egg parasitoid fauna of M.
americanum in the four upstate New York localities; Tetrastichus is clearly
the most abundant egg parasitoid. The same rank order of these four para-
sitoids was found in a study conducted in Ithaca, New York, in 1924 (Liu,
1926), but in that study Telenomus was a much more important mortality

452 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 3. Comparison of the distribution of the 3 most common species of parasitoids within
egg masses of M. americanum. The x? values test the null hypothesis of equal numbers of
parasitoids in marginal and interior eggs classes. Note that the number of eggs in the two
classes are not equal (Table 2). [% of occurrences, (number of individuals). ]

ee

Species Marginal Eggs Interior Eggs x?
Tetrastichus malacosomae .730 (492) .270 (182) IPA
Telenomus clisiocampae .518 (56) .482 (52) .074 n.s.
Ooencyrtus clisiocampae 427 (32) .573 (43) 811 ns.

4 Significant at P = .001; n.s.: P > .25.

factor (30% of reared wasps versus 10%). Parasitoid abundance varies on
a regional basis; parasitism by Ooencyrtus ranged over an order of magni-
tude in this study alone.

Table 2 summarizes the results of the detailed analysis of ten egg masses.
Parasitism rates are significantly higher in exposed than interior eggs (t =
4.69, 9 df, P < .001), and significantly higher in marginal than interior eggs
(t = 8.02, 9 df, P < .0005). Exposed and marginal eggs do not differ sig-
nificantly in parasitism rate (t = .950, 9 df, P > .10).

Inviability rates were significantly higher in exposed than marginal eggs
(t = 3.75, 9 df, P < .01), higher in marginal than interior eggs (t = 4.94, 9
df, P < .0005) and higher in exposed than interior eggs (t = 4.84, 9 df,
P < .0005).

The distribution of the three most common species of parasitoid within
the egg mass was also compared (n = 90 egg masses). Table 3 summarizes
the results. Telenomus and Ooencyrtus were equally common in the interior
and marginal egg classes. Tetrastichus is more closely associated with the
egg mass margin (73%) and this distribution is significantly different from
that of both Ooencyrtus (x? = 28.9, P < .001) and Telenomus (x? = 20.0,
Br=, 00).

DISCUSSION

In Malacosoma americanum egg survival is strongly correlated with the
relative position in the egg mass and with the covering of frothy spumaline.
Exposed eggs, i.e. those without a spumaline covering, average 27% para-
sitism and 25% inviability rates. The most vulnerable position within the

overed portion of the egg mass is along its perimeter (23% parasitism, 7%
inviability). The same general trends appear to be true for M. disstria. Clear-

ly, the best oviposition strategy for a female of Malacosoma is to cover all
eggs with spumaline and to minimize the number of marginal eggs in the
mass.

The number of marginal eggs can be changed by altering the shape of the

VOLUME 84, NUMBER 3 453

Fig. 1. Egg masses of M. americanum (bottom) and M. disstria (top). M. americanum has
a Clasping egg mass; M. disstria has a helical egg mass. Note the deviation of the marginal

eggs of M. americanum from the vertical orientation.
|

‘egg mass. Two general egg mass shapes are found in Malacosoma, clasping
(found in americanum, californicum (Packard), and incurvum (Hy. Ed-
wards) and helical (all other species). These terms refer to the final shape

of the mass and are a result of differences in oviposition behavior. According

-to Stehr and Cook (1968), in all species the female oviposits with the lon-

-gitidunal axis of the body perpendicular to the twig. She swings her abdomen

from side to side, laying eggs in a single row, and then moves forward and

-around the twig. The difference in shape between the two kinds of masses

results from varying numbers of eggs laid in each row. Those species form-

ing a helical mass lay only three to four eggs before moving around the twig.

The female completely circles the twig several times and packs the eggs as

tightly together as possible. The resulting egg mass is roughly cylindrical.

The clasping species lay a row of ten to fifteen eggs. As the female oviposits

she moves around the twig, but generally does not move far along its lon-

gitudinal axis. The egg mass is usually saddle-shaped, occasionally com-
pletely encircling the twig and sometimes even overlapping upon itself

(Fig. 1).

454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The type of egg mass will determine the proportion of eggs placed in the
vulnerable marginal row. Assume that the eggs are cubic in shape with unit
length (u) for each side and unit area for each face. Assume also that the
clasping mass has a circular outline so as to minimize the number of marginal
eggs in the egg mass (note, an elliptical egg mass will have an even greater
number of marginal eggs). Consider a circular clasping egg mass of 260 eggs
(the average for M. americanum, see below) laid so that the mass com-
pletely encircles the twig, but does not overlap itself. The twig would have
a circumference of 18.2 eggs, the diameter of the egg mass (ar? = 260u, r =
9.097u, d = 2r = 18.2u). This egg mass would have 57 marginal eggs along |
the circumference of the mass (circumference = 2 mr = 27 9.09u = 57.7u),
and 22% of the total eggs would be marginal. A helical egg mass laid as a
perfect cylinder would have two circles of marginal eggs, one at each end
of the egg mass. If laid on the same diameter twig, 18.2u, there would be a
total of 36 marginal eggs, with 14% of the total eggs on the margin. Clearly,
the optimal egg mass shape, the one minimizing the number of vulnerable
marginal eggs, is the helical (cylindrical) egg mass.

Considerations of the phylogeny of Malacosoma (see below) indicate that
the clasping egg mass is derived (apomorphic) and the helical mass ancestral
(plesiomorphic). How can we explain the evolution of this change in ovi-
position behavior, considering the relatively high marginal mortality rates
and assuming the presence of frothy spumaline?

One possibility is that we have overestimated the vulnerability of some
of the marginal eggs in clasping egg masses. In those cases in which the
lateral edges of the mass are closely approximated and the space between
them filled with spumaline, eggs that by our definition are marginal are
probably as well protected as interior eggs. If egg masses are laid on very
small twigs the clasping egg mass may completely encircle the twig. The
last laid row(s) are then laid atop the first laid row(s). In the instances in
which we found overlapping eggs, those in the lower layer were rarely
parasitized (1%, n = 76 eggs). Nevertheless, in the majority of masses the
eggs do not encircle the twig (81%, n = 63).

We have also observed that the long axes of the marginal eggs of M.
americanum are usually slanted both away from the perpendicular and away
from the center of the mass, sometimes to the extent that the eggs are lying
parallel to the twig. This is in contrast to the eggs of disstria which are
Imost always parallel to one another and each perpendicular to the surface
of ‘he twig (Fig. 1). This slanting effect increases the distance of the second
row of cggs from the edge of the spumaline layer and should result in a
lower parasitism rate of the interior eggs. The second and third row para-
sitism rates cited by Hodson (1939) for disstria are indeed greater than those
we have found for americanum (Fig. 2).

VOLUME 84, NUMBER 3 455

% of Parasites

10 20 30 40 50 60 70 80 90 100

| Row

Fig. 2. Distribution of parasites by row. Malacosoma americanum (@) [present study, n =
10 egg masses] and M. disstria (4) [from Hodson, 1939, n = 52 egg masses].

_ The egg masses of M. americanum generally contain many more eggs
/(mean = 259, SD = 76.8, n = 13 egg masses) than those of either disstria
|(mean = 155 eggs; Hodson, 1941) or neustria (140-200 eggs: Schwenke,
1978). The clasping mode of oviposition may be necessary if the female is
to carry and lay a large number of eggs. A large egg mass, in turn, may be
needed to provide a large number of caterpillars, either for the construction
of large permanent tents (lacking in disstria), or for communal foraging.
Finally, a clasping egg mass may enable the female to oviposit on large
diameter twigs or even low on tree trunks normally covered by winter
snows. The latter is apparently common in the northern Great Lakes region,
but is very rare in New York State. A comparison of the low temperature
survival rates of pharate larvae of americanum and disstria may, in fact,
provide the key for understanding the evolution of this egg mass shape.

| There are interesting differences in the relative abundances of the para-
| sitoid species of M. americanum and disstria. Species of Tetrastichus,
| Ooencyrtus, and Telenomus are the most abundant egg parasitoids of both
species. In our study of M. americanum 80% of the parasitoids were Te-
trastichus malacosomae, and over 70% of these individuals were reared
from marginal eggs (Table 3). In M. disstria, parasitism is primarily due to

456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ooencyrtus clisiocampae and Telenomus clisiocampae which occur in equal
numbers and account for 80% of the total parasitism with only 20% mortality
due to Tetrastichus silvaticus Gahan (Hodson, 1939). In M. americanum,
Ooencyrtus clisiocampae and Telenomus clisiocampae show no preference
for the marginal egg class (Table 3). The more even distribution of parasit-
ized eggs in M. disstria may reflect the relative importance of these two
parasitoids. The predominance of 7. malacosomae in M. americanum may
be a reflection of the preference for marginal eggs by this species and the
greater numbers in this egg class due to the clasping mode of oviposition.

All species of Malacosoma deposit a glandular secretion with each egg
(Stehr and Cook, 1968). Hodson and Weinman (1945) coined the term *‘spu-
maline’’ to refer to these glandular materials, regardless of the final form.
In many species the spumaline is elaborated into a frothy material which
covers the eggs, referred to here as frothy spumaline. This type of spumaline
is found in all New World species except M. tigris.

Two hypotheses have been proposed for the adaptive significance of
frothy spumaline: (1) Reduction of egg parasitism (Hodson, 1939) and (2)
resistance to desiccation (Hodson and Weinman, 1945). The data of Hodson
and Weinman for M. disstria agree with ours for M. americanum: inviability
rates are significantly higher in exposed eggs than those covered with frothy
spumaline. However, if spumaline elaboration is primarily a response to
desiccation, then one must explain the resistance to desiccation in the arid-
land species M. tigris and in all Old World species.

The high parasitism rates in exposed eggs (27%) versus internal eggs (1%)
suggests that frothy spumaline is a deterrent to egg parasitoids. However,
it is not completely effective in eliminating parasitism in eggs along the
margin of the mass in either M. americanum or disstria. This is not a result
of a more ragged covering near the margin as was suggested by Hodson
(1939). All eggs scored as marginal in our study were completely covered
with a frothy spumaline. In both species over 90% of the parasitized eggs
were found in the outside two rows. This strongly suggests that the spu-
maline does not function as a repellent to parasitic Hymenoptera; rather it
acts as a physical barrier separating the wasps from potential hosts.

The distribution of frothy spumaline in species of Malacosoma can pro-
vide ecological insights only in the context of the phylogeny of the genus.
The phylogenetic relationships of Malacosoma have not been resolved, but
Stehr and Cook (1968) present morphological, ecological and behavioral
data that establish the following groups as monophyletic (Table 4): tigris
+ constrictum, neustria + disstria, and americanum + incur-
vum + californicum. The possible relationships among these three groups
are illustrated in Fig. 3. Alternative A is suggested by the distribution of
character 10, but we prefer to consider the relationships of these three
groups as unresolved pending detailed studies. Species with frothy spuma-

VOLUME 84, NUMBER 3 457

is) 2 QO
2 lla = tag 3 8 Ss
a = i) = = conse =
+ 2 = °° = a. = Ss € =“
=f = ce) s ca) - @ = = on =
= 3 = 2 =. = ” = a — x =.
—— > <= = i) 72) ” ” a“ ma = o fone Q
oa = = & ¢ + = = =e £ ¢£ gQ r=$
—_- — _— —_- = | et
a = oy eS a B.D Dein Sia en are 3.
° 2 ©
: _ a &
a = so
e = 2 25 §
= m = =
+ oO = = oo a
Pa or ” ” < —! OQ
TQ c ot Fa c = —
= =F = 3 3
an =) o », 3. . .

a

Fig. 3. Three possible cladograms of Malacosoma species-groups, with species with a
frothy spumaline indicated (*). Numbers refer to characters in Table 4. [See text for discussion. ]

line are indicated (*) and appear in each of the three groups. No other
members of the genus have frothy spumaline. There are two equally par-
simonious interpretations of the pattern of spumaline elaboration for all
cladograms in Fig. 3, either frothy spumaline evolved in the common ances-
tor of all seven species, and was subsequently lost in neustria and tigris, or
frothy spumaline was independently developed three times, in the ancestor
of americanum + californicum + incurvum, in disstria and in constrictum.

Either interpretation establishes that frothy spumaline is an evolutionarily
labile character, with either convergences or parallel reductions. We favor
the view that a frothy spumaline has evolved independently at least three
times for the following reasons:

1. The loss of frothy spumaline in M. neustria would result in a substantial

458

Table 4.

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Character states of Malacosoma species. Polarity determined by considering the

European species, M. franconicum Esper, castrensis (L.), alpicola (Staudinger) as the out-
group. Data derived from Stehr and Cook, 1968.

Character
oO. Ancestral Derived
l Large, permanent tents Small tents, used only during
moulting
2 Feed on many plants Restricted to oaks
3 Helical egg mass Clasping egg mass
4 Upper posterior corner of ovipositor Upper posterior corner of ovipositor
valves not produced, rounded or valves produced as flattened
angulate in lateral view lobes, strongly angled in lateral
view
5 Upper posterior corner of ovipositor Upper posterior corner of ovipositor
valve rounded or only slightly valve strongly produced
produced
6 Male epiphysis of foreleg small or Male epiphysis large and sickle-
absent shaped
7 Larval setal group L2 with 3 setae Setal group L2 with 2 setae
8 Larval setal group D1 with 5 setae Setal group D1 with 4 setae
9 Large, permanent tents Tent absent
10 Rear margin of male 7th sternum Rear margin of male 7th sternum

sinuous, smooth or slightly
denticulate

strongly and coarsely serrate

increase in mortality due to parasitism, from the 5—10% loss in species with
frothy spumaline (7.0% in this study) to the level of 20-40% observable
today (Tadic, 1965; Romanova and Lozinskij, 1958). We can conceive of no
selective advantage to the elimination of the spumaline coating that could
offset such an increase in egg mortality.

2. The evolution of the clasping egg mass in the americanum group pre-
sents a dilemna only if frothy spumaline occurred either simultaneously or
before the change in egg mass shape. No selective disadvantage due to
marginal egg parasitism would obtain in shifting from a helical to the clasping
egg mass if spumaline was not present; all eggs are equally vulnerable. For
the clasping egg mass to precede spumaline elaboration in the americanum
group requires that spumaline evolved convergently at least three times.

3. }rothy spumaline requires only a modification of existing accessory
gland secretions, primarily by the incorporation of bubbles. It is noteworthy
that the accessory glands of M. tigris, which has unmodified spumaline,
produce as much material as other New World species (Stehr and Cook,
1968).

VOLUME 84, NUMBER 3 459

On the basis of these observations we feel that a frothy spumaline cov-
ering was not lost in neustria and tigris, but was evolved independently in
disstria, constrictum and the americanum species group as a mechanism to
decrease egg parasitism. Parasitoids have been linked several times to the
evolution of parental guarding of eggs by the host species (see, e.g. Eber-
hard, 1975; Odhiambo, 1959; Ralston, 1977). We expect that further studies
on the evolution of mating and oviposition behavior of hosts will continue
to demonstrate the influential role played by chalcidoid and proctotrupoid
egg parasitoids.

ACKNOWLEDGMENTS

We thank J. Carpenter, J. Edgerly, G. C. Eickwort, and J. E. Rawlins
for comments on the manuscript; J. E. Rawlins for evaluating morphological
similarities of Malacosoma species and for many discussions of the phy-
logeny of the genus; and Roslyn Darling for preparation of the manuscript.

LITERATURE CITED

Anderson, J. F. 1976. Egg parasitoids of forest defoliating Lepidoptera. Pp. 233-249. Jn An-
derson, J. F. and H. K. Kaya, eds., Perspectives in Forest Entomology, Academic Press,
New York.

Eberhard, W. G. 1975. The ecology and behavior of a subsocial pentatomid bug and two
scelionid wasps: Strategy and counterstrategy in a host and its parasites. Smithson.
Contrib. Zool. 205, 39 pp.

Hodson, A. C. 1939. Biological notes on the egg parasites of Malacosoma disstria Hbn. Ann.
Entomol. Soc. Am. 32: 131-136.

——. 1941. An ecological study of the forest tent caterpillar, Malacosoma disstria Hbn., in
Northern Minnesota. Minn. Agric. Exp. Stn. Tech. Bull. 148, 55 pp.

Hodson, A. C. and C. J. Weinman. 1945. Factors affecting recovery from diapause and hatch-
ing of eggs of the forest tent caterpillar Malacosoma disstria Hbn. Minn. Agric. Exp.
Stn. Tech. Bull. 170, 31 pp.

Liu, C. L. 1926. On some factors of natural control of the eastern tent caterpillar (Malacosoma
americana Harris), with notes on the biology of the host. Ph.D. thesis, Cornell Uni-
versity, Ithaca, New York.

Odhiambo, T. R. 1959. An account of parental care in Rhinocoris albopilosus Signoret (He-
miptera—Heteroptera: Reduviidae), with notes on its life history. Proc. R. Entomol.
Soc. Lond. (A) 34: 175-185.

Prentice, R. M. 1953. Egg parasitism of forest tent caterpillar. Can. Dept. For. Bi-mon. Prog.
Rep. 9(4): 2.

Ralston, J. S. 1977. Egg guarding by male assassin bugs of the genus Zelus (Hemiptera:
Reduviidae). Psyche (Camb. Mass.) 84: 103-107.

Romanova, Y. S. and V. A. Lozinskij. 1958. Experiments on the practical use of egg parasites
of Malacosoma neustria in forest conditions. Zool. Zh. 37: 542-547.

Schwenke, W. 1978. Familienreihe Bombycoidea. Pp. 421-445. In Schwenke, W., ed., Die
Forstschadlinge Europas, Vol. 3. Schmetterlinge.

Stehr, F. W. and E. F. Cook. 1968. A revision of the genus Malacosoma Hubner in North
America (Lepidoptera: Lasiocampidae): systematics, biology, immatures, and parasites.
U.S. Natl. Mus. Bull. 276, 321 pp.

Tadi¢, M. D. 1965. Telenomus terebrans Ratz., parasite des oeufs du bombyx a livree (Ma-

460 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lacosoma neustria L.). Pp. 568-569. In Freeman, P., ed., Proc. XIIth Int. Congr. Ento-
mol., Royal Entomological Society, London.
Wellington, W. G. 1965. Some maternal influences on progeny quality in the western tent
caterpillar, Malacosoma pluviale (Dyar). Can. Entomol. 97: 1-14.
Witter. J. A. and H. M. Kulman. 1972. A review of the parasites and predators of tent
caterpillars (Malacosoma spp.) in North America. Minn. Agric. Exp. Stn. Bull. 289, 48
Pp-
. 1979. The parasite complex of the forest tent caterpillar in northern Minnesota. En-
viron. Entomol. 8: 723-731.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 461-467

THE BIONOMICS OF TWO SPECIES OF HYDROCHARA
(COLEOPTERA: HYDROPHILIDAE) WITH
DESCRIPTIONS OF THEIR LARVAE

JAMES F. MATTA

Department of Biological Sciences, Old Dominion University, Norfolk,
Virginia 23508.

Abstract.—The habitat, periods of larval and adult occurrence, and pos-
sible larval food sources are described for two species of Hydrochara. Lar-
vae and adults of Hydrochara occulta (d’Orchymont) were found to inhabit
Spartina patens (Ait.) Muhl. salt marshes, while Hydrochara soror Smetana
larvae and adults were found in a variety of non-salt marsh habitats. In
addition the larvae of the two species are described.

In his world revision of the Hydrochara, Smetana (1980) states that the
larvae of only three species (Hydrochara caraboides (Linnaeus), H. affinis
(Sharp), and H. obtusata (d’Orchymont)) are known. Of these only one, H.
obtusata, is found in the Nearctic Region. The larva of this species has been
discussed by several individuals (Bowditch, 1884; Wickham, 1895; Rich-
mond, 1920; Wilson, 1923; Bertrand, 1972), and Richmond (1920) presented
an extensive discussion of larval life history for H. obtusata.

During recent studies on the aquatic insects of Parramore Island, a barrier
island on the Virginia eastern shore, two species of Hydrochara were found
to maintain breeding populations. The larvae of these two species (H. oc-
culta (d’Orchymont) and Hydrochara soror Smetana) and their habitats are
described.

The larvae of the two species were not reared; however, larvae were
collected in association with adults, and there appears to be little or no
‘cross contamination’ of habitat by these two species. Adults were never
collected outside of their preferred habitat on Parramore Island, and H.
occulta was not collected from the Dismal Swamp. While rearing is the
preferred method of associating larvae with adults, the habitat distinctions
appear to be clear-cut enough in this case to warrant the association.

Hydrochara soror Smetana

BIONOMICS

Larvae and adults of this species were found in freshwater woodland
pools and were not as abundant on the island as H. occulta. Hydrochara

462 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

soror was also found in abundance in other woodland habitats in the tide-
water area, and larvae and adults of this species have been collected rou-
tinely from open, woodland pools in the Dismal Swamp, Virginia.

Adults have been collected during all seasons of the year, but they were
most abundant from late spring to fall. First-instar larvae have been col-
lected as early as mid-April and mature third-instar larvae have been col-
lected as late as mid-July. Adults are most abundant in heavily shaded
woodland pools and backwaters, and larvae have been collected under these
conditions on Parramore Island. In the Dismal Swamp, larvae are much
more abundant in large, open temporary pools which form in relatively
unshaded areas (old field or burn areas). There the larvae are usually as-
sociated with clumps of grass or other vegetation with stems hanging in the
water. The rarity of the larvae in woodland pools may be an artifact of
sampling since it is very difficult to separate larvae from detritus which is
invariably associated with woodland pool habitats.

On Parramore Island the woodland pools contain few potential food
species of macroinvertebrates other than Gammarus sp., Anopheles sp.,
Chironomidae, and larvae of several aquatic Coleoptera including Hydro-
porus signatus youngi Gordon, Agabus punctatus Melsheimer, and Hel-
odidae. In the Dismal Swamp a wide variety of macroinvertebrates and
amphibian larvae is available as potential prey, including the larvae of over
20 species of aquatic Coleoptera.

LARVAL DESCRIPTION

Last-instar larva.—Overall length 25.0 mm, greatest width of pronotum
3.5 mm. Body, except head and prothorax, dirty brown, slightly lighter
beneath, especially on thoracic and Ist abdominal segments.

Head as in Fig. |. Greatest width of head capsule 3.4 mm. Length 2.3 mm
from labroclypeus to occipital foramen; almost rectangular, slightly tapering
from base of mandibles to posterior edge. Labroclypeus slightly asymmet-
rical, right side with teeth projecting slightly anteriorly. Side of labroclypeus
with 2 small teeth (not counting tooth at apex). Teeth partially fused in some
specimens, particularly on right side, and rarely labroclypeus appears al-
most semicircular with no distinct teeth. Apex of labroclypeus broadly flat-
tened, slightly concave and indistinctly crenulate, with a slight projection
at each end which might be interpreted as a 3rd tooth. Anterolateral pro-
‘ections of epistoma slightly produced, not attaining level of anterior edge
of |abroclypeus; with 4 or 5 indistinct crenulations.

Oceliar area with 6 ocelli arranged in an ellipse with anterior ocellus larger
and subtriangular. Antenna with Ist segment almost 4x as long as 2nd and
3rd together. First antennal segment with inner margin slightly concave;
with about 50 strong setae on inner margin which vary in length from %4 to
134 the width of Ist antennal segment. Outer margin with about 20 irregularly

463

VOLUME 84, NUMBER 3

Head (dorsal view) of Hydrochara soror. Line indicates 1.0 mm.

Fig. 1.

464 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

placed, weak setae. Second segment narrower than Ist, slightly curved,
with a sensory pit on outer margin at midpoint and a light seta (which is
almost as long as last antennal segment) on inner margin at apex. Third
segment about half as long as 2nd and about half as wide, with 2 setae on
inner margin at apex.

Mandibles large, long and tapering, roughly symmetrical, each with a
large median tooth and a smaller, blunt submedian tooth. Teeth of left man-
dible slightly broader and thicker than right.

Maxillary stipe long and slender, almost as long as mandible, with several
irregularly placed weak setae. Palpifer much shorter, less than '/s as long
as stipes, with | or 2 setae on outer edge anteriorly and a short appendage
on inner edge anteriorly. Appendage with a long terminal bristle. Palpus 3
segmented, with Ist segment slightly more than half as long as 2nd. Third
segment slightly longer than 2nd, tapering at distal % to a rounded tip, with
a single seta located basally.

Labrum projecting beyond tip of median mandibular tooth. Mentum (in-
cluding ligula) and submentum about equal in length. Submentum (best
viewed dorsally) roughly pentagonal, with 2 sharp strong denticles distally
flanking base of mentum and with 4 smaller denticles on sloping sides. The
anterior most of these 4 denticles is extremely minute. Mentum subquad-
rangular, with ligula produced, 2x as long as Ist palpal segment. Labial
palpi 2 segmented, Ist segment short, 2nd segment long and slender, 4X as
long as Ist.

Pronotum broader than long, sloping laterally, broadly margined, with
Sagittal line prominent. Sclerotized portion of mesonotum and metanotum
reduced to subtriangular sclerites, broader anteriorly, tapering posteriorly
with a small transverse sclerite at tip of each triangle. Metanotal sclerites
smaller than mesonotal sclerites. Prosternal sclerite large. Mesosternum and
metasternum without sclerites.

Legs of similar morphology, 4 segmented, widely separated. Coxae, tro-
chanters, femora, tibiotarsi and claws 1.6, 0.6, 2.0, 1.3, and 0.5 mm in length
respectively. Coxae, trochanters and femora with a fringe of fine hairs on
upper and lower surfaces. Tibiotarsi with a fringe of stouter hairs on lower
surface and with a single large seta on upper inner face at basal % and
1 stout seta at apex. Claws single, gently curved with 2 stout setae in ba-
sal 2.

Abdomen with 8 distinct segments. Segments 9 and 10 reduced. Segment
| with a pair of small sclerites dorsolaterally at anterior margin. Segments
|—~7 secondarily divided by transverse folds; 2nd fold of each segment bear-
ing 4 setose tubercles. Epipleurite and hypopleurite of each segment (1-7)
with a short setose tubercle and the epipleurite with a long setose filament.

Eighth tergum, representing the dorsal valve of stigmatic atrium, bears a

VOLUME 84, NUMBER 3

ar
Fig. 2. Head (dorsal view) of Hydrochara occulta. Line indicates 1.0 mm.
quadrate sclerotized plate which is usually entire but occasionally projects
weakly distally and is rarely notched posteromedially so that rear margin

appears bidentate; posterior margin of stigmatic atrium with 6 distinct lobes.
Mesocercus prominent and sclerotized, bearing 2 long and | short setae;

466 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

paracerci less distinctly sclerotized, bearing 2 terminal setae. Posteroventral
gill-like appendages as long as the stigmatic atrium (ventromedially).

Hydrochara occulta (d’Orchymont)
BIONOMICS

Adults of this species were found in the low salinity (2 to 12 ppt) Spartina
patens (Ait.) Muhl. marshes on Parramore Island. They were most fre-
quently encountered in deep (1% to | m), detritus filled sloughs which were
heavily shaded with shrubs. Adults were collected during all seasons of the
year but were most abundant during late summer and fall.

Larvae were also found in the Spartina patens marshes and were usually
found clinging to the mat of S. patens stems which hang in the water at the
margins of the pools. First- and second-instar larvae were collected in late
May and mature third-instar larvae were collected as late as mid-July. In
the laboratory larvae fed on anything small enough for them to subdue,
including mosquito larvae, damselfly larvae, isopods, amphipods, small fish,
and amphibian larvae. They rejected hard bodied insects such as adult bee-
tles and some hemipterans. In the field, food available included Gammarus
sp. and small fish (Fundulus heteroclitus (Linnaeus) and Cyprinodon var-
iegatus Lacépede) as well as a variety of smaller crustaceans and insects.
Other aquatic Coleoptera found in association with this species were the
Dytiscidae, Hygrotus impressopunctatus (Schaller), Liodessus affinis (Say),
and Thermonectus basillaris (Harris), and the Hydrophilidae Tropisternus
quadristriatus Horn, Enochrus hamiltoni (Horn), and Berosus fraternus
LeConte. In addition Helodidae larvae were also abundant.

LARVAL DESCRIPTION

Last-instar larva.—Similar to H. soror; differing in shape of labroclypeus
(Fig. 2) which has a large distinct tooth on side and anterior margin ap-
pearing tridentate with middle tooth reduced. Gill-like posteroventral ap-
pendages of stigmatic atrium reduced, shorter than ventromedian length of
stigmatic atrium. In addition H. occulta usually are lighter than H. soror,
being straw yellow rather than brown.

ACKNOWLEDGMENTS

The author thanks The Virginia Environmental Endowment (grant 78-02;
An "Ecological Study of Parramore Island) and the Barrier Islands Program
of Old Dominion University for support. Thanks are also due to James
Cowan, Bill Sutphin, and Harold Marshall for their help in obtaining spec-
imens.

VOLUME 84, NUMBER 3 467

LITERATURE CITED

Bertrand, H. 1972. Larves et nymphes des coléopteres aquatiques du globe. Paris. 804 pp.

Bowditch, F. C. 1884. Hydrocharis obtusatus. J. Boston Zool. Soc. 3: 1-6.

Richmond, E. A. 1920. Studies on the biology of the aquatic Hydrophilidae. Bull. Am. Mus.
Nat. Hist. 42: 1-94.

Smetana, A. 1980. Revision of the genus Hydrochara Berth. (Coleoptera: Hydrophilidae).
Mem. Entomol. Soc. Can. 111: 1-100.

Wickham, H. F. 1895. On the larvae of Hydrocharis obtusatus and Silpha surinamensis.
Entomol. News 6: 168-171.

Wilson, C. B. 1923. Life history of the water scavenger beetle Hydrous (Hydrophilus) trian-
gularis, and its economic relation to fish breeding. Bull. U.S. Bur. Fisheries 39: 9-38.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 468-474

SMITTIA LASIOPS (MALLOCH): A REDESCRIPTION OF THE
ADULTS WITH A DESCRIPTION OF THE IMMATURE STAGES
(DIPTERA: CHIRONOMIDAE)

DONALD W. WEBB

Section of Faunistic Surveys and Insect Identification, Illinois Natural
History Survey, Champaign, Illinois 61820.

Abstract.—The adults of Smittia lasiops (Malloch) are redescribed, with
a description of the immature stages. The terrestrial nature of the immature
stages is discussed.

Malloch (1915) described the adults of Smittia lasiops (as Camptocladius
lasiops) from adults collected around garbage at a home in Urbana, Illinois.
The place of occurrence of the adults indicated to Malloch that the larva
was terrestrial. During an investigation of the bionomics of the northern and
western corn rootworms by the Section of Economic Entomology, Illinois
Natural History Survey, numerous larvae of a terrestrial chironomid were
collected from soil samples taken in an experimental corn plot on the Uni-
versity of Illinois South Farm area at the south edge of Urbana. Soil samples
collected for egg counts of corn rootworms had been washed and separated
using an apparatus (Shaw et al., 1976) that would process a pint of soil in
34 minutes. Using this apparatus, I recovered numerous live chironomid
larvae from subsequent soil samples taken in the experimental corn plots.
Larvae were also collected on the roots of corn plants, although there was
no evidence of damage to the roots. Several larvae were reared, and I
identified the adults as Smittia lasiops.

Malloch’s description of the male and female of Smittia lasiops was very
general. Sublette (1967, 1970) added to the description of Smittia lasiops in
his review of the type-specimens of the Chironomidae in the collections of
the Illinois Natural History Survey and the Canadian National Collection.
In his examination of the lectotype male (Frison, 1927), lectoallotype, and
paratypes at the Illinois Natural History Survey, Sublette mounted only the
male terminalia on slides, and based the remainder of his description on
pinned material. In my examination of slide material of the reared adults
and a slide preparation of the lectotype, I found some variation from Sub-
lette’s description, in addition to certain undescribed features of the adults.

VOLUME 84, NUMBER 3 469

With the discovery of the immature stages, the determination of their
terrestrial nature, and because of the incomplete earlier description of the
adults, I feel it is appropriate to redescribe the adults and to describe the
immature stages. No comparison is made at this time between other eastern
species of Smittia.

The terminology and abbreviations follow that of Saether (1980). All mea-
surements are given in millimeters, with the range and mean.

Smittia lasiops (Malloch)

Male.—Length 2.17-2.51, 2.41 + 0.10 (N = 10). Head (Fig. 1) dark
brown, expanded laterally. Eyes dark brown; dichoptic; facets of equal size;
covered with short fine setae; dorsal separation between eyes 1.63 x ventral
distance. Coronal triangle reduced, narrowed anteriorly, |.1—1.7, 1.4 long-
er than wide; coronal setae absent. Antenna dark brown; scape flattened,
ring-shaped, 1.2—1.4, 1.3 longer than wide; pedicel globose, macrosetae
absent, 1.4—2.2, 1.9 wider than long; flagellomeres plumose, flagellomeres

2,3, and 13 (Fig. 2) with 2 flat hyaline lateral blades, flagellomere 13 with
_ distinct apical seta, flagellomere lengths: 0.048-0.054, 0.052; 0.018-0.024,

0.021; 0.021-0.027, 0.024; 0.018-0.024, 0.021; 0.018-0.024, 0.020; 0.018-
0.024, 0.020; 0.018-0.024, 0.020; 0.018-0.024, 0.020; 0.018—-0.024, 0.021;
0.021-0.024, 0.022; 0.018-0.024, 0.022; 0.021-0.024, 0.022; 0.540-0.600,

0.578; AR 1.87-2.16, 2.03. Clypeus dark brown, broad, ventral 43 tapered

to broad point; clypeal setae 6-8, 7, dark brown, elongate. Maxillary palps
membranous, pale brown, cylindrical, length of palpal segments: 0.016—
0.030, 0.023; 0.037—0.051, 0.043; 0.076-0.092, 0.085; 0.069-0.092, 0.097;
0.085-0.099, 0.091; setae dark brown, elongate, scattered. Inner vertical
setae 0-2, 1; outer vertical setae 0-4, 2; frontal setae absent; postorbital
setae 0-3, 1. Cibarial pump (Fig. 3) elongate, tapered ventrally, apex acute,
0.145-0.170, 0.159 long, 0.048-0.074, 0.060 wide, 2.2-3.2, 2.7 longer than
wide. Tentorium (Fig. 3) 0.127—0.150, 0.141 long.

Thorax dark brown, subshiny; vittae indistinct, Antepronotum in lateral
view narrow, elongate, expanded ventrally; in dorsal view median fissure
contiguous; antepronotal setae absent. Acrostichial setae absent; dorsocen-
trals 5-12, 8; prealar setae 2-6, 4. Pleura pale brown, anepisternal and Ka-
tepisternal setae absent. Halter pale yellowish brown to pale brown. Scu-
tellum dark brown, subshiny, rounded posteriorly; scutellar setae 6—7, 6 in
single transverse row.

Wing (Fig. 4) length (from arculus) 1.47—1.79, 1.68; 3.6 longer than wide.
Membrane hyaline; microtrichia absent; brachiolar setae 1-2, 1; basal half
of radius with 4-6, 5 setae; R, with | seta; veins pale yellowish brown.
Costa ending anterior to apex of wing, distinctly distad to apex of Ry,;. Ro+s
reaches costa 0.49-0.55, 0.53 of way between R, and R,,;. Squama glabrous.
OR 1.2—1.3, 1.3.

470 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-7. Smittia lasiops. 1, Head. 2, Male flagellomeres 1-3, apex of flagellomere 13. 3,
Cibarial pump and tentorium. 4, Male wing. 5, Male terminalia. 6, Female antenna. 7, Female
terminalia.

VOLUME 84, NUMBER 3 47]

Legs pale brown, concolor, linear. Tibial spur on foreleg 0.046-0.051,
0.048. Inner tibial spur on midleg 0.012—0.018, 0.015, outer tibial spur 0.018—
0.028, 0.022. Inner tibial spur on hindleg 0.009-0.023, 0.015, outer tibial
spur 0.041—0.054, 0.048. Empodium filiform, pale yellow. Pulvilli reduced,
indistinct. Apical claws on tarsomere 5 fuscus, paired, simple. Lengths (in
mm) and proportions of legs:

fe tl ta,
|e 0.536-0.643, 0.604 0.673—0.826, 0.760 0).352-0.444, 0.393
P, 0.612-—0.689, 0.656 0.536—-0.719, 0.635 0.245—0.321, 0.292
Py 0).627-0.719, 0.689 0.673-0.796, 0.736 0.352-—0.459, 0.424
tay ta3 ta,
Pe 0.199-0.245, 0.226 0.138—0.168, 0.158 0.077—0.107, 0.096
|e 0.153—0.184, 0.166 0.107—0.138, 0.119 0.061-0.077, 0.075
P; 0.1840.245, 0.226 0. 168—0.184, 0.177 0.077—0.092, 0.090
ta; LR BV
Leg 0).046—0.092, 0.081 0.46—-0.57, 0.52 2.90-4.04, 3.23
ae 0.061—0.077, 0.073 0).40-0.54, 0.46 3.62—3.74, 3.65
P, 0.077—0.092, 0.086 O352—0:62, 0:57 3.24-3.41, 3.26
SV
Pi 3913-381, 3-48
PS 4.02-4.90, 4.40
re 521-370, 3:40

Abdomen and terminalia (Fig. 5) dark brown; setae dark brown to fuscus,
elongate, scattered. Anal point distinct, acute. Gonostylus broad, reflexed,
apex truncate, length 0.08—0.10, 0.095; apical tooth thick, length 0.01. HR
N7=1.95 1.75:

Female.—Coronal triangle indistinct. Antenna (Fig. 6); scape 0.06—0.10,
0.08 long, 0.05-0.07, 0.06 wide, 1.1-1.6, 1.3 longer than wide; pedicel
0.06—-0.07, 0.062 long, 0.07—0.08, 0.072 wide, 1.0—1.3, 1.2 wider than long;
flagellomeres pilose, flagellomeres I—5 with 2 broad hyaline blades, lengths
0.078-0.090, 0.084, flagellomere lengths: 0.066—0.072, 0.067; 0.048—0.060,
0.053; 0.060—-0.072, 0.062; 0.060-0.072, 0.062; 0.090-0.114, 0.097. Clypeal
setae 6-13, 8. Maxillary palpal lengths: 0.018—0.024, 0.023; 0.024—0.042,
0.032; 0.060-0.078, 0.067; 0.072—0.084, 0.076; 0.066—-0.096, 0.079; setae pale
brown. Inner vertical setae 2; outer vertical setae 1-2, 2; postorbital setae
absent. Cibarial pump 0.108—0. 162, 0.140 long, 0.054—0.066, 0.060 wide, 1.8-
2.6, 2.3x longer than wide. Tentorium 0.108—0.138, 0.128 long.

472 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Antepronotal setae 3-4, 4; dorsocentrals 9-12, 11; prealars 3-4, 4; scu-
tellar setae 5—7, 6.

Wing length 1.42—1.64, 1.53, 2.7-3.7, 2.9x longer than wide. Brachiolar
setae 2; setae on basal half of radius 10-13, 11; setae on R, 7-8, 8; setae on
R,,; 13-18, 16. R,,; lies alongside of R,, almost fused. VR 1.2-1.3, 1.26.

Length of tibial spur on foreleg 0.018—0.037, 0.029; length of inner spur
on midleg 0.009-0.018, 0.014, outer spur 0.018—0.025, 0.022; length of inner
spur on hindleg 0.012—0.016, 0.014, outer spur 0.051—0.046, 0.045. Lengths
(in mm) and proportions of legs:

Fe ae Ta,

et 0.413-0.536, 0.477 0.536—0.658, 0.594 0.260-0.306, 0.288

Es 0.4740.612, 0.536 0.52020-627,, 0.557 0.214—0.275, 0.236

EB 0.58 1-0.643, 0.600 0.551—0.719, 0.624 0.321-0.428, 0.361
Ta; Tas Ta,

jet 0.122—0.184, 0.165 0.107—0.199, 0.135 0.077—0.077, 0.077

PS 0.107-0. 138, 0.125 0.092-0. 168, 0.113 0.061—0.077, 0.064

P; 0.153-0.199, 0.174 0. 138-0. 168, 0.147 0.061—0.077, 0.071
Ta; LR BV

E 0.061—0.077, 0.064 0.40.5, 0.48 2.6—3.3, 3.1

P, 0.046—0.061, 0.058 0.40.4, 0.40 3.0-4.0, 3.6

P; 0.061—0.077, 0.071 0.60.6, 0.60 3.24.02. 3.5
SV

|e 3.53937

ES 4.44.8, 4.6

lee 3.23.5, 3.4

Abdomen and terminalia (Fig. 7) pale brown. gonopophysis VIII mem-
branous, oblong. Gonocoxite LX with 4 setae. Rami of gonapophysis IX
0.207—0.248, 0.228 long. Cerci reduced, length 0.048-0.060, 0.054. Sper-
matheca oval, dark brown, length 0.081, width 0.044—-0.048, 0.046, 1.8x
longer than wide.

Pupa.—Length 2.4-3.0, 2.7 (N = 3). Exuvium pale yellow. Thoracic horn
and cephalic tubercle absent. Precorneal setae 1-3 in short arc. Median
antepronotal setae |. Dorsocentral, metanotal, prealar, supraalar, postor-
bital, and vertical setae absent. Wing sheath broad, apex rounded, nose and
pearl row absent. Tergum I with fine spinules scattered across anterior half.
Terga II—-VII1 with fine spinules across each entire tergum, generally coarser
anteriorly on each tergum. Lateral and dorsal setae absent. Pedes spurii B

VOLUME 84, NUMBER 3 473

Figs. 8-12. Smittia lasiops, larva. 8, Antenna. 9, Mentum. 10, Mandible. 11, Premandible.
12, Abdominal segments 7-9.

and posterolateral tubercle on tergum VIII absent. Anal lobe without mac-
rosetae or fringe of natatory setae.

Larva.—Head capsule dark yellow, body pale white. Length 4.24.6,
4.3 + 0.07 (N = 5). Head capsule length 0.306—0.360, 0.322, width 0.246—

474 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

0.276, 0.263, 1.2 longer than wide. Postoccipital margin broadly rounded
anteriorly, extending anteriorly 0.108—0.138, 0.121 from posterior margin of
head. Antenna (Fig. 8), length of segments: 0.028—0.035, 0.031; 0.037—0.041,
0.039; 0.001; 0.002, AR = 0.74; width of basal segment 0.018, 1.6—1.9, 1.7
longer than wide, 0.7—0.9, 0.8 length of 2nd segment; distance from base
to annular organ 0.012—0.018, 0.015; length of antennal blade 0.032-0.044,
0.039; lauterborn organs reduced at apex of 2nd segment, extend to apex of
antennal segment 4. Mentum (Fig. 9), length 0.053-0.067, 0.061, width
0.104—0.113, 0.107, 1.6-2.0, 1.8 wider than long; anterior convex, median
tooth broad, simple, 5 pairs of lateral teeth, descending in height from me-
dian tooth. Ventromental plate and setae absent. Mandible (Fig. 10) length
0.104—0.110, 0.107; width 0.053-0.064, 0.057; 1.7-2.0, 1.9x longer than
wide; outer lateral broadly rounded, not crenulate; mediolateral margin
smooth; apical tooth 0.5—0.6, 0.55 combined width of lateral teeth; setae
subdentalis indistinct; setae interna reduced, branched; lateral setae 2. Pre-
mandible (Fig. 11) bifid, length 0.053—0.064, 0.059; inner blade 1.0—1.5, 1.3 x
basal width of outer blade; premandibular brush absent. Setulae I dentate.
Fore prolegs reduced, fused medially, claws simple. Procerci and posterior
prolegs absent (Fig. 12). Anal gills reduced, 2 pairs, rounded apically. Pos-
terior margin of abdominal segment 9 (Fig. 12) with numerous coarse, simple
spinules.

LITERATURE CITED

Frison, T. H. 1927. A list of the insect types in the collections of the Illinois State Natural
History Survey and the University of Illinois. Ill. Nat. Hist. Surv. Bull. 16: 137-309.

Malloch, J. R. 1915. The Chironomidae, or midges, of Illinois, with particular reference to the
species occurring in the Illinois River. Ill. State Lab. Nat. Hist. Bull. 10: 275-543.

Saether, O. A. 1980. Glossary of chironomid morphology terminology. (Diptera: Chironomi-
dae). Entomol. Scand. Suppl. 14: 1-51.

Shaw, J. T., R. O. Ellis, and W. H. Luckmann. 1976. Apparatus and procedure for extracting
corn rootworm eggs from soil. Ill. Nat. Hist. Surv. Biol. Notes 96: 1-4.

Sublette, J. E. 1967. Type specimens of Chironomidae (Diptera) in the Canadian National

Collections, Ottawa. J. Kans. Entomol. Soc. 40: 290-331.

. 1970. Type specimens of Chironomidae (Diptera) in the Illinois Natural History Survey

collection, Urbana. J. Kans. Entomol. Soc. 43: 44-95.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 475-479

A REVIEW OF THE GENUS CHAETOCOELUS LeCONTE
(COLEOPTERA: MALACHIIDAE)

ADRIEAN J. MAYOR

Department of Entomology, University of California, Riverside, California
92521.

Abstract.—Chaetocoelus LeConte and C. setosus LeConte are rede-
scribed. Chaetocoelus is characterized by the presence in the male of a
pectinate comb of stout, black teeth on the anterior margin of segment I of
the protarsus. This character, not mentioned in the original description,
places the genus in the subfamily Carphurinae. Chaetocoelus is closely re-
lated to Carphuroides Champion from which it differs primarily by the great-
ly reduced female elytra, a character of dubious generic value. Formal syn-
onymy is not currently recommended; however, C. setosus should be
considered in any revision of the genus Carphuroides. A lectotype is des-
ignated for C. setosus.

The monotypic genus Chaetocoelus was described by LeConte in 1880.
The genus was briefly characterized by the brachyelytrous female, and the
presence in both sexes of long, stiff bristles at the sides of the abdomen.
Marshall (1954), relying on LeConte’s description, placed Chaetocoelus in
the subfamily Malachiinae. Recently I was able to examine the type-series
of Chaetocoelus setosus and discovered several characters not mentioned
by LeConte in his original description. The most important of these, the
presence of a pectinate comb of stout, black teeth on the anterior margin
of segment I of the male protarsus, suggests that Chaetocoelus belongs in
the subfamily Carphurinae. Redescriptions of Chaetocoelus and C. setosus
and a discussion of the taxonomic placement of the genus are given below.

Genus Chaetocoelus LeConte

Chaetocoelus LeConte, 1880: 194 [Type-species: Chaetocoelus setosus
LeConte; fixed by monotypy]; LeConte and Horn, 1883: 214; Marshall,
1948: 125, 1954: 75; Wittmer, 1961: 612.

Description.—Small, graciliform. Surface shining. Yellow brown to red-
dish brown. Pubescence conspicuous; long erect setae on head, pronotum,
elytra, and abdomen.

Head short, frons slightly impressed. Clypeus short, appearing entirely

476 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

membranous; epistomal suture tangential to antennal sockets. Antenna I1-
segmented, submoniliform to subserrate.

Pronotum quadrate, as wide as head, apex not produced; anterior angles
variable; sides slightly and evenly produced; posterior angles broadly round-
ed, not reflexed; disc smooth, lacking sulci or other impressions. Elytra
varying in length, fully developed in male, greatly reduced in female; apex
unmodified in both sexes. Wings present, fully developed in male, reduced
to small pads in female. Tarsi 5-segmented in both sexes; segment I of male
protarsus with a pectinate comb of stout, black teeth on anterior margin;
segments II-IV of both sexes narrow, rounded at apex, not bilobed.

Abdominal sterna completely sclerotized medially; terminal abdominal
sternum broadly rounded in both sexes; pygidium truncate in male, broadly
rounded in female. Aedeagus of male genitalia with median lobe elongate,
bluntly bilobed apically (dorsal aspect); internal sac armed with large acute
spines.

Remarks.—The family Malachiidae has been divided into two subfamilies,
the Carphurinae and Malachiinae (Champion, 1923; Evers, 1968). The Car-
phurinae are characterized by the presence of a pectinate comb of stout,
black teeth on the anterior margin of segment I of the male protarsus, the
presence of eight small pairs of eversible vesicles (one between head and
prothorax, one on metepimeron, and one each on the lateral margins of
abdominal sterna I-VI), the presence of seven visible abdominal sterna, and
the bilobed apex of the median lobe of the aedeagus. The Malachiinae are
characterized by the absence of a comb on segment I of the male protarsus,
the presence of two large pairs of eversible vesicles (one between head and
prothorax and one between metasternum and first abdominal sternum), the
presence of six visible abdominal sterna, and the simple apex of the median
lobe of the aedeagus. The presence of a tarsal comb on segment I of the
male protarsus, a character overlooked by LeConte in his description of
Chaetocoelus, as well as concurrence of other characters mentioned above,
places the genus in the subfamily Carphurinae.

Chaetocoelus, with the exception of a specimen from Ceralbo Island in
the Gulf of California identified by Marshall (1951) as Carphuroides atra-
tulus (Gorham), is the only genus of Carphurinae known from North Amer-
ica. It appears to be most closely related to the genus Carphuroides Cham-
pion (1923). In both taxa tarsal segments HI-IV are narrow and rounded at
the apex rather than broad and more or less bilobed as they are in most
other carphurine genera, and the pronotum is unmodified, without sulci or
other impressions. Males of Chaetocoelus cannot be distinguished from
males of Carphuroides by any character or combination of characters. Fe-
males differ only in that those of Chaetocoelus are brachyelytrous and bra-
chypterous while females of Carphuroides tend to have the elytra and wings

a

VOLUME 84, NUMBER 3 477

3

Figs. 1-3. Chaetocoelus setosus. 1, Male elytron, dorsal aspect. 2, Female elytron, dorsal
aspect. 3, Male protarsus, anterior aspect.

478 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

fully developed. These characters have not often been used as a basis for
characterizing genera of Malachiidae, and in some (Endeodes and Attalu-
sinus) there is considerable interspecific variation with respect to the length
of the elytra and the development of the wings. This suggests that the two
taxa may be congeneric. However, Carphuroides is a large, widely distrib-
uted genus, and, until its numerous species can be studied along with Chae-
tocoelus setosus in the context of a generic revision, formal synonymy is
not recommended.

Chaetocoelus setosus LeConte
Figs. 1-3

Chaetocoelus setosus LeConte, 1880: 194 [Lectotype, ¢, herein designated
from Columbus, Texas, in the Museum of Comparative Zoology, Harvard
University, Cambridge, Mass. (Type No. 3465, examined)]; LeConte and
Horn, 1883: 214; Marshall, 1948: 125.

Description.—Length 1.8 mm, male; 0.8—1.1 mm, female (measured from
anterior margin of pronotum to apex of elytra). Pale yellow brown to dark
reddish brown; elytra yellow brown, apex darker in female. Pubescence
conspicuous; long, erect, tan to reddish-brown setae on head, pronotum,
elytra, and abdomen; lateral margins of abdomen with some setae 1.5X as
long as eye or shorter in male, and 4 to 5X as long as eye or longer in
female.

Head 1.6, male; 1.3—1.4, female, as wide as interocular distance; frons,
interocular margins, and much of vertex with finely sculptured ridges; ridges
transverse on vertex, and more or less longitudinal on interocular margins
and frons; median apex of vertex smooth, finely, sparsely punctate. Antenna
short, stout, barely surpassing elytral humeri in male, shorter in female;
segments III—X subserrate in male, submoniliform in female; VII 0.8—1.0 as
long as wide. Eyes coarsely faceted in both sexes; male eyes nearly twice
as large as female eyes.

Pronotum 0.78, male; 0.80-0.85, female, as long as wide; anterior angles
truncate in male, broadly rounded in female; lateral margins diverging from
base in male, widest at apical angles; lateral margins subparallel in female,
widest near middle. Elytra of male (Fig. 1) long, sutural margin entire,
4 abdominal terga exposed; female elytra (Fig. 2) short, sutural margin
divergent apically, 7 abdominal terga exposed.

Tarsal comb (Fig. 3) on segment I of male protarsus with 7 teeth.

Abdomen wider than elytra in female; segments II and III widest, IV to
VII tapering to apex. Aedeagus of male genitalia with a pair of long sharp
spines at apex of internal sac.

VOLUME 84, NUMBER 3 479

Geographic distribution.—Southeastern North America from Florida west
to Texas.

Specimens examined.—FLORIDA: 4 2 Punta Gorda, II.7/10.40. TEXAS:
1 36,2 2 Columbus, I-17 June.

ACKNOWLEDGMENTS

I thank Armin Coray, biological illustrator, Basel, Switzerland, for the
drawings, and John D. Pinto, University of California, Riverside, for sug-
gestions concerning the manuscript.

LITERATURE CITED

Champion, G. C. 1923. A revision of the Malayan and Indian species of the melyrid subfamily
Carphurinae. Ann. Mag. Natl. Hist. (9) 12: 1-54.

Evers, A. M. J. 1968. Carphurinae oder Carphuridae. Entomol. Bl. Biol. Syst. Kaefer 64: 17-
Die

LeConte, J. 1880. Short studies of North American Coleoptera. Trans. Am. Entomol. Soc. 8:
194.

LeConte, J. and G. Horn. 1883. Classification of the Coleoptera of North America. Smithson.
Misc. Collect. 264: 169-348.

Marshall, M. Y. 1948. Studies in the Malachiidae II. Entomol. Am. 28: 113-144.

. 1951. Studies in the Malachiidae III. Proc. Calif. Acad. Sci. 27(4): 77-132.

. 1954. A Key to the world genera of Malachiidae. Coleopt. Bull. 8(5&6): 59-82.

Wittmer, W. 1961. Malachiinae, Fasc. 67, p. 612. Jn Arnett, R. H., The Beetles of the United
States (a manual for identification). Catholic University of America Press, Washington,
DIG i2tpp:

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 480-488

SIX NEW SPECIES OF NEOTROPICAL ARADIDAE
(HEMIPTERA)

NICHOLAS A. KORMILEV

Research Associate in Entomology, Bernice P. Bishop Museum, Hono-
lulu, Hawaii; mailing address: 87-17 Myrtle Ave., Glendale, New York
11385.

Abstract.—Six new taxa are proposed, Aneurus aterrimus, n. sp. (Costa
Rica), A. brailovskyi, n. sp. (Mexico), Mezira guanacastensis, n. sp. (Costa
Rica), M. hondurensis, n. sp. (Honduras), M. proxima, n. sp. (Peru), and
Notapictinus piliger, n. sp. (Colombia).

By the kind offices of Dr. Harry Brailovsky, Instituto de Biologia, Uni-
versidad Nacional Autonoma de Mexico, Mexico 20, Mexico, and Dr. John
A. Chemsak, University of California, Berkeley, I had the privilege of study-
ing Neotropical Aradidae under their care, for which I express my sincere
gratitude.

Among these Aradidae six species were new and are described in this
paper. Of particular interest is a species of Notapictinus Usinger and Mat-
suda, 1959, whose body, antennae and legs are covered with stiff, erect
bristles, the first case of such pilosity in this genus.

All measurements in this paper were taken with micromillimeter eyepiece,
25 units = | mm. In ratios, the first figure represents the length and the
second the width of measured portion.

Subfamily ANEURINAE
Genus Aneurus Curtis, 1825
Aneurus brailovskyi Kormilev, NEw SPECIES
Figs. 1-3

Female.—Elongate ovate; head and pronotum partially granulate; looks
scabrous.

Head: Almost as long as its width across eyes (15.5:16.0); anterior process
rounded anteriorly, not reaching tip of antennal segment I; antenniferous
tubercles truncate anteriorly; postocular blunt, reaching outer border of
eyes. Eyes semiglobose, moderately protruding. Vertex transversely striate.

:

VOLUME 84, NUMBER 3 481

Antenna strong; antennal segment I barrel-shaped; II subobovate; III ta-
pering toward base; IV fusiform; relative length of antennal segments I to
IV are: 5.0:5.0:5.5:11.5. Labium reaching line connecting hind borders of
eyes.

Pronotum: Less than '% as long as its maximum width (15:35); collar
sinuate anteriorly; anterior borders rounded and produced forward beyond
collar; lateral notch slightly sinuate; lateral borders of hind lobe parallel;
hind border sinuate medially. Fore disc granulate and with 4 (2+2) curved
callosities; hind disc transversely striate and with 2 (1+1) transverse cal-
losities.

Scutellum: Semicircular, shorter than its basal width (15:23); basal 4% of
disc with rough longitudinal striation, surrounded with concentrical stria-
tion.

Hemelytra: Reaching hind border of tergum VI; corium reaching basal
¥3 of scutellum.

Abdomen: Ovate, longer than its maximum width across segment IV
(70:52); posteroexterior angles of connexiva barely protruding; paratergites
very short, not reaching tip of segment IX; the latter truncate posteriorly.
Spiracles II, VI, and VII lateral and visible from above; III to V ventral,
equidistant from border; VIII terminal. Sublateral fold reaches hind border
of sternum VI.

Posterior lobe of propleuron swollen and granulate, visible from above.

Male.—Similar to female, but smaller; paratergites small, reaching tip of
rounded, posteriorly, small hypopygium.

Measurements.—head 15.0:15.5; relative length of antennal segments I to
IV are: 5:5:5:11; pronotum 14:31; scutellum 15:20; abdomen 59:48; hypo-
pygium 3:5.

Color.—Yellow brown (¢) to dark brown (@ )

Total length.— 2? —4.68, 6 —4.20; width of pronotum: 2—1.40, ¢d—1.24;
width of abdomen: 2?—2.08, d—1.92 mm.

Holotype.—?, Mexico, Vera Cruz, Las Cabanas; 10.VII.1972, H. Brail-
ovsky leg.; deposited at the Instituto de Biologia, UNAM, Mexico 20, Mex-
ico.

Allotype—é , collected with holotype; same collection.

Paratypes.—1 2, | 3 (without head), collected with holotype; same col-
lection and Kormilev collection.

Remarks.—Aneurus brailovskyi is closely related to A. arizonensis Picchi,
1977, and runs to it in Picchi’s key, but may be separated from it by the
following: Anterior process of head not reaching tip of antennal segment I
(produced beyond tip in A. arizonensis); postocular tubercles only reaching,
or barely protruding, beyond outer border of eyes (distinctly protruding in
A. arizonensis); hind lobe of propleuron more swollen and visible from

482 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

above (less swollen and not visible from above in A. arizonensis); and ratio
between length of antennae and width of head across eyes is 1.635 (1.535
in A. arizonensis).
The holotype was compared with a paratype of A. arizonensis from the
collections of the American Museum of Natural History, New York, N.Y.
It is a pleasure to dedicate this species to its collector Dr. Harry Brailov-
sky, Instituto de Biologia, UNAM, Mexico, Mexico.

Aneurus aterrimus Kormilev, NEW SPECIES
Figs. 4-6

Male.—Elongate with subparallel sides; head, pronotum, scutellum and
connexivum partially, finely granulate.

Head: Longer than its width across eyes 16.0:15.5; anterior process
rounded anteriorly; genae short, by far not reaching tip of clypeus; almost
reaching tip of antennal segment I. Antenniferous tubercles truncate ante-
riorly, acute anterolaterally. Postocular tubercles blunt, not reaching outer
border of eyes. Eyes semiglobose, protruding. Vertex with 2 (1+1) large,
ovate infraocular callosities. Antenna strong; antennal segment I elongate
obovate, II obovate, III tapering toward base, IV fusiform; relative length
of antennal segments I to IV are: 5.5:4.5:5.0:10.0; ratio length of antenna:
width of head across eyes as 1.61:1.00. Labium reaching line connecting
hind borders of eyes.

Pronotum: Half as long as its maximum width (16:33); collar sinuate an-
teriorly; anterolateral angles rounded, neither produced forward, nor side-
ways. Lateral notch forming obtuse angle; lateral borders of hind lobe sub-
parallel, slightly rounded; hind border slightly sinuate medially. Fore disc
with 2 (1+1) larger callosities and 4 (2+2) smaller laterad of them; hind disc
finely granulate.

Scutellum: Semicircular, shorter than its basal width (15:22); disc densely
granulate on basal 4%, less densely around basal 4%.

Hemelytra: Almost reaching hind border of tergum VII; corium reaching
basal % of scutellum.

Abdomen: Elongate, with slightly rounded sides; longer than its maximum
width across segment IV (61:41); posteroexterior angles of connexiva not
protruding. Paratergites short, rounded posteriorly, reaching tip of a small
hypopygium, which is shorter than its maximum width (3.0:5.5). Spiracles
Il, VI and VII lateral and visible from above; III to V sublateral, placed on
sublateral fold; VHI terminal. Sublateral fold extends from III to hind border
of VI.

Female.—Similar to male, but larger; paratergites very short, rounded
posteriorly, reaching tip of truncate segment IX.

Measurements.—Head 16.0:15.5; relative length of antennal segments I

VOLUME 84, NUMBER 3 483

to IV are: 6:5:5:11; pronotum 19:36; scutellum 15:23; abdomen across seg-
ment IV (67:44); width of tergum VIII, 14.

Color.—Black; base of membrane dark brown. On some specimens
pronotum laterad of collar yellow brown.

Total length— ¢—4.40, 2—4.72 mm; width of pronotum: ¢—1.32,
2?—1.44 mm; width of abdomen: ¢—1.64, 2—1.76 mm.

Holotype.—d , Costa Rica, Monte Verde, Ponta Arenas; 9.II.1981, H.
Brailovsky leg., deposited at the Instituto de Biologia, UNAM, Mexico 20,
Mexico.

Allotype.—@ , Same locality and date; E. Barrera leg.; same collection.

Paratypes.—3 36, 2 2, same locality and date, H. Brailovsky leg.; 3 6,
2 2, same locality and date, E. Barrera leg. Same collection and Kormilev
collection.

Remarks.—Aneurus aterrimus runs to A. slateri Picchi, 1977, in Picchi’s
key but may be separated from it by: Head slightly longer than its width
across eyes; antennal segment III tapering toward base, not fusiform; larger
size; and different color, black.

Subfamily MEZIRINAE
Genus Notapictinus Usinger and Matsuda, 1949
Notapictinus piliger Kormiley, NEw SPECIES
Figs 7, 8

Male.—Elongate ovate; body, particularly on the borders, antennae and
legs, covered with stiff, erect bristles.

Head: Shorter than its width across eyes (21.0:22.5); anterior process
with parallel sides, rounded anteriorly, reaching basal 4 of antennal seg-
ment I. Antenniferous tubercles short, blunt, with parallel outer borders.
Postocular borders rounded and granulate. Eyes moderately convex, shorter
than preocular portion (5:7). Vertex moderately convex, granulate. Anten-
nae long and thin, more than 24% as long as width of head across eyes
(2.69:1.00); relative length of antennal segments I to IV are: 16.0:11.0:
21.0:12.5. Labium not reaching hind border of labial groove, which is open
posteriorly.

Pronotum: Less than % as long as its maximum width (27:58); collar
truncate anteriorly, granulate. Anterolateral angles rounded and produced
forward, reaching hind border of collar. Fore border between collar and
anterolateral angles sinuate and with a cluster of granules in the middle.
Lateral borders of fore lobe and hind lobe in front of humeri expanded and
crenate. Lateral borders of hind lobe roughly granulate. Hind border twice
(1+1) sinuate laterad of scutellum. Fore disc with 2 (1+1) flat callosities,
granulate between and laterad of them. Hind disc irregularly granulate.

Scutellum: Crushed by pin.

484 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-3. Aneurus brailovskyi. 1, Head, pronotum, and scutellum, 2. 2, Tip of abdomen
from above, 2.3, Tip of abdomen from above, ¢. Figs. 4-6. A. aterrimus. 4, Head, pronotum,
and scutellum, 2. 5, Tip of abdomen from above, ¢. 6, Tip of abdomen from above, ¢. Figs.
7,8. Notapictinus piliger, 6.7, Head and pronotum. 8, Tip of abdomen from above. Figs. 9-
11. Mezira hondurensis. 9, Pronotum, 3. 10, Hypopygium and paratergites, ¢. 11, Tip of
abdomen from above, 2. Fig. 12. M. proxima, 2, tip of abdomen from above. Figs. 13, 14.
M. guanacastensis. 13, Tip of abdomen from above, 3. 14, Tip of abdomen from above, °?.

Hemelytra: Reaching hind border of tergum VII; corium reaching hind
border of connexivum II; its basolateral border straight, slightly expanded
and granulate; apical angle acute and produced backward; apical border
sinuate. Membrane with anastomosed veins.

VOLUME 84, NUMBER 3 485

Abdomen: Ovate, longer than its maximum width across segment V
(90:70); posteroexterior angles of connexiva II to IV not protruding; V
slightly produced, rounded; VI more protruding, rounded; exterior border
between VI and VII sinuate; VII rounded, forming subangular lobe. Para-
tergites clavate, reaching *%4 of hypopygium; the latter pointed posteriorly
and with a stout median ridge, almost reaching tip of disc. Spiracles II to
V ventral, placed far from border; VI sublateral, but not visible from above;
VII and VIII lateral and visible from above. Sterna III to VI with swollen
hind border.

Legs: Unarmed.

Color.—Reddish brown; connexiva with pale yellow spots; hypopygium
dark brown; antenna and legs yellow brown.

Total length.—6.72 mm; width of pronotum 2.32 mm; width of abdomen
2.80 mm.

Holotype.—d , Colombia, Ville de Cauca, 4 km NW San Antonio, 6500’;
A.H. Miller leg. 1.X1.1958. Deposited at the California Academy of Sci-
ences, San Francisco, California.

Remarks.—Notrapictinus piliger runs to N. quadraticeps (Champion),
1898, in my key (Kormilev, 1967: 7) but may be separated at once by the
long, erect bristles on the body, antennae, and legs.

Genus Mezira Amyot and Serville, 1843
Mezira hondurensis Kormilevy, NEw Species
Figs. 9—11

Male.—Elongate ovate, partially covered with short, golden, curled hairs;
connexivum tricolored, ferrugineous, yellow, and black.

Head: As long as its width across eyes (24:24); anterior process with
parallel sides, incised anteriorly, reaching 2 of antennal segment I; anten-
niferous tubercles acute and slightly divaricating; postocular small, not
reaching outer border of eyes. Eyes large, strongly protruding. Vertex with
M-shaped rows of granules. Antennae moderately strong, twice as long as
width of head across eyes (49:24); relative length of antennal segments I to
IV are: 13:11:15:10. Labium reaching hind border of labial groove, which
is closed posteriorly.

Pronotum: Short and wide (28:54); collar sinuate anteriorly, granulate;
anterolateral angles produced forward as rounded lobes as far as collar;
lateral notch forming an obtuse angle; lateral borders of hind lobe parallel,
rounded and strongly converging anteriorly; hind border twice sinuate. Fore

disc with 4 (2+2) ridges; hind disc granulate.

Scutellum: Shorter than its basal width (25:30); all borders carinate, lateral
sinuate before apex, tip rounded. Disc with 2 (1+1) transverse, naked ele-

vations along basal border; median ridge narrow.

486 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Hemelytra: Reaching hind border of tergum VI; corium reaching % of
connexivum III, its basolateral border slightly sinuate and reflexed, apical
border sinuate interiorly, rounded exteriorly; apical angle rounded.

Abdomen: Ovate, longer than its maximum width across segment IV
(89:67): lateral borders rounded; posteroexterior angles of connexiva II to
VI slightly protruding, VII rounded. Paratergites clavate, reaching % of
hypopygium; the latter cordate, median ridge reaching tip of disc. Spiracles
II to VII ventral, placed far from border, VIII lateral and visible from above.

Female.—Similar to male, but slightly larger. Paratergites large, rounded
posteriorly, reaching 2 of incised posteriorly segment IX.

Measurements.—Head 25:25; relative length of antennal segments I to IV
are: 13:11:15:10; pronotum 28:55; scutellum 25:32; abdomen 95:69; width of
tergum VIII, 29.

Color.—Ferrugineous; connexivum with yellow round spots and anterior
4 of exterior border; posterior 34 of exterior border is black.

Total length.—颗6.76, 2—7.00 mm; width of pronotum: d—2.16,
2—2.20 mm; width of abdomen: ¢—2.68, 2 —2.76 mm.

Holotype.—¢d, Honduras, 10 km N Siguatepeque; 20.VIII.1978, J.A.
Chemsak, E.G. & J.M. Linsley leg. Deposited at the California Academy
of Sciences, San Francisco.

Allotype.— , collected with holotype; same collection.

Paratypes.—6 , 2 2, collected with holotype; same collection and Kor-
milev collection.

Remarks.—Mezira hondurensis is related to M. hyperlobata Kormilev,
1962, from Brasil, but the head is as long as the width across the eyes; the
anterolateral lobes of the pronotum are directed more forward and only
reach level of the collar; the lateral notch of the pronotum is obtuse; and
the (°) paratergites are large, reaching % length of segment IX.

Mezira proxima Kormilev, NEw SPECIES
Fig. 12

Female.—Elongate ovate, abdomen with subparallel sides; prosternum,
propleura and venter laterally, with thin layer of white incrustation. Pilosity
short and curled.

Head: Shorter than its width across eyes (21.0:23.5); anterior process
rounded and slightly incised anteriorly, genae contiguous in front of clypeus,
reaching *% of antennal segment I; antenniferous tubercles wide, acute, di-
varicating; postocular dentiform, reaching or almost reaching, outer border
of eyes. The latter large, protruding. Vertex with V-form rows of granules.
Antennae moderately stout, less than 142X as long as width of head across
eyes (32.5:23.5); relative length of antennal segments I to IV are:
10.0:7.0:8.0:7.5. Labium not reaching hind border of labial groove, which
is closed posteriorly.

VOLUME 84, NUMBER 3 487

Pronotum: Less than % as long as its maximum width (22:48); collar
slightly sinuate anteriorly and granulate; anterolateral angles rounded, cren-
ulate and slightly expanded; lateral notch shallow; lateral borders of hind
lobe parallel, converging anteriorly; hind border sinuate medially. Fore disc
with 4 (2+2) oblique ridges; hind disc granulate.

Scutellum: Shorter than its basal width (20:25); disc crushed by pin.

Hemelytra: Reaching hind border of tergum VI; corium reaching % of
connexivum III; its hind border rounded, apical angle also rounded.

Abdomen: Longer than its maximum width (80:54); lateral borders sub-
parallel from II to V, then converging in arquate line; connexiva II and III
semifused together; posteroexterior angles of connexiva not protruding; par-
atergites subangular, rounded posteriorly, reaching 12 of rounded poste-
riorly segment IX. Spiracles II to VI ventral, placed far from border; VII
also ventral, placed nearer to border; VIII lateral and visible from above.

Legs: Unarmed; femora and tibiae roughly granulate.

Color.—Testaceous; membrane infuscate, white at base.

Total length.—5.80 mm; width of pronotum 1.92 mm; width of abdomen
2.16 mm.

Holotype.—@ , Peru, Loromayo, 4-10.1X.1962, L.E. Pena leg. Deposited
at the California Academy of Sciences, San Francisco.

Remarks.—Mezira proxima is closely related to M. paraensis Kormilev
and Hess, 1979, but is larger; the labium is relatively shorter, not reaching

- the hind border of the labial groove; the basolateral border of the corium is

straight; spiracle VII is placed near the margin; and the ventral side of the
body is partially incrusted.

Mezira guanacastensis Kormiley, NEw SPECIES
Figs 13, 14

Male.—Closely related to Mezira neotropicalis (Champion), 1898, but
smaller; antennae relatively shorter, only 1.52< as long as width of head
across eyes (2X in M. neotropicalis); antennal segment III relatively short-
er, only slightly longer than I and only by % longer than IV (much longer
than I (23:18) and almost twice as long as IV (23:13) in M. neotropicalis);
antenniferous tubercles relatively shorter and not so pointed as in M. neo-
tropicalis. Hypopygium relatively wider, ratio length:width as 13:18 (13:16
in M. neotropicalis). Paratergites (2) relatively shorter, reaching 2 of seg-
ment IX (almost reaching tip of [IX in M. neotropicalis).

Measurements.—Head: ¢—22:24, 9—23:25; relative length of antennal

| segments I to IV are: 6—10.0:8.0:11.0:7.5, 2—11:8:12:8; pronotum: ¢—

25:49, 2—26:50; scutellum: ¢—18:26, 92—18:28; abdomen: d—73:56,

| 2—88:62; hypopygium 13:20; width of tergum VIII (2), 27.
Paratergites—(¢) clavate, reaching slightly beyond ’% of hypopygium;

the latter cordate, shorter than its maximum width. Rounded posteriorly in

488 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

female reaching 4% of segment IX. Spiracles II to VII ventral, placed far
from border; VIII ventral, placed near border.

Color.—Ferrugineous, membrane black.

Total length.—d¢—5.64, 2—6.20 mm; width of protnotum: ¢—1.96,
2?—2.04 mm; width of abdomen: ¢—2.24, 2—2.48 mm.

Holotype.—d , Costa Rica, Guanacaste Prov., Comelco Areal, 5 km W
Bagaces; under bark; 28.1X.1973, P. A. Opler leg. Deposited at the Cali-
fornia Academy of Sciences, San Francisco.

Allotype.—2 , collected with holotype; same collection.

Paratypes.—S 3, collected with holotype; same collection and Kormilev
collection.

ACKNOWLEDGMENT

I express my sincere thanks to L. Jessop, Department of Entomology,
British Museum (Natural History), London, who kindly examined the types
of Mezira neotropicalis (Champion), 1898, and made drawings of the genital
segments.

LITERATURE CITED

Champion, G. C. 1898. Biologia Centrali-Americana. London. Heteroptera 2: 65-117, 3 pls.

Kormilev, N. A. 1962. Notes on Aradidae in the Naturhistoriska Riksmuseum, Stockholm
(Hemiptera-Heteroptera). Ark. Zool. (2) 15(14): 255-273, 2 pls.

—. 1967. On some Aradidae from Brasil, Argentina and Laos; (Hemiptera-Heteroptera).
Opusc. Zool. (Munich) 100: I-10.

Kormilev, N. A. and E. Heiss. 1979. Four new species and lectotype designation of some
neotropical Aradidae (Insecta, Heteroptera). Entomol. Arb. Mus. G. Frey Tutzing bei
Muench. 28: 93-118, 34 figs.

Picchi, V. D. 1977. A systematic review of the genus Aneurus of North and Middle America
and the West Indies (Hemiptera: Aradidae). Quaest. Entomol. 13: 255-308, 233 figs, [X
maps.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 489-494

TWO NEW SPECIES OF ESENBECKIA (DIPTERA: TABANIDAE)
FROM MEXICO’?

RICHARD C. WILKERSON AND G. B. FAIRCHILD
Department of Entomology and Nematology, University of Florida, In-

stitute of Food and Agricultural Sciences, Gainesville, Florida 32611; Re-
search Associates, Florida State Collection of Arthropods.

Abstract.—Two new species of Esenbeckia from Mexico are described
_and figured, EF. (Ricardoa) biclausa, n. sp., from Hidalgo State is compared
to E. (R.) scionodes Philip, and E. (Esenbeckia) hirsutipalpus, n. sp., from

Nuevo Leon State is compared to E. (E.) illota (Williston).

_ The genus Esenbeckia Rondani with its four subgenera is an entirely
American group of relatively primitive Pangoniini. The genus as a whole
ranges from Argentina to southern United States, but the subgenera, except
Esenbeckia itself, have more restricted ranges. Philip (1978) has recently
reviewed, with keys to both sexes, the Mexican species, most of which
belong to the subgenus Ricardoa Enderlein, a group which does not range
south of Panama. Fairchild and Wilkerson (1981) give a detailed review of
the subgenus Proboscoides Philip and have nearly completed a similar treat-
-ment of the nominate subgenus. The remaining subgenus, Palassomyia Fair-
child contains but one peculiar Chilean species. Very little is known of the
early stages or habits of any of the species. For the most part they seem to
be rare and local; many are haematophagous, especially in the subgenus
Esenbeckia and probably Proboscoides, while those in Ricardoa are often
taken feeding on flowers. Philip’s paper contains references to the pertinent
literature, but is without figures, though most of the species have been
figured by him or others in earlier publications.
The two species described here are peculiar in several ways and form
interesting additions to the Mexican fauna. Esenbeckia (Ricardoa) biclausa
is apparently close to E. (R.) scionodes Philip, having the same, hitherto

' Florida Agricultural Experiment Station Journal Series No. 2962.
* This material is based upon work supported by the National Science Foundation under
Grant DEB-78-10121.

490 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

unique, Scione-like venation, while males of E. (Esenbeckia) hirsutipalpus
were reported by Philip (1978) as possibly the first records of E. (E.) illota
from Mexico. The female collected by H. V. Weems, Jr., shows little sim-
ilarity to E. illota, except for general coloration, but seems surely correctly
associated with the sympatric males.

Esenbeckia (Ricardoa) biclausa Wilkerson and Fairchild, NEw SPEcIEs
Figs. 1[A—D

A blackish, yellow-haired, medium-sized, robust, beelike fly with an or-
ange antennal flagellum, and short, wholly hairy, pointed palpi. Legs bicol-
ored, femora blackish, tibiae yellow. Wings unpatterned but veins margined
brown, Ist and 4th posterior cells closed and fork of 3rd vein with an ap-
pendix.

Female.—Length 12.5 mm; of wing 11.5 mm. Head structures as figured.
Frons blackish in ground color, covered with gray pollinosity which is dens-
est on the lower 4. Frontal hairs as figured, all pale lemon yellow. Callus
absent but a median raised ridge present. Ocellar tubercle prominent and
black, bearing 3 reddish-brown ocelli. Subcallus, genae, and frontoclypeus
blackish in ground color covered with dense gray pollinosity. Area around
antennal sockets yellowish-brown pollinose. Oculogenal hairs pale lemon
yellow. Beard of dense, pale lemon-yellow hairs. Antennal segments | and
2 yellowish brown in ground color, sparsely pale grayish-yellow pollinose
with pale lemon-yellow hairs. Segment 3 orange, dusky at apex of last an-
nulus. Palpal segment | blackish in ground color, pale yellow pollinose;
segment 2 dark reddish brown which shows through thin yellowish polli-
nosity and shiny, pale yellow, bristle-like hairs. Second segment with deep
groove on outer aspect. Proboscis wholly sclerotized, black, surface shiny
but rough.

Mesonotum and scutellum black in ground color with dark gray pollinosity
and quite dense, long, pale lemon-yellow hairs. Tufts of hairs at wing base
and in front of scutellum whitish yellow. Ground color of pleura and coxae
dark reddish brown with dark gray pollinosity and dense, long, pale lemon-
yellow hairs. Femoral ground color dark reddish brown, yellowish brown
at apical tips. Femora grayish pollinose with long, pale lemon-yellow hairs.
Tarsi and apical tips of tibiae brown with pale lemon-yellow hairs. Tibiae
yellow with pale lemon-yellow hairs. Basicosta bluntly pointed and blackish.
Wings hyaline except for yellowish line along leading edge, especially in
costal, subcostal, and Ist basal cells. All veins brown and brown margined.
First and 4th posterior cells closed, fork of 3rd appendiculate. Halter reddish
brown, dusky at tip of and base of knob.

Abdomen very dark shiny brown except for Ist segment which is dark
gray pollinose. Setal vestiture of quite dense, pale lemon-yellow hairs which

cr

VOLUME 84, NUMBER 3 49]

Figs. |A-D. Esenbeckia (Ricardoa) biclausa. 1A, Head, side view. 1B, Palp. 1C, Antenna,
1D, Frons. Figs. 2A-D. Esenbeckia (Esenbeckia) hirsutipalpus. 2A, Antenna. 2B, Palp, 2C,
Frons. 2D, Head, side view.

492 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

are most numerous along segmental borders. Yellow color along borders
accentuated by narrow strips of underlying pale yellow integumental color.
Abdomen below as above but with hairs slightly longer and more number-
ous.

Male.—Length 15 mm; of wing 13 mm. Eyes contiguous. Ocellar tubercle
at vertex prominent, bearing 3 distinct reddish-brown ocelli. Antennal fla-
gellae missing. Palpi as female but slightly longer and curved. Proboscis
more slender than female. Thorax, legs, and wings as female but hairs longer
and more numerous and pollinosity with yellowish hue. Abdominal hairs as
female but longer and more numerous both above and below. Ground color
blackish on Ist tergum, middle % of the 2nd and 3rd, and on all but apical
borders of remaining terga. Lateral portions of terga 2 and 3 and apical
borders of following segments reddish yellow. All segments grayish-yellow
pollinose. Abdomen below as above but sternum 3 with lateral reddish-
yellow areas.

Type material.—Holotype, 2, Mexico, Hidalgo, Tepeapulco, 1-[X-1974,
G. Bohart and W. Hanson coll. Allotype, ¢ , Mexico, Hidalgo, Tepeapulco,
18-I[X-1974, G. Bohart and W. Hanson coll. To be deposited in Florida State
Collection of Arthropods (F.S.C.A.)

Discussion.—Esenbeckia scionodes Philip is apparently sympatric with
and closely allied to E. biclausa, n. sp.; however, biclausa is more robust
and lacks the lateral reddish areas on the first two abdominal terga. It also
has shorter palpi which are blackish, with long pale yellow hairs, not orange
with black hairs. The first annulus (basal plate) of the flagellum is also much
wider than in scionodes, being half again as wide as the scape. In other
respects, including leg coloration, overall body vestiture and coloration, and
closure of the fourth posterior cell, the two species are quite close.

We place it in the subgenus Ricardoa on the basis of the short grooved
palpi and the slender proboscis even though it, like E. (Ricardoa) scionodes,
does have oculongenal hairs.

All specimens of the type-series of E. scionodes Philip do not have a
completely closed 4th posterior cell. Of the 19 specimens, 7 were strongly
narrowed and 3 ‘‘variously more opened”’ (Philip, 1973). One might expect
this same variability in a larger series of E. biclausa.

Esenbeckia (R.) incisuralis (Say) is also a robust, yellow-haired, beelike
species with darkened femora. The palpi of E. incisuralis are longer with a
bare median groove, not short and wholly hairy; the antennal basal plate is
narrower than E. biclausa and the 4th posterior cell is always open.

Esenbeckia (Esenbeckia) hirsutipalpus Wilkerson and Fairchild, New SPEcIEs

Figs. 2A—D

Esenbeckia illota: Philip 1978:350. In part, 2 6 reported from Nuevo
Leon.

VOLUME 84, NUMBER 3 493

A medium-sized yellowish-brown species lacking a distinct callus, but
with sabre-shaped palpi covered completely with long brown hairs; probos-
cis about equal to head height; pale fore- and midtibiae, and smoky brown
wings. Abdomen brown, Ist segment contrastingly pale translucent yellow-
ish white.

Female.—Length 14 mm; of wing 14.5 mm. Head structures as figured.
Frons yellowish-brown pollinose, bearing on its lower /% a slight depression
with a barely evident median ridge which extends upward into a small slight-
ly denuded callus. Tubercle at vertex raised and bearing 3 dark red ocelli.
Subcallus, genae, and frontoclypeus with brownish integument and yellow-
ish-brown pollinosity; beard and other facial hairs dark brown. Antennal
pedicel and scape yellowish-brown pollinose, flagellum pale yellow orange;
antennal hairs dark brown. Palpi dark yellowish brown, densely covered
with long dark brown hairs. Proboscis shiny brown.

Integument of mesonotum, scutellum, pleura, and coxae brown; yellow-
ish-brown pollinose. Setal vestiture of numerous brown hairs, longer lat-
erally, on scutellum and pleura. Hairs of thorax nearest head pale, yellow-
ish. Femora and hindtibia and tarsus brown and brown haired; hindfemur
slightly darker. Fore- and midtibiae and tarsi pale yellowish brown and short
yellow haired. Halter yellowish brown. Wings with brownish tinge, darker
in costal, subcostal, Ist basal, and base of marginal cells. Wing venation
normal for genus.

Abdomen above brown and brown haired with Ist tergum translucent pale
yellowish white and pale yellowish-brown haired. Extreme lateral borders
of terga 2—7 blackish; posterior borders of terga 3—7 darkened. Below as
above but without lateral darkening and with posterior borders of sterna 2—
6 dark.

Male.—Length 13 mm; of wing 13 mm. Similar to female but paler overall.
Head, thoracic, and leg hairs long, pale yellowish brown; those of Ist ab-
dominal tergum pale yellow, remaining abdominal hairs pale brown. Eyes
contiguous, a distinct raised tubercle bearing 3 ocelli at vertex. Palpi similar
to female, but shorter, slender, and curved. Palpal hairs numerous, long,
and reddish brown. Proboscis short, littke more than ’% head height, only
Ys again as long as palp. Abdomen as female, segment | pale yellow trans-
lucent, lateral borders on terga 2—7 darkened.

Type material.—Holotype, 2, Mexico, Nuevo Leon, Chipinque Mesa,
18-VI-1976, H. V. Weems coll. To be deposited in F.S.C.A. Allotype, ¢,
Mexico, V. deSantiago, N. L. (Nuevo Leon), H. Hedderich coll. Cornelius
B. Philip collection in California Academy of Sciences San Francisco
(C.A.S.). Determined as Esenbeckia sp. nr. illota by C. B. Philip, 1957.
Paratype, 6, Mexico, Apodaca, N.L., A. Salazar coll., in collection C. B.
Philip in C.A.S. determined as E. illota in 1954 and E. sp. nr. illota in 1957
by C. B. Philip.

Discussion.—Esenbeckia hirsutipalpus is similar to E. illota (Williston)

494 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

in that both are medium sized straw-colored to brown species with smoky
brown wings and the first abdominal segment translucent horn colored. The
palpi of the two are strikingly different in both sexes; however, those of
hirsutipalpus are wholly long haired, and those of illota are short haired
and mostly bare on the outer aspect. In addition, hirsutipalpus has a stouter
and much shorter proboscis, and lacks a distinct shiny frontal callus.

LITERATURE CITED

Fairchild, G. B. and R. C. Wilkerson. 1981. New species of Esenbeckia (Proboscoides) (Dip-
tera: Tabanidae) with a key to the subgenus. Fla. Entomol. 64(1): 158-175.

Philip, C. B. 1973. New North American Tabanidae (Diptera). XX. Six new species of Esen-
beckia subgenus Ricardoa from Mexico. Ann. Entomol. Soc. Am. 66(5): 1141-1145.

. 1978. New North American Tabanidae (Insecta: Diptera). XXIV. Further comments

on certain Pangoniinae in Mexico with special reference to Esenbeckia. Proc. Calif.

Acad. Sci. (4) 41(14): 245-356.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 495-508

AN ANNOTATED CHECKLIST OF THE CADDISFLIES
(TRICHOPTERA) OF MISSISSIPPI AND
SOUTHEASTERN LOUISIANA. PART I:

INTRODUCTION AND HYDROPSYCHOIDEA

PAUL K. LAGO, RALPH W. HOLZENTHAL, and STEVEN C. HARRIS

(PKL) Department of Biology, University of Mississippi, University,
Mississippi 38677; (RWH) Department of Entomology, Fisheries and Wild-
life, Clemson University, Clemson, South Carolina 29631; (SCH) Environ-
mental Division, Geological Survey of Alabama and Department of Biology,
University of Alabama, University, Alabama 35486.

Abstract.—Distributional records for 46 species of caddisflies in the su-
perfamily Hydropsychoidea (Hydropsychidae, Philopotamidae, Polycentro-
podidae, and Psychomyiidae) are presented, and the seasonal distribution of
each is indicated. Annotations for many species include habitat notes. Of the
species reported, 38 represent new state records for Mississippi, and 18 are
new records for Louisiana.

Faunal lists of caddisfly species have been prepared for several of the
states in the southeastern United States. Studies conducted in Arkansas
(Unzicker et al., 1970), Florida (Blickle, 1962), Kentucky (Resh, 1975),
North and South Carolina (Morse, 1970; Morse et al., 1980; Unzicker et al.,
in press), Tennessee (Etnier and Schuster, 1979), Texas (Edwards, 1973),
and Virginia (Parker and Voshell, 1981) have provided preliminary to fairly
complete checklists. Although more than 450 species have been reported
from the southeast (Morse, personal communication), the caddisfly faunas
of the gulf coastal states (Texas, Louisiana, Mississippi, Alabama, and Flor-
ida) and of Georgia are virtually unknown and records available from these
states are few and scattered. Only 10 species have been reported from Mis-
sissippi and only 14 from Louisiana.

The following list represents a compilation of two separate surveys. Hol-
zenthal (1980) recently completed a study of the caddisflies of the southern
third of Mississippi and the southeastern ‘‘Florida”’ parishes of Louisiana,
and Lago and Harris have been conducting a general survey of the Tri-
choptera of Mississippi since 1977. The information we have accumulated
will be presented in three parts, each covering one of the three trichopteran

496 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

superfamilies, the Hydropsychoidea, Rhyacophiloidea, and Limnephiloidea
(Ross, 1967).

GENERAL PHYSIOGRAPHY OF THE STUDY AREA

Both Mississippi and Louisiana lie in the Nearctic Coastal Plain Physio-
graphic Province with only the extreme northeastern corner of Mississippi
belonging to the Interior Low Plateau Province. This latter area is small and
the state is usually considered entirely Coastal Plain (e.g. Berner, 1977). Lowe
(1919) divided Mississippi into ten physiographic regions differing in soil
types and natural vegetation. These regions can be generally grouped into
four areas containing fairly distinct aquatic habitat types (Stanford, 1980).
The areas, as shown in Figure |, are the Tennessee River Hills (TRH), the
North Central Plateau (NCP), the Yazoo Delta (YD), and the Coastal Plain
(CP). The TRH area presents the roughest terrain in the state and is char-
acterized by swift, clear streams with rocky-sandy bottoms. The NCP region
has sandy or silty streams and many man-made lakes with mud or leaf-
littered bottoms. Gravel-bottomed streams are occasionally encountered
throughout the area. West of this lies the YD area which is characterized
by meandering bayous, ox-bow lakes and small silt-bottomed streams. Most
of the southern half of Mississippi and the portion of Louisiana included in
the study area (the southeastern ‘‘Florida’’ parishes) is CP. Streams and
rivers here have sand and gravel bottoms, and streams stained with tannic
acid (black water) are ubiquitous.

LIsT OF COLLECTION SITES

Figure | shows the distribution of 132 localities in the study area from
which specimens were obtained. The following list will serve as an index to
collecting sites for the three parts of our checklist and will not be repeated
in subsequent parts.

Tennessee River Hills

Tishomingo County

1. Iuka.

2. Tishomingo State Park.
Monroe County

3. Hamilton.
Lowndes County

4. Columbus.

North Central Plateau

Marshall County
5. 4 mi. N Holly Springs (T3S-R3W-Sec. 13).

VOLUME 84, NUMBER 3 497

Clary
Ne Lf my
Sec
1 era

Vee

Ll

2)
~“

| Fig. 1. Collecting sites and major physiographic regions of the study area. CP—Coastal
| Plain; NCP—North Central Plateau; TRH—Tennessee River Hills; YD—Yazoo Delta.

498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

6. Holly Springs.

7. Wall Doxey State Park, Spring Lake.
Benton County

8. Hickory Flat.
Union County

9.8.5 mi NW New Albany (T6S-R2E-Sec. 5).
Lafayette County

10. 11 mi. NW Oxford, Sardis Lake.

11. 4.5 mi. SW Abbeville, Dunlap Branch.

12. 4 mi. NW Oxford, Cataraci Branch.

13. 11 mi. NE Oxford, Puskus Lake.

14. 8 mi. NE Oxford, Bay Springs Branch.

15. 3 mi. NW Oxford, Davidson Creek.

16. Oxford.

17. 6 mi. E Oxford, Hopewell Lake.

18. 6 mi. S Oxford, Morris Creek.
Lee County

19. Tupelo.
Calhoun County

20. 13 mi. NW Bruce (T11S-R2W-Sec. 25).
Clay County

21. Cedar Bluff.
Oktibbeha County

22. Adaton.

23. State College (Mississippi State University).
Winston County

24. 13 mi. W Louisville, Branch of Labutcha Creek.
Madison County

25. 10 mi. NE Canton, Dry Creek.
Leake County

26. Carthage.

27. 5 mi. S Carthage, Pearl River.
Neshoba County

28. 1.5 mi. N Dixon.

29. 7 mi. SW Philadelphia.
Scott County

30. Golden Memorial State Park.

31. Roosevelt State Park.
Newton County

32. Roberts.

33. 4.5 mi. E Newton, small pond.

34. 6 mi. SE Newton, Falema Creek.

VOLUME 84, NUMBER 3

Lauderdale County
35. Meridian.
Smith County
36. | mi. W Lake Marathon, Branch of Ichusa Creek.
Jasper County
37. 6 mi. NW Montrose, Cedar Creek.
Clarke County
38. 3 mi. N Enterprise, Chunky Creek.
39. Clarke Co. State Park, Moore Mill Creek.

Yazoo Delta

Bolviar County

40. 2 mi. W Rosedale, Mississippi River.
Quitman County

41. 2 mi. S Sledge, Flagg Lake.
Washington County

42. Stoneville.

43. Wayside.

Coastal Plain
Mississippi
Warren County
44. Bovina.
45. Vicksburg.
46. 6 mi. S Yokena (T13N-R3E-Sec. 30).
Hinds County
47. Big Black River x Interstate 20.
48. 4.5 mi. NW Clinton.
49. 4 mi. N Clinton, Bogue Chitto Creek.
50. Clinton.
51. Jackson.
Rankin County
52> Pearl.
Claiborne County
53. Rocky Springs.

54. Little Sand Creek, nr Rocky Springs (Natchez Trace Parkway).

55. 3 mi. S Rocky Springs, Owens Creek.
56. Bayou Pierre at Carlisle.

Simpson County
57. 2 mi. W Pinola, Strong River x Hwy 28.
58. Westville Creek <x Hwy 43, nr Pinola.
59. 2 mi. SE Pinola, Tanyard Creek x Hwy 28.
60. 4 mi. SE Pinola, Mill Creek x Hwy 472.

499

500 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

61. 3 mi. E Magee, Ocatoma Creek Hwy 28.
Jasper Co.

62. 2.5 mi. E Bay Springs, Tallahoma Creek x Hwy 528.
Adams County

63. Natchez.

64. 5 mi. S Natchez.

65. Homochitto National Forest, Sandy Creek.
Franklin County

66. 7 mil. SW Meadville, Clear Springs Lake.
Lincoln County

67. 4 mi. W Caseyville, Homochitto River x Hwy 550.

68. Brookhaven.

69. E Fork Amite River X Hwy 98, nr Auburn.

70. 7 mi. N Summit on Interstate 55.
Covington County

71. Okatoma Creek X Hwy 598, nr Sanford.
Wayne County

72. Waynesboro.
Wilkinson County

73. Buffalo River x Hwy 61.

74. Fort Adams.

75. 1 mi. SW Pond, Clark Creek.

76. Centerville.
Amite County

77.3 mi. SW Liberty, W Fork Amite River X Hwy 48.

78. Wagoner Creek X Hwy 48, nr Liberty.

79. 9 mi. S Liberty, E Fork Amite River.

80. 9 mi. SSW Liberty, W Fork Amite River.
Pike County

81. .5 mi. W Pricedale, Topisaw Creek x Hwy 44.

82. Percy Quinn State Park, Percy Quinn Lake.

83. 3 mi. S Magnolia, Tangipahoa River.
Walthall County

84. 5 mi. SE Lexie, Bogue Chitto River.
Marion County

85. Lake Columbia.
Lamar County

86. 5 mi. N Baxterville, Half-Moon Creek.
Forrest County

87. Hattiesburg.

88. Paul B. Johnson State Park.

89. 9.5 mi. S Brooklyn.

VOLUME 84, NUMBER 3 501

Perry County
90. | mi. E Janice, Cypress Creek x Hwy 29.
91. Leaf River Game Mang. Area.
Pearl River County
92. 5 mi. S Crossroads, Chinquapin Creek x Hwy 43.
93. | mi. SW Silver Run, Wolf River.
94. Picayune.
Stone County
95. 5 mi. W Wiggins, Red Creek x Hwy 26.
96. Wiggins.
97. 5 mi. E Wiggins, Flint Creek x Hwy 26.
98. Perkinston.
99. 13 mi. SW Wiggins, Sandy Creek.
100. University of Mississippi forest lands, headquarters.
101. Red Creek x Hwy 15.
George County
102. 7 mi. S Benndale.
Hancock County
103. 4 mi. NW Kiln, Orphan Creek <x Hwy 43.
104. NASA Missile Testing Facility.
105. Bayou LaTerre.
106. 6 mi. WNW Waveland.
107. Bay St. Louis.
Harrison County
108. 2 mi. N Lyman, Little Biloxi River.
109. Long Beach.
110. Handsboro.
iD tberville:
112. Biloxi.
113. Keesler Air Force Base.
114. Big Biloxi River (exact locality unknown).
Jackson County
115. Ocean Springs.

Louisiana
West Feliciana Parish
116. 2 mi. NNW Weyanoke, Little Bayou Sara x Hwy 66.
East Feliciana Parish
117. 4.5 mi. NNE Felixville, confluence of E and W Forks Amite River.
118. 10 mi. E Clinton, confluence of E and W Prongs Amite River.

St. Helena Parish
119. 3 mi. S Pine Grove, Claiborne Branch x Hwy 449.

502 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Tangipahoa Parish
120. 2.5 mi. E Holton, Chappepeela Creek x Hwy 16.
121. 6 mi. E Holton, Tchefuncte River x Hwy 16.
122. 8 mi. NW Folsom, Tchefuncte River X Parish Rd 19.
123. 4.5 mi. W Folsom, Tchefuncte River x Hwy 40.
Washington Parish
124. 11 mi. N Folsom, Bonner Creek X Hwy 25.
St. Tammany Parish
125. 6 mi. N Folsom, Bogue Falaya River X Hwy 25.
126. | mi. S Folsom, Morgan Branch Bogue Falaya River x Hwy 25.
127. Talisheek Creek x Hwy 435, nr Talisheek.
128. Talisheek Creek X Hwy 41, E of Talisheek.
129. 4.5 mi. E Abita Springs, Abita Creek x Hwy 435.
130. Slidell, barrow pit at Hwy 433.
Livingston Parish
131. Hornsby Creek x Hwy 1025, .5 mi. E of Hwy 49.
Orleans Parish
132. Biological Sciences Annex, University of New Orleans.

LIST OF SPECIES

The Hydropsychoidea are represented in the study area by 46 species in |
15 genera and four families. In the following list those records obtained from
literature sources are indicated by the citation. Since most of the species |
have rather broad temporal distributions, a range of collection dates is pre-
sented when more than three dates are involved. Widely disjunct collection
dates are also indicated. Voucher specimens have been deposited in collec-
tions at the University of Mississippi, Clemson University, and Louisiana
State University.

Philopotamidae

Chimarra aterrima Hagen. Sites 2, 70. 5 April and 8 Sept. Rare in our
collections. Several specimens were collected near a small, swift, rocky
stream.

Chimarra feria Ross. Sites 6, 55, 70. 20 March-2 June. Uncommon. !

Chimarra florida Ross. Sites 54, 86, 97, 99, 100, 107, 108, 110, 120, 123, |
125, 126, 127, 129, 131. 28 March—5S June, 18 Sept. and 17 Oct. Locally |
abundant in small to medium black water streams in the CP. The widely
disjunct collection dates may indicate two cohorts per year for this species
in the study area.

Chimarra obscura (Walker). Sites 14, 16, 57, 58, 59, 66, 67, 101. 4 May—26
Sept. Although Ross (1944) stated that this species frequents rapid, clear
streams, several of the adults we have collected came from light traps
near small spring-fed lakes. Not common.

VOLUME 84, NUMBER 3 503

Chimarra socia Hagen-C. moselyi Denning *“‘complex.’’ Sites 2, 16, 25, 26,
NO. 7s 365-59, Gl, 79,180, 8183564, 93, 955°97, 99, 1005" 10151021197,
118, 120, 126, 129. 11 April-29 Sept. It seems that this complex is rep-
resented in the study area by two or three variable species, or perhaps a
complex of described and undescribed species. Many more specimens
will need to be studied in order to understand the variation presented by
this series. This group is widespread and common throughout the CP, and
specimens were collected along slowly flowing, as well as rapid, streams.

Wormaldia moesta (Banks). Sites 16, 17, 100. 10 April—14 June. Rare in our
collections.

Psychomylidae

Lype diversa (Banks). Sites 25, 36, 60, 80, 83, 90, 121, 122, 123, 124, 125,
127. 20 March—10 June, 15 Sept. and 17 Oct. The spring and fall collection
dates for this CP species indicate it may be bivoltine in the study area.

Polycentropodidae

Cernotina calcea Ross. Sites 57, 62, 67, 69, 71, 80, 83, 106, 119, 120, 131.
23 May-—18 Sept. Specimens of C. calcea were not encountered in large
numbers, but the species is widespread in the CP.

Cernotina spicata Ross. Sites 7, 25, 31, 66, 71, 86, 100. 6 May—26 May, 30
Aug.—9 Sept. Less common than C. calcea, but considerably more wide-
spread in the study area. The split collection dates suggest a bivoltine life
cycle here. Specimens were collected near spring-fed lakes, a swift black-

water river and small sand bottom creeks, confirming the wide variety of

habitats noted for this species by Hudson et al. (1981).

Cyrnellus fraternus (Banks). Sites 5, 7, 10, 11, 14, 16, 23, 28, 40, 41, 49, 82,

| 85, 86, 87, 106, 126, 129. 21 April—-26 Sept. Widespread and common;
specimens were collected from a wide variety of aquatic habitats in the

study area.

| Neureclipsis crepuscularis (Walker). Sites 16, 24, 25, 40, 42, 44, 47, 48, 52,
57, 73, 79, 83, 86, 87, 120, 125. 6 May—10 Sept. Widespread and common.

| In the study area, capture nets of larvae were observed on the tops of

| submerged logs, on firm gravel substrate and on Vallisneria leaves in
moderate current.

| Neureclipsis melco Ross. Site 128. 4 June. This is a black-water, coastal

| plain endemic, and was previously known from Georgia and South Car-
olina. It is not common here.

| Nyctiophylax affinis (Banks) Sites 6, 7, 14, 15, 16, 21, 29, 32, 40, 41, 50, 62,

| 100, 119, 120. 21 April-6 July, 9 Sept., 27 Sept. Widely distributed and

_ fairly common. Specimens were collected near tiny streams, the Missis-

| sippi River and several spring-fed lakes. Morse (1972) reported this
species from Mississippi, but gave no specific locality.

|

504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Nyctiophylax banksi Morse. Sites 101, 102, 108. 24 and 25 May. Rare in our
collections. Specimens were collected near a large, swift black-water riv-
er, a smaller sand bottomed creek and a marshy area where little water
flow was discernable.

Nyctiophylax celta Denning. Sites 57, 97. 7 June, 20 June. Rare in our
collections.

Nyctiophylax denningi Morse. Sites 2, 57. 20 June, 21 July, 8 Sept. This
species was previously known from the southern Appalachians (Morse,
1972); consequently, the record from the TRH area was not unexpected.
However, the Simpson County record (CP) represents an interesting
range extension.

Phylocentropus carolinus Carpenter. Sites 60, 77, 78, 86, 95, 121, 126. 20
April—July, 17 October. Uncommon and apparently restricted to the CP
here. Several specimens were collected along black-water streams.

Phylocentropus lucidus (Hagen). Sites 2, 16, 83. 4 June, 10 June, 8 Sept.
More widely distributed but less common than P. carolinus in the study
area.

Phylocentropus placidus (Banks). Sites 14, 17, 23, 60, 61, 62, 71, 79, 81, 83,
119, 120,-122, 124, 127,129, 131,21 March—17, October:. Thisiis:the most
common species of Phylocentropus in the study area.

Polycentropus blicklei Ross and Yamamota. Sites 7, 88. 29 March, 21 May.
The Marshall County record (21 May 1957) was reported by Ross and
Yamamoto (1965). They had one male from Mississippi which was des-
ignated a paratype, and they felt that the Mississippi record represented
a relict population. A male from Forrest County represents the only other
specimen known from the study area.

Polycentropus centralis Banks. Site 53. 18 April. Only one specimen has
been collected here.

Polycentropus cinereus Hagen. Sites 103, 128. 26 May, | October. Rare in
our collections.

Polycentropus clinei (Milne). Site 100. 10 April, 15 April. Rare in our col-
lections.

Polycentropus confusus Hagen. Sites 81, 86, 97. 20 April, 7 June, 29 Sept.
Most specimens were collected near swift black-water creeks.

Polycentropus crassicornis Walker. Sites 4, 16, 19, 23, 24, 36, 41, 53, 63,
86, 112, 115. 2 April-15 June. The most widely distributed and common
species of Polycentropus in the study area. Collected from a variety of |
aquatic habitats, including swift black-water rivers and small streams.

Hydropsychidae

Cheumatopsyche burksi Ross. Sites 2, 11, 12, 14, 15, 17, 20, 21, 23, 25, 36,
40, 41, 44, 45, 49, 50, 51, 52, 59, 63, 64, 65, 66, 73, 83, 100. 28 March-10 |
October. Common in all regions of Mississippi, but no specimens were

VOLUME 84, NUMBER 3 505

collected in the ‘‘Florida’’ parishes of Louisiana. The extreme diversity
of habitat types from which specimens were collected indicates that the
species is tolerant of a wide range of environmental conditions.

Cheumatopsyche geora Denning. Sites 2, 66. 28 May, 22 July. Rare in our
collections. All specimens were collected near streams where they en-
tered small impoundments. These represent the western-most records for
C. geora.

Cheumatopsyche pasella Ross. Sites 2, 5, 16, 23, 25, 26, 29, 33, 35, 38, 40,
AAAs A952; 565,57; 595; 60,162; 65,69; 80/81 783) 84) 875 Dig 9295,
S900) MOT 2103007 , 11951209122, 1235125, 126129, 130. Apml=19
Sept. Reported from Site 23 by Gordon (1974). Abundant throughout the
study area. Shows about the same habitat specificity as C. burksi.

Cheumatopsyche petersi Ross, Morse and Gordon. Site 108. 25 May. Pre-
viously known only from Florida. About 20 specimens were collected
from this site near a small sandy-bottomed river.

Cheumatopsyche pettiti (Banks). Sites 4, 7, 10, 11, 12, 14, 16, 17, 18, 19,
PO 22322425 132,1335'34036, 37 9445495 50551952 759)163 5 655166;,70;
73, 75, 86, 91, 92, 99, 100, 103, 108, 112, 116, 119, 124, 131. 28 March—28
October. Widely distributed and abundant, but apparently absent from
the YD area. Perhaps the larvae of C. peftiti are not as tolerant as those
of C. burksi and C. pasella to the relatively warm water, silty conditions
present in the streams of that region.

Cheumatopsyche pinaca Ross. Sites 2, 12, 14, 16, 18, 86, 87, 100, 120, 125,
127. 20 March-17 October. Widely distributed but uncommon. Collected
from a variety of aquatic habitats, but most commonly from small black-

water streams.

Cheumatopsyche sordida (Hagen). 16, 21, 23, 49, 57, 59, 65, 66, 67, 71, 73,
81, 84, 86, 87, 90, 93, 102, 108, 114, 116. 23 May—20 October. This is the

| only common species of Cheumatopsyche in the study area which seems
to be restricted to the CP. We have only three records north of that area.
In light of its known distribution, widespread east of the Mississippi River,
the virtual absence of C. sordida from the northern half of the study area

| is somewhat puzzling.

ee virginica Denning. Sites 100, 125. 1 April-24 May. Known

previously from the Atlantic Coastal states (Gordon, 1974). Rare in our
| collections.

ee Coron modesta Banks. Sites 2, 16, 66, 72, 75, 86, 100. 7 April-8 Sept.

| Widespread but uncommonly encountered in the study area. Specimens

were collected near spring-fed lakes, black-water streams and small, clear
rocky streams.

Hydropsyche alvata Denning. Sites 2, 26, 51, 59, 73, 77, 79, 83, 91, 93, 95,

97, 99, 100, 101, 102, 103, 120, 127, 128, 129. 20 March, 30 April-8 Sept.
Denning (1949) described this species from specimens collected in Jack-

506 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

son, Mississippi. Principally a coastal plain species in the study area.
Specimens are seldom encountered in large numbers, but on one occasion
(Site 101, 24 May) several hundred were collected at a blacklight in 45
minutes. Most specimens were obtained from medium to large rivers.

Hydropsyche betteni Ross. Sites 2, 16, 18, 20, 59, 73, 75, 124. 7 April-8
Sept. Widely distributed but not common. Most specimens were collected
near small or medium streams with sand or gravel bottoms.

Hydropsyche ellisoma Ross. Sites 124, 125. 28 March, | April. Previously
known from the type locality and vicinity in Georgia, and South Carolina
(Morse et al., 1980). This species appears to be associated with spring-
fed black-water streams of the CP.

Hydropsyche mississippiensis Flint. Sites 7, 11, 16, 25, 26, 30, 58, 59, 65,
67,69, 71, 72:73, 77,, 785 815.825.8386, 90, 93,°95;.97 , 1005, 101 hO2 aaa
108, 120, 122, 123. 8 May—9 Sept. Flint (1972) described this species from
Wayne County, Mississippi (Site 72). Schuster and Etnier (1978) exam-
ined larvae of H. mississippiensis from Jones County, Mississippi. Fairly
common in the CP near streams of various types, also collected around
small spring-fed lakes. Often collected with H. alvata.

Hydropsyche orris. Ross. Sites 4;/5;.6, 7; 11, 16,417; 19, 21..23, 26,3255.
38, 40, 41, 43, 44, 45, 47, 48, 49, 50, 51, 52, 53, 56, 60, 62, 63, 64, 65, 66,
67,, 695, 73. 74,, 77, 785. 795 815, 82583 84, 87, 92;.95, 100s 110. JB eailioe
120, 127, 129. 8 April-16 October. Schuster and Etnier (1978) reported H.
orris from our study area (St. Tammany and East Baton Rouge parishes,
Louisiana). This is our most common species of Hydropsyche and is
found throughout the study area. It is particularly abundant along the
Mississippi River and near the large reservoirs of the NCP.

Hydropsyche phalerata Hagen. Sites 38, 84. 16 June, 13 October. Rare in
our collections, only two specimens were taken.

Hydropsyche rossi Flint, Voshell, and Parker. Sites 2, 5, 7, 10, 11, 13, 14,
16; 17). 18, 23, 24, 25; 26, 36;:41,:43, 44; 4546; 48,49, 50; 515)52. 56a
59, 60, 65, 79, 83, 86, 90, 92, 114, 116, 120, 127, 128. 11 March—24 October.
The holotype of H. rossi is from Hattiesburg, Forrest County, Mississippi
and was described by Flint et al. (1979). Paratypes are listed from various
locations in Mississippi and Louisiana as well as several other states. This
species is almost as widely distributed in the study area as H. orris,
however H. rossi is rarely encountered in the YD, an area where orris
abounds. Throughout the rest of the study area the two species are com-
monly collected together, but H. rossi is usually much less common.

Hydropsyche simulans Ross. Site 4. 16 April. Considered a Central States
species by Ross (1944), H. simulans is known from the study area by one
male specimen (det. O. S. Flint, Jr.).

Macronema carolina Banks. Sites 2, 3, 16, 23, 31, 32, 35, 51, 56, 57, 59, 62, |
65, 67, 69, 71, 75,77, 78, 81, 82, 83,84; 86, 87,92, 94,/95,96,:97. 99510 iee |

|

VOLUME 84, NUMBER 3 507

102, 103, 104, 106, 108, 112, 114, 117, 119, 121, 128, 129, 130. 7 April—23
October. Ross (1944) recorded M. carolina from Louisiana without spe-
cific locality data. This species is not found in the YD, but is widely
distributed throughout the rest of the study area, and is abundant in the
CP. Specimens were collected near a wide variety of aquatic habitats,
including springs, small streams, and impoundments.

Macronema transversa (Walker). Site 57. 20 June. This record is based on
two female specimens (det. D. G. Denning). The species was previously
known only from Georgia and Indiana.

Potamyia flava (Hagen). Sites 4, 5, 6, 8, 15, 16, 17, 19, 21, 23, 32, 40, 41,
BA AS AQ SONI 2, 7s O25805,° 1 19 eo ls BO Or gL. AEapru—LS
October. Very common along larger rivers, and particularly abundant
along the Mississippi River.

Symphitopsyche sparna (Ross). Sites 2, 57, 65, 66, 71, 79, 80, 81, 83, 84,
87, 88. 29 March—29 Sept. This uncommon species seems to be restricted
to the CP in the study area. We collected specimens near a variety of
aquatic habitats, including small sandy bottomed streams, spring-fed im-
poundments, and black-water rivers.

ACKNOWLEDGMENTS

In addition to the specimens collected during the surveys mentioned
above, many records were obtained from Bryant Mather who has collected
in various portions of Mississippi for many years. These records were of
great value in defining the distribution of each species in the state, and we
wish to thank Mr. Mather for making them available to us.

John Morse was of particular assistance throughout our work both by
identifying specimens or verifying our identifications and by acting as a
sounding board for our ideas. Michael Poirrier also offered valuable assis-
tance with various aspects of this study. Several other individuals assisted
us by making or verifying identifications, or supplying distributional records,
including Robert Blickle, Jay Chapin, Donald Denning, David Etnier, Oliver
S. Flint, Jr., Steven Hamilton, Ken Manuel, and John Unzicker.

Paul Hartfield, Bill Stark, and Ed Zuccaro supplied us with many speci-
mens. We thank the host of graduate and undergraduate students from the
University of Mississippi and University of New Orleans who assisted with
various aspects of fieldwork.

LITERATURE CITED

Berner, L. 1977. Distributional patterns of southeastern mayflies (Ephemeroptera). Bull. Fla.
State Mus. Biol. Soc. 22: I-56.

Blickle, R. L. 1962. Hydroptilidae (Trichoptera) of Florida. Fla. Entomol. 45: 153-155.

Denning, D. G. 1949. New species of Nearctic caddis flies. Bull. Brooklyn Entomol. Soc. 44:
37-48.

Edwards, S. W. 1973. Texas caddisflies. Tex. J. Sci. 24: 491-516.

|

508 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Etnier, D. A. and G. A. Schuster. 1979. An annotated list of Trichoptera (caddisflies) of
Tennessee. J. Tenn. Acad. Sci. 54: 15-22.

Flint, O. S., Jr. 1972. Three new caddisflies from the southeastern United States. J. Ga.
Entomol. Soc. 7: 79-82.

Flint, O. S., Jr., J. R. Voshell, Jr., and C. R. Parker. 1979. The Hydropsyche scalaris group
in Virginia, with the description of two new species (Trichoptera: Hydropsychidae).
Proc. Biol. Soc. Wash. 92: 837-862.

Gordon, A. E. 1974. A synopsis and phylogenetic outline of the Nearctic members of Cheu-
matopsyche. Proc. Acad. Nat. Sci. Phila. 129: 117-160.

Holzenthal, R. W. 1980. A survey of the caddisflies (Insecta: Trichoptera) of southeastern
Louisiana and southern Mississippi. MS thesis, Univ. of New Orleans. 75 pp.

Hudson, P. L., J. C. Morse, and J. R. Voshell, Jr. 1981. The larva and pupa of Cernotina
spicata (Trichoptera: Polycentropodidae). Ann. Entomol. Soc. Am. 74: 516-519.

Lowe, E. N. 1919. Mississippi: its geology, geography, soils and minerals resources. Miss.
State Geol. Surv. Bull. 14: 1-345.

Morse, J. C. 1970. Caddisflies, or Trichoptera, from northwestern South Carolina. MS thesis,
Clemson Univ. 56 pp.

——. 1972. The genus Nyctiophylax in North America. J. Kans. Entomol. Soc. 45: 172-
181.

Morse, J. C., J. W. Chapin, D. D. Herlong, and R. S. Harvey. 1980. Aquatic insects of Upper
Three Runs Creek, Savannah River Plant, South Carolina. Part 1: Orders other than
Diptera. J. Ga. Entomol. Soc. 15: 73-101.

Parker, C. R. and J. R. Voshell, Jr. 1981. A preliminary checklist of the caddisflies (Trichop-
tera) of Virginia. J. Ga. Entomol. Soc. 16: 1-7.

Resh, V. H. 1975. A distributional study of the caddisflies of Kentucky. Trans. Ky. Acad.

Sci. 36: 6-16.

Ross, H. H. 1944. The caddisflies, or Trichoptera, of Illinois. Ill. Nat. Hist. Sur. Bull. 23(1):
1-326.

—. 1967. The evolution and past dispersal of the Trichoptera. Annu. Rev. Entomol. 12:
169-206.

Ross, H. H. and T. Yamamoto. 1965. New species of the caddisfly genus Polycentropus from
eastern North America (Trichoptera, Psychomyiidae). Proc. Biol. Soc. Wash. 78: 241-
245.

Schuster, G. A. and D. A. Etnier. 1978. A manual for the identification of the larvae of the
caddisfly genera Hydropsyche Pictet and Symphitopsyche Ulmer in Eastern and Central
North America (Trichoptera: Hydropsychidae) U.S. Environ. Prot. Agency Rep. 600/4-
78-060. 128 pp.

Stanford, D. F. 1980. The immature dragonflies (Odonata: Anisoptera) of north Mississippi.
MS thesis, Univ. of Mississippi. 98 pp.

Unzicker, J. D., L. Aggus, and L. O. Warren. 1970. A preliminary list of the Arkansas
Trichoptera. J. Ga. Entomol. Soc. 5: 167-174.

Unzicker, J. D., V. H. Resh, and J. C. Morse. Jn press. Trichoptera, Chapter 9. Jn Brigham,
A. R., W. U. Brigham, and A. Gnilka, eds., Aquatic Insects and Oligochaetes of North
and South Carolina. Midwest Aquatic Enterprises, Mahomet, III.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 509-512

AN ANNOTATED CHECKLIST OF THE CADDISFLIES
(TRICHOPTERA) OF MISSISSIPPI AND
SOUTHEASTERN LOUISIANA. PART II:

RHYACOPHILOIDEA

STEVEN C. HARRIS, PAUL K. LAGO, AND RALPH W. HOLZENTHAL!

(SCH) Environmental Division, Geological Survey of Alabama and De-
partment of Biology, University of Alabama, University, Alabama 35486;
(PKL) Department of Biology, University of Mississippi, University, Mis-
sissippi 38677; (RWH) Department of Entomology, Fisheries and Wildlife,
Clemson University, Clemson, South Carolina 29631.

Abstract.—Distributional records for 30 species of caddisflies in the su-
perfamily Rhyacophiloidea (Glossosomatidae, Hydroptilidae, Rhyacophili-
dae) are presented, and the seasonal occurrence of each is indicated. An-

- notations include information on species habitat and relative abundance. All

26 of the species reported from Mississippi are new state records, and all
but eight species are newly recorded from Louisiana.

This paper is the second of a three part series dealing with the Trichoptera
of Mississippi and southeastern Louisiana. Representatives of all three fam-
ilies in the superfamily Rhyacophiloidea were collected during the surveys,
but only members of the Hydroptilidae were common and represented by
more than a single species. This superfamily was more poorly represented
than the Hydropsychoidea of Part I and the Limnephilodea of Part III in
the study area.

List OF SPECIES

The Rhyacophiloidea were represented in the study area by 30 species in
eight genera and three families. Of the reported 30 species, 28 belong to the
Hydroptilidae and one each to the Rhyacophilidae and Glossosomatidae.
Only eight species, all hydroptilids, were previously reported from the study
area: Hydroptila waubesiana Betten, Neotrichia minutisimella (Chambers),
Oxyethira glasa Ross, O. verna Ross, and O. zeronia Ross (as O. walteri)

es

|

1 Address reprint requests to junior author (PKL).

510 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

by Denning (1947a); Orthotrichia cristata Morton by Denning (1947b); Ox-
yethira janella Denning by Denning (1948); and Orthotrichia aegerfasciella
(Chambers) (as O. americana Banks) by Kingsolver and Ross (1961).

Collecting sites and seasonal occurrence designations in the following list
follow that outlined in Part I. Since most of these species, particularly within
the Hydroptilidae, have poorly known distributions, those with significant
range extensions are indicated.

Glossosomatidae

Protoptila lega Ross. Sites 35, 57, 71, 80, 81. 23 May—13 Oct. Locally com-
mon. Collected along fast, clear streams with sand-gravel substrates. At
several of the localities aquatic mosses and algae were encrusted on the
rocks of the rapids.

Hydroptilidae

Hydroptila armata Ross. Sites 24, 60. 27 March, 8 May. Uncommon. Col-
lected near swift, sand-bottom streams.

Hydroptila berneri Ross. Site 127. 20 March. A single specimen was col-
lected along a small blackwater stream.

Hydroptila grandiosa Ross. Sites 60, 61. 7 Feb., 11 April. Rare. Collected
along small, spring-fed streams. Primarily known from the northeastern
and northcentral United States (Blickle, 1979).

Hydroptila gunda Milne. Sites 57, 61, 67. 11 April, 20 June, | Aug. Uncom-
mon. Collected along Coastal Plain (CP) streams with moderately fast
flow and sand-gravel bottoms.

Hydroptila hamata Morton. Sites 34, 60, 125. 30 Jan., 7 Feb., 1 April, 10
May. Occasional but probably widespread in study area. Collected along
a variety of streams.

Hydroptila molsonae Blickle. Sites 99, 127. 20 March, 22 May. Uncommon.
Collected along CP blackwater streams with abundant vegetation. Only
known previously from the type-locality in Florida (Blickle, 1962).

Hydroptila n. sp. Sites 2, 38, 67, 71, 80, 81. 8 Sept., 1 Aug.—13 Oct. Com-
mon. Widely distributed in study area, usually occurring near clear, cool,
fast-flowing streams. This species is being described by the junior author
(RWH).

Hydroptila novicola Blickle and Morse. Site 97. 7 June. Uncommon. Col-
lected near a small, sand-bottom blackwater stream. Beds of Vallisneria
were abundant in stream. Primarily known from the northeastern United
States (Blickle, 1979).

Hydroptila quinola Ross. Sites 24, 37, 60, 83, 86, 97, 103. 7 Feb., 7-28 May,

10 June. Common. Widely distributed in study area occurring along a
variety of streams.

VOLUME 84, NUMBER 3 511

Hydroptila remita Blickle and Morse. Sites 37, 61, 120, 127. 11 April, 7-31
May, 17 Oct. Occasional. Collected near slow- to moderate-flowing
streams primarily on the CP.

Hydroptila strepha Ross. Site 97. 7 June. Uncommon. Collected with H.
novicola. Previously known only from northeastern United States (Blic-
kle, 1979).

Hydroptila tusculum Ross. Sites 83, 122, 129. 9 April, 5-10 June. Uncom-
mon. Associated with small, spring-fed blackwater streams. Previously
only known from the type locality in Tennessee (Ross, 1947).

Hydroptila waubesiana Betten. Sites 51, 79, 86. 23 May, 9 June, 23 June.
Uncommon. This widespread eastern species was collected near clear,
fast-flowing streams with sand-gravel substrates.

Mayatrichia ayama Mosely. Sites 95, 97. 7 June. Locally common. This
widespread North American species was collected near small, moderately
flowing blackwater streams with sand substrates.

Neotrichia minutisimella (Chambers). Site 40. 23 Aug., 10 Oct. Locally
common. Collected along the Mississippi River.

Neotrichia vibrans Ross. Sites 71, 95. 7 June, 30 Aug. Locally common.
Collected along small blackwater streams with moderate flow and sand-
gravel bottoms.

Ochrotrichia tarsalis (Hagen). Sites 2, 57, 67, 116. 20 June—20 Oct. Com-
mon. Collected near clear, fast-flowing streams with sand-gravel bottoms.
This species as well as the two preceding are fairly widespread in the
eastern United States (Blickle, 1979).

Orthotrichia aegerfasciella (Chambers). Sites 2, 16, 24, 25, 37, 40, 41, 49,
66,71, 103, 126, 129, 131..25 Apnl—5 June, 16 July, 30 Aug—10 Get:
Common. Widely distributed in study area associated with a variety of
streams and lakes.

Orthotrichia cristata Morton. Sites 31, 66. 28 May, 31 Aug. Uncommon.
Collected near spring-fed, sand-bottom lakes. As with O. aegerfasciella,
this is a widespread eastern species (Blickle, 1979).

Orthotrichia curta Kingsolver and Ross. Site 129. 5 June. Rare. Collected
near a small, spring-fed blackwater stream with little flow. Previously
only known from the type-locality in Florida (Blickle, 1962).

Orthotrichia dentata Kingsolver and Ross. Site 103. 26 May. A single spec-
imen was collected along a large, swift-flowing blackwater stream of the
CP. Previously only known from the type-locality in Florida (Blickle,
1962).

Oxyethira glasa (Ross). Sites 86, 99, 102, 129. 22 May-S June. Locally
common. Collected primarily along small, slow-moving blackwater
streams of the lower CP.

Oxyethira janella Denning. Sites 71, 106, 129. 26 May, 5 June, 30 Aug.

2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Locally common. Collected from the CP near blackwater streams of var-
ious sizes and flow regimes.

Oxyethira novasota Ross. Sites 28, 37, 61, 83, 86, 129. 11 April, 7-23 May,
5-10 June. Locally common. Widespread in study area occurring along
a variety of streams.

Oxyethira pallida (Banks). Sites 25, 106, 129. 6 May, 26 May, 5 June. Lo-
cally common. This widespread North American species was collected
from both clear and blackwater streams of varying size and flow regimes.

Oxyethira sininsigne Kelley. Site 129. 5 June. Rare. Collected along a small,
spring-fed blackwater stream with little flow.

Oxyethira verna Ross. Site 132. 14 May. Locally common.

Oxyethira zeronia Ross. Sites 31, 99, 102. 22-24 May, 31 Aug. Locally
common. This widespread eastern species was associated with a variety
of aquatic habitats including a lake, a marsh, and a swift-flowing, sand-
bottom stream.

Rhyacophilidae

Rhyacophila glaberrima Ulmer. Site 2. 8 Sept. Rare. This primarily northern
species was collected along a small, fast-flowing stream with rocky bottom
in the Tennessee River Hills (TRH) region.

ACKNOWLEDGMENTS

We thank Steve Hamilton of Clemson University for his review of the
paper. We also thank Irene Thompson for her typing and editing of the
manuscript.

LITERATURE CITED

Blickle, R. L. 1962. Hydroptilidae (Trichoptera) of Florida. Fla. Entomol. 45: 153-155.

. 1979. Hydroptilidae (Trichoptera) of America north of Mexico. N. H. Agric. Exp.

Stn. Bull. 509: 1-97.

Denning, D. G. 1947a. Hydroptilidae (Trichoptera) from southern United States. Can. Ento-
mol. 79: 12-20.

—. 1947b. New species and records of North American Hydroptilidae (Trichoptera).

Psyche (Camb. Mass.) 54: 170-177.

. 1948. New species of Trichoptera. Ann. Entomol. Soc. Am. 41: 397-401.

Kingsolver, J. M. and H. H. Ross. 1961. New species of nearctic Orthotrichia (Hydroptilidae,
Trichoptera). Ill. St. Acad. Sci. Trans. 54: 28-33.

Ross, H. H. 1947. Descriptions and records of North American Trichoptera, with synoptic
notes. Trans. Am. Entomol. Soc. 73: 125-168.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 513-520

AN ANNOTATED CHECKLIST OF THE CADDISFLIES
(TRICHOPTERA) OF MISSISSIPPI AND
SOUTHEASTERN LOUISIANA. PART III:
LIMNEPHILOIDEA AND CONCLUSIONS!

RALPH W. HOLZENTHAL, STEVEN C. HARRIS, AND PAUL K. LAGO?

(RWH) Department of Entomology, Fisheries and Wildlife, Clemson Uni-
versity, Clemson, South Carolina 29631; (SCH) Environmental Division,
Geological Survey of Alabama and Department of Biology, University of
Alabama, University, Alabama 35486; (PKL) Department of Biology, Uni-
versity of Mississippi, University, Mississippi 38677.

Abstract.—Distributional records for 63 species of caddisflies in the su-
perfamily Limnephiloidea (Phryganeidae, Brachycentridae, Limnephilidae,
Lepidostomatidae, Sericostomatidae, Odontoceridae, Molannidae, Helico-
psychidae, Calamoceratidae, Leptoceridae) from Mississippi and southeast-
ern Louisiana are presented and seasonal occurrence of each is indicated.
Annotations for many species include notes on habitat and relative abun-
dance. Of the species reported, 58 represent new state records for Missis-
sippi and 60 are newly recorded from Louisiana. Faunal composition and
distribution of the order in the study area are discussed.

This is the third and final contribution in a three part series on the cad-
disflies of Mississippi and southeastern Louisiana. In this section records
are presented for 63 species in 22 genera from the families Phryganeidae,
Brachycentridae, Limnephilidae, Lepidostomatidae, Sericostomatidae,
Odontoceridae, Molannidae, Helicopsychidae, Calamoceratidae, and Lep-
toceridae. Sixty-two percent of the species reported belong to the Lepto-
ceridae.

Literature records of Limnephiloidea from Mississippi and Louisiana are
very few and scattered, and include the following: Hagen (1861) described
Platycentropus amicus (as Hallesus) from New Orleans, Louisiana; Ross

1 Technical Contribution No. 1997 of the South Carolina Agricultural Experiment Station,
Clemson University.
2 Address reprint requests to junior author (PKL).

514 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(1962) described Agarodes stannardi (as Sericostoma) from Marshall Co.,
Mississippi; Flint (1972) described the male of Jronoquia kaskaskia (Ross)
(as I. brysoni) from Oktibbeha Co., Mississippi; Ross and Scott (1974) de-
scribed Agarodes libalis from Jackson Parish, Louisiana; and Haddock
(1977) recorded Nectopsyche spiloma (Ross) from Louisiana and Mississip-
pi and N. candida (Hagen) and N. pavida (Hagen) from Mississipp1.

Additional records indicated in the text were obtained from the Louisiana
State University Collection (LSUC). Collection site numbers correspond to
those listed in Part I.

LIST OF SPECIES
Phryganeidae

Agrypnia vestita (Walker). Sites 2,9, 10, 13, 14, 16, 19, 22, 23, 44, 90, 100.
7 Apr.—12 July, 11 Sept.-24 Oct. Fairly common in the northern half of
the study area.

Ptilostomis ocellifera (Walker). Sites 3, 16. 16 June, 2 July, 26 July. Un-
common.

Ptilostomis postica (Walker). Sites 3, 4, 16, 18, 19, 23, 26, 85, 104. Addi-
tional record from Baton Rouge, East Baton Rouge Parish, LA (LSUC).
16 Apr.-14 July, 27 Aug.—27 Sept. Collected near rocky streams, muddy
ponds and sand-bottomed reservoirs. Widespread but not common.

Brachycentridae

Brachycentrus numerosus (Say). Larvae collected from sites 81, 123, 125.
Adult records from Magnolia, Plaquemines Parish, LA, 28 Feb., (LSUC).
This species is fairly common and widespread in the CP. Larvae were
collected from submerged logs and aquatic vegetation in strong currents
of clear, fast flowing streams. This was the earliest emerging caddisfly in
the study area.

Micrasema wataga Ross. Sites 71, 92, 97, 108, 120. 25 May—30 Aug. Wide-
spread in the CP but not common. Most often collected near clear, cool,
blackwater streams having abundant aquatic vegetation.

Limnephilidae

Tronoquia kaskaskia (Ross). Site 23. Additional record from Fluker, Tan-
gipahoa Parish, LA (LSUC). 14 Oct.—7 Nov. Rare.

Tronoquia punctatissima (Walker). Sites 6, 7, 10, 11, 13, 16, 19, 23, 44, 116.
9 Sept.-8 Nov. Widespread but not common. Collected along a wide
variety of aquatic habitats.

Limnephilus prob. submonilifer Walker. Site 7. A single small larva was
collected in May. Our identification is tentative until we can associate it
with adults. L. submonilifer is the only Limnephilus recorded from the

VOLUME 84, NUMBER 3 515

neighboring states of Arkansas (Unzicker et al., 1970) and Tennessee
(Etnier and Schuster, 1979). The larva was described by Flint (1960).

Platycentropus amicus (Hagen). The female holotype is from New Orleans.
Ross (1938) described the male (as P. plectrus) from Michigan and Wis-
consin. This rare species has not been collected from the study area since
Hagen’s time.

Platycentropus radiatus (Say). Based on a single male from Moss Point,
Hancock Co., Miss. 26 May.

Pycnopsyche indiana (Ross). Site 127. 17 Oct. Rare. Collected from a small,
spring-fed, black-water stream.

Pycnopsyche lepida (Hagen). Site 23. 22 Oct.—28 Oct.

Pycnopsyche luculenta (Betten). Sites 11, 14, 16, 17. 1 Oct.-—30 Oct. Col-
lected along small streams, ponds and bogs.

Pycnopsyche scabripennis (Rambur). Sites 10, 11, 13, 14, 17, 44, 103, 127.
1 Oct.-30 Oct. The most common and widespread Pycnopsyche in the
study area, the majority of specimens were collected near small reser-
voirs.

Lepidostomatidae

Lepidostoma sp. Site 7. Based on larvae collected in May.

Sericostomatidae

Agarodes crassicornis (Walker). Sites 16, 21, 86, 89, 93, 96, 97, 98, 100,
102, 108, 114. 18 May-—30 June. Fairly common. Associated with cool
streams and springs.

Agarodes libalis Ross and Scott. Sites 86, 108, 120, 128. 23 May—4 June.
This CP endemic has a narrow emergence period in the spring of the year
and was always associated with spring-fed, black-water streams.

Agarodes stannardi (Ross). Sites 7, 16. 16 May, 21 May, 30 May. This rare
species is known only from the type-locality and vicinity and is associated
with cool, sandy springs.

| Odontoceridae

Psilotreta rufa Hagen. Sites 1,7, 17. 16 May, 21 May, 28 May. Not common.
Limited to cool, rocky springs. Previously known only from the Northeast
(Ross, 1944).

| Molannidae

Molanna blenda Sibley. Sites 7, 16, 124. 28 March-12 June. Larvae and
adults were collected from springs.
| Molanna tryphena Betten. Sites 70, 127, 128. 21 March, 29 March, 4 June.
Associated with CP black-water streams. Larvae were very common at
| sites 127 and 128.

516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Molanna ulmerina Navas. Sites 87, 125. 1 April, 25 Sept. Rare. The collec-
tion of adults in September suggest that this species may have two gen-
erations per year in the study area.

Helicopsychidae

Helicopsyche borealis (Hagen). Sites 38, 57, 58. 20 June, 22 Sept., 13 Oct.
In Mississippi larvae were collected in shallow, gravel expanses of clear,
well aerated streams.

Calamoceratidae

Anisocentropus pyraloides (Walker). Sites 39, 59, 60, 86, 97. 19 April—7
June. Collected near small, cool, black-water streams.

Leptoceridae

Ceraclea cancellata (Betten). Sites 33, 114. 17 April, 10 May.

Ceraclea diluta (Hagen). Site 100. 6 April. Collected near a shallow, sand-
bottomed pond.

Ceraclea flava (Banks). Sites 40, 44, 59, 63, 65, 73, 80, 83, 102. 23 May—20
June. Usually associated with large, slow moving rivers.

Ceraclea maculata (Banks). Sites 2;5,7, 10, 11,16, 17,203 233 253265373
33, 34, 37, 40, 44, 45, 51, 52, 56, 57, 58,.59,.62, 65, 67,69, 732 80neamaa
91; 92, 93, 95, 97, 100, 101; 106; 108; 117; 119; 121, 122, 126, 27
April-17 Oct. The most common and widespread Ceraclea in the study
area. Collected along a wide diversity of aquatic habitats. |

Ceraclea nepha (Ross). Sites 2, 11, 16, 19, 23, 25, 45, 59, 86. 18 April—27
Oct. Collected from a variety of habitats in the study area. Previously
known from the central states (Morse, 1975).

Ceraclea ophiodera (Ross). Sites 16, 40, 47, 58, 71, 79, 80, 83, 84, 97, 101,
117, 120. 23 May—22 Sept. Widespread and common in the southern half
of the study area.

Ceraclea protonepha Morse and Ross. Sites 2, 11, 16, 24, 25, 28, 34, 66,
86. 19 April-27 June. Specimens were commonly collected near small, |
sandy streams. |

Ceraclea resurgens (Walker). Sites 122, 124, 125. 28 March, | April, 9 April.
This obligate sponge feeder was always associated with medium-sized, |
black-water streams. Adult emergence period was very brief.

Ceraclea slossonae (Banks). Site 51. 27 May.

Ceraclea spongillovorax (Resh). Sites 7, 118, 128. 6 Aug., 18 Sept., 1 Oct. |
Several specimens were collected near a small, spring-fed reservoir.

Ceraclea tarsipunctata (Vorhies). Sites 5, 6, 7, 15, 16, 19, 23, 26, 28, 29,
32, 33, 36, 44, 49, 51, 59, 63, 73, 80, 85, 88, 93, 100, 115, 132. 29 March—
11 Nov. Widespread and common across the study area. There is prob-
ably more than a single generation per year for this species in our area.

VOLUME 84, NUMBER 3 517

Ceraclea transversa (Hagen). Sites 28, 84. 9 May, 16 June. Collected from
small, sandy streams. Rarely encountered.

Leptocerus americanus (Banks). Sites 16, 24, 25, 32, 44, 51, 52, 66, 110,
120, 126. 19 April-31 May. Usually associated with streams and ponds
having abundant beds of aquatic vegetation, especially Ceratophyllum.

Nectopsyche albida (Walker). Sites 16, 19, 23, 32. 8 May—17 June.

Nectopsyche candida (Hagen). Sites 2, 7, 16, 17, 23, 40, 43, 44, 45, 51, 52,
SGMO TOU Nol y 6252909925 100; 103; 105,12, 21450115, M6; 95 1202126,
127, 129, 131. 17 April—-20 Oct. Widespread and common.

Nectopsyche exquisita (Walker). Sites 2, 16, 35, 38, 49, 57, 59, 71, 73, 87,
91, 110. 21 May—13 Oct. As widespread but somewhat less common than
N. candida in the study area.

Nectopsyche pavida (Hagen). Sites 2, 51, 57, 58, 61, 62, 67, 80, 81, 90, 92,
119. 11 April—29 Sept. Collected mostly near CP streams.

Nectopsyche spiloma (Ross). Site 51. 7 July.

Oecetis avara (Banks). Sites 83, 84, 87. 10 June, 16 June, 10 Sept., 20 Sept.
Rarely encountered.

Wecetis cinerascens (Hagen). Sites 2, 8, 10, 16, 20, 25, 30, 31, 32, 35, 37,
44, 50, 51, 52, 65, 66, 73, 87, 114, 126, 132. 11 April-8 Oct. Collected near
small reservoirs and a variety of stream types throughout the study area.

Oecetis daytona Ross. Sites 91, 102. 24 May, 5 Aug. An apparent CP en-
demic. Previously known only from the type-locality in Florida (Ross,

im 1947).

Wecetis ditissa Ross. Sites 2, 5, 11, 13, 16, 24, 25, 28, 34, 40, 49, 62, 63, 83,
fe 84.'85, 86, 100; 127, 128. 15 April-17 Oct. Widespread and common.
Oecetis georgia Ross. Sites 118, 127. 6 Aug.—17 Oct. A rare CP endemic.
Collected only along spring-fed, black-water streams. Known previously

from Georgia (Ross, 1941).

Mecetis inconspicua (Walker). Sites 2, 4, 9, 10, 11, 13, 14, 16, 17, 18, 19,
POMP 238 244255. 265 285 3003's 32533; 35):36 540; 43, 44549} SOS 59,
62, 63, 65, 66, 68, 69, 71, 73, 79, 80, 81, 82, 83, 84, 85, 86, 87, 88, 90, 92,
BS /995 1001027103): 107; 1093 110;-111, 112; 1145 11S p6; 120125 126,

127, 128, 129, 131, 132. 15 March-21 Nov. One of the most common

| caddisflies occurring in the study area.

Oecetis nocturna Ross. Sites 2, 24, 26, 34, 83, 114, 127. 8 May-17 Oct.
Taken from a variety of habitats including reservoirs and small, sandy
streams.

Oecetis osteni Milne. Sites 7, 16, 28, 37, 56, 62, 66, 69, 71, 90, 92, 93, 95,
97, 100, 101, 102, 103, 126, 129, 131. S April—18 Oct. Although widespread,
this species was more common in the CP.

Oecetis persimilis (Banks). Sites 2, 14, 28, 34, 52, 61, 77, 79, 83, 84, 91, 92,

| 97, 122, 126. 9 April-13 Oct. The distribution and abundance of this

species in the study area is similar to those of O. osteni.

518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Oecetis sphyra Ross. Sites 60, 67, 78, 80, 81, 83, 90, 93, 95, 97, 103, 117,
120, 127, 128. 23 May—18 Oct. Limited in distribution to the CP.

Setodes dixiensis Holzenthal. Sites 61, 79, 80, 117, 118. 11 April—6 Aug.
Associated with cool, clear, fast flowing, medium-sized streams in the
GP:

Triaenodes abus Milne. Sites 19, 34, 122.9 April, 10 May, 12 May. Collected
near small, sandy streams. Most members of this genus were rare in our
area and records for many species represent rather significant range ex-
tensions.

Triaenodes flavescens Banks. Sites 38, 87. 19 Sept.—13 Oct.

Triaenodes ignitus (Walker). Sites 16, 18, 29, 51, 57, 60, 69, 79, 80, 84, 97,
101, 103, 122, 127, 128. 9 April—28 July, 17 Oct. Common and taken along
a variety of aquatic habitats but never in large numbers.

Triaenodes inflexus Morse. Site 16. 14 June. Very rare. Our specimens were
collected rather far from water. Previously known only from South Car-
olina (Morse, 1971; Morse et al., 1980).

Triaenodes marginatus Sibley. Sites 11, 14. 3 May, 30 May, 17 July. A
widespread eastern species. Specimens were collected near a shallow,
sandy stream and a muddy stream.

Triaenodes melacus Ross. Sites 28, 30. 8 May, 9 May. Collected along a
small stream and a sand bottomed reservoir. Previously known only from
Illinois (Ross, 1947).

Triaenodes nox Ross. Site 29. 9 May. These specimens were collected near
a small farm pond. Very rare.

Triaenodes ochraceus (Betten and Mosely). Sites 24, 29, 34, 86, 100. 19
April-18 May. Rare. Formerly known only from Burke Co., Georgia and
Aiken Co., South Carolina (Morse et al., 1980). Our specimens were
collected near small, sandy streams and small ponds.

Triaenodes pernus Ross. Sites 11, 14, 29, 69, 92, 129, 131. 9 May-17 July, |
18 Sept. Fairly common near small to medium-sized sandy streams. '

Triaenodes smithi Ross. Sites 14, 33, 44, 79. 10 May—23 June. Reported |
previously from Illinois (Ross, 1959). We collected specimens in an area |
of spring-fed ponds, and near small sandy streams.

CONCLUSIONS

In our three part checklist we recorded a total of 139 species of caddisflies
from 43 genera and 17 families from Mississippi and southeastern Louisiana.
Most of the more than 450 species of Southeastern caddisflies occur in areas |
of high topographic relief, particularly in the southern Appalachian region, |
where lotic habitats include large, meandering rivers, torrential mountain
streams, cool springs and seeps. The majority of streams in the study area |
have sand and gravel bottoms, low gradients and warm waters. These con- |
ditions result in fewer microhabitats available to many species and limit the |

VOLUME 84, NUMBER 3 ail)

occurrence of cool-adapted species. It is not surprising then that the families
Leptoceridae, Hydropsychidae, Hydroptilidae, and Polycentropodidae ac-
counted for 76% of the species reported (28%, 20%, 15% and 13% respec-
tively). Species in these families, especially the Leptoceridae, are noted for
their ability to exploit warm water habitats (Wiggins, 1977). This factor in
large part accounts for the high numbers of species from these four families
in our area.
The generally cool-adapted families Philopotamidae, Psychomyiidae,
~Glossosomatidae, Phryganeidae, Brachycentridae, and Limnephilidae were
represented by species in genera with members secondarily adapted to warm
waters, e.g. Chimarra, Lype, Protoptila, Agrypnia, Ptilostomis, Brachy-
centrus, Micrasema, Ironoquia, and Pycnopsyche (Wiggins, 1977). Species
from other cool-adapted families, including Rhyacophila glaberrima (Rhy-
acophilidae), Psilotreta rufa (Odontoceridae), and Lepidostoma sp. (Lepi-
_dostomatidae) were restricted to the extreme northern part of the NCP or
the TRH where cooler streams are more frequent. Species in the Sericos-
~tomatidae, Molannidae, and Calamoceratidae were almost always associ-
ated with cool springs and spring-fed streams, especially in the CP.
Approximately 35 southeastern caddisfly species appear to be restricted
in distribution to the Gulf and Atlantic Coastal Plains where they usually
are associated with black-water streams. Morse et al. (1980) recorded many
of these species and Ross and Scott (1974) and Kelley and Morse (1982)
noted the endemicity of several species of Agarodes and Oxyethira, re-
spectively, to this region of North America. In the study area these coastal
plain endemics included, Chimarra florida, Neureclipsis melco, Cheuma-
_topsyche petersi, Cheumatopsyche virginica, Hydropsyche elissoma, Hy-
droptila molsonae, Orthotrichia curta, Orthotrichia dentata, Oxyethira ja-
nella, Oxyethira novasota, Oxyethira sininsigne, Agarodes libalis, Oecetis
daytona, Oecetis georgia, and Setodes dixiensis.
A few typically northern species were collected only from the northern
half of the NCP an the TRH and included Hydropsyche simulans, Rhyaco-
phila glaberrima, Ptilostomis ocellifera, Limnephilus prob. submonilifer,
| Pycnopsyche lepida, Pycnopsyche luculenta, Lepidostoma sp., Psilotreta
rufa, and Triaenodes marginatus.
Only Neotrichia minutisimella, a widespread eastern species, was re-
stricted to the YD, where it was collected along the Mississippi River.

ACKNOWLEDGMENTS

We thank John Weaver of Clemson University for his review of the manu-
script. We are grateful to Eileen McEwan, Savannah River Ecology Lab,
for providing larval material from Wall Doxey State Park; Ken Manuel,
Duke Power Co., for the record of Platycentropus radiatus; and Joan Chap-
in, Louisiana State University, for the records from the LSUC.

|

520 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

LITERATURE CITED

Etnier, D. A. and G. A. Schuster. 1979. An annotated list of Trichoptera (Caddisflies) of
Tennessee. J. Tenn. Acad. Sci. 54: 15-22.

Flint, O. S., Jr. 1960. Taxonomy and biology of Nearctic limnephilid larvae (Trichoptera),
with special reference to species in eastern United States. Entomol. Am. 40: 1-117.

———. 1972. Three new caddisflies from the southeastern United States. J. Ga. Entomol.
Soc. 7: 79-82.

Haddock, J. D. 1977. The biosystematics of the caddis fly genus Nectopsyche in North Amer-
ica with emphasis on the aquatic stages. Am. Midl. Nat. 98: 382-421.

Hagen, H. A. 1861. Synopsis of the Neuroptera of North America, with a list of the South
American species. Smithson. Misc. Collect., 347 pp.

Kelley, R. W. and J. C. Morse. 1982. A key to females of the genus Oxyethira (Trichoptera:
Hydroptilidae) from the southern United States. Proc. Entomol. Soc. Wash. 84: 256—
269.

Morse, J. C. 1971. New caddisflies (Trichoptera) from the southeastern United States. J. Ga.

Entomol. Soc. 6: 77-84.

. 1975. A phylogeny and revision of the caddisfly genus Ceraclea (Trichoptera, Lep-

toceridae). Contrib. Am. Entomol. Inst. (Ann Arbor) 11: 1—97.

Morse, J. C., J. W. Chapin, D. D. Herlong, and R. S. Harvey. 1980. Aquatic insects of Upper
Three Runs Creek, Savannah River Plant, South Carolina. Part I: Orders other than
Diptera. J. Ga. Entomol. Soc. 15: 73-101.

Ross, H. H. 1938. Descriptions of Nearctic caddis flies (Trichoptera) with special reference
to the Illinois species. II]. Nat. Hist. Surv. Bull. 21: 101-183.

—. 1941. Descriptions and records of North American Trichoptera. Trans. Am. Entomol.
Soc. 67: 35-126.

——. 1944. The caddis flies, or Trichoptera, of Illinois. Ill. Nat. Hist. Surv. Bull. 23: 1-326.

—. 1947. Descriptions and records of North American Trichoptera, with synoptic notes.

Trans. Am. Entomol. Soc. 73: 125-168.

. 1959. The relationships of three new species of Triaenodes from Illinois and Florida.

Entomol. News 70: 39-4S.

—. 1962. Three new species of Trichoptera from eastern North America. Entomol. News
73: 129-133.

Ross, H. H. and D. C. Scott. 1974. A review of the caddisfly genus Agarodes, with descriptions
of new species (Trichoptera: Sericostomatidae). J. Ga. Entomol. Soc. 9: 147-155.
Unzicker, J. D., L. Aggus, and L. O. Warren. 1970. A preliminary list of the Arkansas

Trichoptera. J. Ga. Entomol. Soc. 5: 167-174.

Wiggins, G. B. 1977. Larvae of the North American caddisfly genera (Trichoptera). Univ. of |

Toronto Press, Toronto. 401 pp.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 521-528

A DESCRIPTION OF THE LARVA OF DIXELLA NOVA (WALKER)
(DIPTERA:DIXIDAE)!

T. MICHAEL PETERS AND DAVID ADAMSKI

| Department of Entomology, University of Massachusetts, Amherst, Mas-
sachusetts 01003; (DA) current address: Department of Entomology, Mis-
_sissippi State University, Mississippi 39762.

Abstract.—The mature fourth instar of Dixella nova (Walker) is described
and illustrated. Terms applied to larval dixid structures by authors of earlier
| descriptions are synonymized with those used in this paper.

All 44 species of Dixidae known to occur in the Nearctic Region have
been described in the imaginal stage (Peters and Cook, 1966; Peters and
Barbosa, 1970; Peters, 1981b). Knowledge of immature stages is sparse;

only nine species have been described or illustrated (Johannsen, 1903, 1933;
Nowell, 1951; Peters and Barbosa, 1970; Smith, 1928).

Adult dixids spend the majority of their time concealed in habitats along
the edges of water dominated by overhanging vegetation. Hence, they are
seldom taken by the general collector. However, adult males form swarms
in late afternoon and evening along the water’s edge and can be collected
by wading along the shore.

Although a few immature dixids may be found in broad spectrum collec-
tions gathered for environmental impact statements, they are seldom col-
lected except by specialized techniques. The pupa is attached to emergent
vegetation or rocks above the waterline. The larva rests at the waterline
-with its body in an inverted U position so that only its head and caudal
respiratory apparatus is in the meniscus. Thus, aquatic netting or use of

dippers seldom catch immature dixids. However, larvae in lotic waters
undergo diel drifting (Waters, 1962) and are caught by drift sampling tech-
niques. Both lentic and lotic forms may be washed into an aquatic net from
their positions on emergent objects and the water’s edge. If the net is emp-

' Research supported by project MS-27 of the Massachusetts Agricultural Experiment Sta-
tion. Published with the aid of a grant from the Guy Chester Crampton Research Fund of the
University of Massachusetts.

522 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tied into a white pan full of water, the larvae soon swim to the surface and
may be picked out.

At present, the only way most larvae can be identified to species is by
rearing to the adult stage and then identifying them. However, most spec-
imens are preserved as they are caught and are thereby not identifiable to
species except by comparison with the few published larval descriptions.
Here too, a problem is encountered because of the diverse terminology used
for the morphological structures employed in the specific descriptions.

The purpose of this paper is twofold. The first objective is to stabilize the
terminology so that workers can use the previous larval descriptions with
some confidence that the structures being compared between articles are
the same. The second objective is to describe the mature larva of Dixella
nova (Walker) and to illustrate the structures that are of value in comparing
it to previously known larvae. The complete chaetotaxy is not presented
here, but it will be covered in a future paper comparing all the known larval
dixids from the Nearctic Region.

Description of the Mature Fourth Instar of Dixella nova (Walker)
Figss 10/225

Length 2.8-3.4 mm (N = 10). Head and caudal respiratory apparatus
medium brown, heavily sclerotized. Body cylindrical, predominantly slate
gray, including thoracic segments and abdominal segment 8. Abdominal
segments I-7 countershaded, with venter lighter gray. Mesothorax with 2
paired subcircular non-pigmented areas, one on each pleural area with a
second contiguous to it toward the venter. Metathorax with single non-
pigmented subcircular area on each side, slightly smaller than those of me-
sothorax.

Head as in Fig. 3A. Eye posterior to and slightly below insertion of an-
tenna, with heavy sclerotization of head capsule around dorsoanterior mar-
gin but very lightly sclerotized immediately below and behind eye, becoming
heavily sclerotized just anterior to seta 10 (following system of Belkin et al.,
1970). Seta 1 as long as mouth brushes but of greater diameter, moderately
sinusoidal, blunt-ended. Seta 3 shorter than mouth brushes, flattened, point-
ed, widest in middle. Antenna with numerous pointed terminal sensilla. A
very lightly sclerotized, pear-shaped sensillum located in depression bound-
ed by ring of pointed terminal sensilla. This sensillum often destroyed in
permanently mounted specimens (see insert in Fig. 3C).

Mandible as in Fig. 3A. Dorsal tooth between mandibular brush and seta
2. Mandibular brush tapering from long distal hair, shortest basally. Two
prominent, heavily sclerotized ventral teeth, the distal one largest. A row
of accessory teeth between inner ventral tooth and articulation. An elongate
tuft of branched, slender hairs forms the mandibular sweeper, arising from
basal edge of accessory tooth ridge, at 90° angle to row of accessory teeth

VOLUME 84, NUMBER 3 523

7

£3
pe

PROLEGS
with
CROCHETS

@

< ~
WE

al

*

Bi

\/
SS

L YT
VX ]
y
Kp

W
A

Na soon

a:

Bt eas
Wy AMBULATORY
AN COMB
A | 2
Fig. |. Mature fourth-instar of Dixella nova. A, Ventral view. B, Dorsal view.

(see insert of Fig. 3A). Lateral seta prominent. Maxilla as in Fig. 3A. Palp
antenniform.

Anterior ventral setae of prothorax prominent, extending beyond mouth
brushes of head, arranged as 1-1-4-1-1-4-1-1. Each group of 4 share a com-
mon sclerotized base.

524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

PARASPIRACULAR HAIRS

SPIRACLE

POSTSPIRACULAR Ws e )

PROCESS yp iy
METASPIRACULAR

PLATE -
DORSAL ANTEROMEDIAL ae

SETA Ye

MEDIAN PLATE

ANTEROLATERAL PLATE

AT

PECTEN

POSTEROLATERAL
PROCESS

‘1. \___veNTROLATERAL
ail PLATE
Iq \
KN }+———ana_ come

ine POSTANAL
i a) PROCESS
are

2

Fig. 2. Posterior abdominal structures of Dixella nova larva, slightly distended; left half
dorsal, right half ventral.

Venter of abdominal segments | and 2 each with a pair of prolegs, tipped
with anteriorly directed, uniordinal, triserial crochets (see insert, Fig. 1A).
Base of prolegs slate gray, lighter distally. Segments 5—7 with ambulatory
comb (see insert, Fig. 1A). Medial plate of comb subrectangular, postero-
lateral corners expanded. Spines of comb in 2 distinct rows, the smaller
slightly appressed to surface of segment, the stouter row at a less acute
angle to body surface.

Segment 8 simple ventrally, with row of 8 long stout setae. Dorsally a
row of multiple-branched paraspiracular setae arise from 4 bases, lateral
bases with 2 setae each, medial bases with | seta each. Metaspiracular plate
located medially between spiracles; subtriangular, with apex directed pos-

VOLUME 84, NUMBER 3 525

MANDIBULAR BRUSH

VENTRAL TEETH Pr

ACCESSORY ent

SETA: 2
DORSAL TOOTH

LATERAL SETA
MAXILLARY PALP

MANDIBULAR SWEEPER—4

MAXILLARY BRUSH

Fig. 3. Dixella nova. A, Left mandible, ventral view. B, Left maxilla, ventral view. C,
Head, left half dorsal, right half ventral.

teriorly, center of plate elevated into a slight, blunt peak. Posterior to each
spiracle is a laterally projecting, dorsoventrally flattened postspiracular pro-
cess fringed with articulated setae.

Segment 9 with paired anterolateral plates bearing a complex pecten on
the posterior margin; pecten of many small quadriradiate processes and a
single large triradiate spine near the most ventral apex (see insert, Fig. 2).
On dorsum a large median plate with slight sculpturing along mid-dorsal
area in center of plate. Sculpturing formed by a contiguous group of elongate
ovoids. Posterior medial margin of median plate with irregular row of very

|

526 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

short complex, multiple-branched spines. Median plate extends into a pair
of dorsoventrally flattened posterolateral processes fringed with articulated
setae. Each process terminates posteriorly in a spine directly beneath the
fringe. At base of each posterolateral process is a small dorsally projecting
tubercle bearing a long, dorsal anteromedial seta.

Segment 10 conical; sclerotized dorsally; venter lightly sclerotized, bear-
ing a pair of heavily sclerotized subtriangular ventrolateral plates, each with
3 very long posteriorly projecting setae. Just anterior to anus is a ventral
anal comb of stout posteriorly projecting spines. Anterior to anal comb is
an area of shorter delicate aculeae extending from one side of the segment
ventrally to the opposite side. Behind anus, postanal process is entirely
sclerotized with a sharp break in sclerotization at its narrowest diameter.
Expanded and sparsely covered with moderately long microtrichia posterior
to break. Caudal tip bears a posteriorly projecting spine on dorsum. Three
pairs of long posteriorly projecting setae at caudal end. One pair projects
dorsoposteriorly, another dorsolaterally, the 3rd posteroventrally.

TERMINOLOGY

Terms previous authors have used for morphological structures in de-
scriptions of larval dixids vary widely. The only attempts to standardize
terminology have been by Belkin (1962, 1968) when he interpreted dixid
morphology considering them to be a subfamily of the Culicidae. He syn-
onymized much of dixid morphology with that used for anopheline mos-
quitoes.

The terminology I apply to the larval head and mouthparts follows that
of Harbach and Knight (1980) for mosquitoes. However, I have not at-
tempted to use culicid terminology for the caudal and respiratory complex
of dixids since homologies are not certain. This is not surprising in light of
the morphological (Rohdendorf, 1964) and cytogentic (Frizzi et al., 1966)
evidence accumulating to support the separation of Dixidae from the Culic-
idae.

The following is a list of published names, by author, for those structures
described and illustrated in the D. nova description above. Each author’s
name appears only if the structure was assigned a name in the listed refer-
ences

Paraspiracular hairs or setae (=prespiracular hairs of Nowell, 1951, Smith,
1928).

Postspiracular process (=anterolateral lobes of Belkin, 1962, Contini, 1965;
=spiracular valves, plates, or appendages of Edwards, 1932, Smith, 1928,
Tonnoir, 1921-3, Brindle, 1963, Disney, 1975; =ear-shaped lobes of Now-
ell, 1951; =Seitenklappe of Martini, 1929).

Metaspiracular plate (=anterior median lobe of Belkin, 1962; =posterior

VOLUME 84, NUMBER 3 527

median plate of Contini, 1965; =chordate chitinized area of Smith, 1928;
=median plate of Peters, 198la; =interspiracular disc of Disney, 1975).
Median plate (=median plate of Belkin, 1962, Contini, 1965; =basal plate

of Brindle, 1963, Disney, 1975; =Mittelstiick of Martini, 1929).

Posterolateral process (=posterolateral lobes of Belkin, 1962, Contini, 1965;
=lateral plates of Tonnoir, 1921-3, Smith, 1928; =posterior articulated
paddle of Nowell, 1951; =posterior appendage of Brindle, 1963; =ciliated
lateral lobe of Johannsen, 1933; =large posterior valves of Edwards, 1932:
=Seitenklappe of Hennig, 1950; =Hinterklappe of Martini, 1929; =pos-
terior paddle of lateral plate of Disney, 1975).

Anterolateral plate with pecten (=pecten of Belkin, 1962, Johannsen, 1903,
Martini, 1929, Smith, 1928; =pectinate comb of Nowell, 1951).

Ventrolateral plate (=Lateralplatte of Hennig, 1950; =Subventralplatte of
Martini, 1929).

Anal comb (=supraanal spicules of Belkin, 1962; =guards for gill chamber
of Smith, 1928).

Postanal process (=tail-process of Johannsen, 1933, Edwards, 1932; =cau-
dal appendage of Tonnoir, 1921-3, Contini, 1965, Disney, 1975; =cap of
Nowell, 1951; =Schwanzfortsatz of Martini, 1929, Hennig, 1950; =anal
appendage of Brindle, 1963; =cylindrical caudal process of Belkin, 1962).

Ambulatory comb (=ambulacral plate of Nowell, 1951; =sclerotized pec-
tinate plate of Johannsen, 1933; =locomotory plate of Contini, 1965;
=spiney plate of Smith, 1928; =rows of spines of Belkin, 1962; =short,
hook-like spines of Brindle, 1963; =plate set with rows of hooks of Ed-
wards, 1932; =Fusstummel of Hennig, 1950; =ambulacral comb of Belk-
in, 1968; =ventral comb of Disney, 1975).

Prolegs with crochets (=pseudopods of Edwards, 1932, Contini, 1965;
=pseudopods with spines, hooks, or claws of Nowell, 1951, Brindle, 1963,
Smith, 1928; =prolegs of Johannsen, 1933; =prolegs with curved spines
of Belkin, 1962; =Kriechwulste of Hennig, 1950).

LITERATURE CITED

Belkin, J. N. 1962. The Mosquitoes of the South Pacific. Univ. of Calif., Press, Berkeley. 2
vols. 608 pp., 412 pls.

— . 1968. Mosquito studies (Diptera, Culicidae) VII. The culicidae of New Zealand. Con-
trib. Am. Entomol. Inst. (Ann Arbor) 3: 1-182.

Belkin, J. N., S. J. Heinemann, and W. A. Page. 1970. Mosquito studies (Diptera, Culicidae)
XXI. The culicidae of Jamaica. Contrib. Am. Entomol. Inst. (Ann Arbor) 6: 1-458.

Brindle, A. 1963. Taxonomic notes on the larvae of British Diptera. Entomologist (Lond.) 96:
237-243.

} Contini, C. 1965. I Dixidae della Sardegna, Nuovi reperti e descrizione di Palaeodixa frizzii

N. Gen. N. Sp. Mem. Soc. Entomol. Ital. 44: 95-108, 6 pls.
Disney, R. H. L. 1975. A key to the British dixidae. Freshw. Biol. Assoc. Sci. Publ. No. 31,

78 pp.

528 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Edwards, F. W. 1932. Family Culicidae. Jn Wytsman, P., ed., Genera Insectorum Fasc. 194,
pp. 8-15.

Frizzi, G., C. Contini, and M. Mameli. 1966. Ulteriori richerche citogenetiche sui Dixidae
della Sardegna. Atti Assoc. Genet. Ital. Pavia XI: 286-291.

Harbach, R. E. and K. L. Knight. 1980. Taxonomists’ Glossary of Mosquito Anatomy. Plexus
Publ. Co., Marlton, N.J. 413 pp.

Hennig, W. 1950. Die Larvenformen der Dipteren. Teil 2, pp. 66—67.

Johannsen, O. A. 1903. Aquatic nematocerous Diptera. N.Y. State Mus. Bull. 68: 429-432.

. 1933. Aquatic Diptera, Part I. Nemosera exclusive of Chironomidae and Ceratopo-

gonidae. Cornell Univ. Agric. Exp. Stn. Mem. 164: 35-37.

Martini, E. 1929. Subfamilie Dixidae. Pp. 20-43. Jn Lindner, E., Die Fliegen der Palaeark-
tischen region Lief 33, Culicidae.

Nowell, W. R. 1951. The dipterous family Dixidae in western North America. Microento-
mology 16: 187-270.

Peters, T. M. 198la. Family Dixidae. Jn McAlpine, J. F., B. V. Peterson, G. E. Shewell, H.

J. Teskey, J. R. Vockeroth, and D. M. Wood, Manual of Nearctic Diptera. Vol. I. Res.

Branch, Agric. Can., Monogr. No. 27. 674 pp.

. 1981b. A new Nearctic Dixa (Diptera: Dixidae). Proc. Entomol. Soc. Wash. 83: 516—

519.

Peters, T. M. and P. Barbosa. 1970. A new Nearctic species of Dixinae (Culicidae: Diptera).
Entomol. News 81: 11-17.

Peters, T. M. and E. F. Cook. 1966. The Nearctic Dixidae. Misc. Publ. Entomol. Soc. Am.
Se PBN 703,

Rohdendorf, B. 1964. The Historical Development of Diptera. Tr. Paleontol. Inst. Akad.
Nauk USSR, Vol. 100 (English translation, Univ. of Alberta Press, Edmonton. 360 pp.)

Smith, F. K. 1928. Larval characters of genus Dixa. J. N.Y. Entomol. Soc. 36: 263-286.

Tonnoir, A. L. 1921-1923. Australian Dixidae. Pap. Proc. R. Soc. Tasmania, pp. 58-71.

Waters, T. F. 1962. Diurnal periodicity in the drift of stream invertebrates. Ecology 43: 316—
320.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 529-534

NOMENCLATURAL NOTES ON POLYNEMA (HYMENOPTERA:
MYMARIDAE), WITH DESCRIPTION OF A NEW SPECIES!

M. E. SCHAUFF AND E. E. GRISSELL

(MES) Maryland Center for Systematic Entomology, Department of
Entomology, University of Maryland, College Park, Maryland 20742; (EEG)
Systematic Entomology Laboratory, IIBIII, Agricultural Research Service,
USDA, % National Museum of Natural History, Washington, D.C. 20560.

Abstract.—Polynema ema, new species, a parasite of the lily planthopper,
(Megamelus davisi Van Duzee), is described and illustrated. Psilus ciliatus
Say, previously reported as a Polynema and believed to be the parasite of
the lily planthopper, is a nomen dubium. Polynema ciliatum Perkins is con-
sidered a valid species and not a homonym of ciliatus Say.

In attempting to identify a species of Polynema parasitic upon eggs of the
lily planthopper (Megamelus davisi Van Duzee), we noticed that the name
Polynema ciliata (Say) had been used by Zimmerman (1948) for the parasite
of this planthopper. We found also that Perkins (1910) had described a
Polynema ciliata, thus creating an apparent homonym. After studying the
literature and available specimens, however, we believe that Say’s ciliatus
is not recognizable to genus, that Perkins’ ciliata is not a homonym, and
that the parasite of the lily planthopper is an undescribed species. We take
this opportunity to clarify the nomenclature of P. ciliata (Say) (of authors)
and to describe the incorrectly named parasite of the lily planthopper.

Morphological terminology used here is that of DeBauche (1948), except
for face height, which is the distance from the oral cavity to the median
carina. The abbreviation LMC is used for the length of the longest marginal
cilia of the forewing. Measurements of body length, ratios of body regions,
etc., were made dorsally along the midline. Wings, antennae, and legs were
measured in their extended form. Measurements were made with the aid of
both a compound (100 and 160) and a dissecting microscope (ca. 60).
Ranges and means were calculated from a random sample of 15 individuals
from the type-series.

! Scientific Article No. A-3024, Contribution No. 6087 of the Maryland Agricultural Exper-
iment Station, Department of Entomology, University of Maryland.

530 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Psilus ciliatus Say, nomen dubium

Psilus ciliatus Say 1828: 80.

Galesus ciliatus: Ashmead 1887: 195 (list).

Diapria ciliatus: Cresson 1887: 251 (list); Ashmead 1893: 428 (list, probably
a mymarid); Dalla Torre 1898: 436 (list); Kieffer 1916: 76 (repeat of orig.
desc. in German).

Trichopria ciliata: Kieffer 1911: 64 (list).

Polynema ciliatum: Peck 1951: 417 (n. comb.).

Psilus ciliatus Say has a long history of varied placement in two different
superfamilies of Hymenoptera (Proctotrupoidea and Chalcidoidea). Most of
the citations of this species are mere listings, and it is probable that none
of the authors except Say ever saw the type-specimen. This specimen is
lost, and based upon Say’s original description it is not possible to assign
ciliatus to a genus. For this reason we do not accept Peck’s placement
(1951) of ciliatus as a Polynema. Because there is no way positively to
associate Say’s description with a known genus or species, we relegate
ciliatus to the status of a nomen dubium. However, we agree with Ashmead
(1893) and Peck (1951) that ciliatus is probably a mymarid on the basis of
the enlarged antennal club and ciliate wing margins.

We do not know how or why the name ciliata (Say) of authors became
applied to the Polynema that attacks Megamelus davisi Van Duzee (Van
Duzee, 1896: 18; Zimmerman, 1948: 248). There is no reason to associate
this parasite with the name ciliata, because, with its extremely long ovi-
positor, it is quite distinct among all Polynema. On the basis of the ‘“‘oblong
oval acute club’’ as described by Say, his specimen would have been a
female, and he certainly would have mentioned the ovipositor if it were
prominent. Say mentioned no host, so that host-association is not possible.

Polynema ema Schauff and Grissell, NEw SpEcIES
Figs. 1-3

Cosmocoma ciliata (Say): Van Duzee, 1896: 18 (misidentification).

Polynema ciliata (Say): Zimmerman, 1948: 248 (misidentification).

Polynema ciliata (Say): Wilson and McPherson, 1981: 346 (misidentifica-
tion).

Holotype female.—Length 0.63 mm excluding ovipositor (0.60-0.66, x =
0.64). Ratio head:thorax:abdomen:ovipositor 8:19:20:30. Head, thorax (ex-
cept pronotum), and abdomen brown; funicle segments, club, last tarsal seg-
ment of legs lighter brown; scape, pedicel, prothorax, legs, petiole yellow.
Head slightly wider than thorax (12:10), vertex alutaceous; median, frontal
supraorbital carinae complete, occipital suture reaching foramen; posterior
ocelli placed at junction of supraorbital carina and occipital suture; POL:
OOL 7:1; frontal grooves converging below toruli, ending on either side of

VOLUME 84, NUMBER 3 531

_ oF
y
) 7.
=
Sar
-
x

mt a
| Ve
i!

Fig. 1. Polynema ema, female, habitus.

clypeus, interocular distance:face height 5:7; torulus removed ca. | diameter
from median carina, laterally against frontal carinae; antennal ratio (Fig. 2)
beginning with scape 33:19:8:21:15:12:11:12:36; club with 7 sensory ridges;
ratio pronotum:scutum:scutellum:propodeum 5:5:5:4. Pronotum aluta-

532 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ceous, becoming coriaceous laterally, divided medially, each side with 5
setae on anterior margin, 3 setae at posterior margin, posterior % of pro-
sternum divided medially, scutum alutaceous, notauli interrupted by large
fovea in anterior '/5; scutellum alutaceous (weaker than scutum), placoid
sensilla removed ca. 3 diameters from anterior margin, separated ca. 5 di-
ameters, transverse row of foveae in posterior '/s; propodeum smooth, de-
scending, median carina broken, ending in a tooth above petiolar insertion
(viewed laterally), with a seta posterolaterally above hindcoxa; forewing
(Fig. 1) length 0.82 mm (0.82-0.88, <¢ = 0.84), width 0.13 mm (0.12-0.14,
x = 0.13), LMC 0.20 mm (0.18—0.20, ¢ = 0.19); hindwing length 0.72 mm
(0.67-0.75, « = 0.71), width 0.02 mm (0.019-0.024, x = 0.20); petiole 2X as
long as wide, produced into a tooth ventrally, (Fig. 3); ratio femur: tibia: tarsi
as follows: foreleg 11:11:13, midleg 10:14:17, hindleg 12:16:22, coxa with
small group of setae on inner surface, hindbasitarsus 4x as long as tarso-
mere 2. Abdomen ovate elliptic, length 0.30 mm (0.30-0.41, x = 0.36), faint-
ly striate dorsally; ovipositor exserted 0.45 mm (0.35—0.50, x = 0.42).

Allotype male.—Length 0.67 mm (0.60-0.75, « = 0.66). Similar to female
except the following: antennal ratio beginning with scape
30:22:25:26:28:27:29:28:28:27:27:26:26; petiole not produced into a tooth
ventrally; abdomen ovate, length 0.22 mm (0.18-0.25, x = 0.22)

Types.—Holotype 2 on point (antenna and wings on slide) with data as
follows: ‘‘Ill., Jackson Co., 3 mi. N. of Pomona, Etherton Pond, 25 Aug.,
1979, S. W. Wilson, ex. eggs of Megamelus davisi on Nuphar advena.”
Deposited in the National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C., Type No. 100015. Allotype and paratypes (22
3 and 36 2) with same data as above. Paratypes deposited in the British
Museum (Natural History), London, Canadian National Collection, Ottawa,
and National Museum of Natural History.

Other specimens.—Two ¢ and 4 9 from Honolulu, Hawaii, reared from
the lily planthopper, E. C. Zimmerman collector (date unknown, probably
1945); 2 2, Md., Prince Georges Co., Laurel, Patuxent Wildlife Research
Center, 25 June, 1980, L. Masner, pan trap in pondside vegetation.

Variation.—Quantitative estimates of variation are given in the species
description. Color varied between series of specimens and between speci-

mens of the same series. The specimens from Hawaii are nearly uniformly |

dark brown, with the legs, pronotum, scape, and pedicel light brown. Most

specimens in the type-series are similar to the holotype. However, a few |

females are lighter in color with the yellow faded to very light yellow (coxae
occasionally nearly white), and males may have the abdomen very dark
brown or black mesad. The occipital suture does not reach the foramen in
some specimens, ending only slightly past the lateral ocellus. The structure
of the median propodeal carina is very difficult to see in this species both

VOLUME 84, NUMBER 3 533

Fig. 2,3. Polynema ema, female. 2, Antenna, lateral view. 3, Petiole, lateral view.

in slide-mounted and pointed specimens. Scanning electron micrographs
show the area to be slightly sunken and composed of irregular broken carina
that forms a keel posteriorly. When viewed laterally under the compound
microscope, this area appears as a raised ‘‘tooth’’ above the petiolar inser-
tion.

Diagnosis.—Females of this species are distinct from other Polynema by
virtue of the long slender forewing (6 times as long as broad), the ovipositor
exserted the length of the abdomen or more, and the presence of the ventral
tooth on the abdominal petiole. Female Polynema normally have the fore-

- wing only 3 or 4 times as long as broad, the abdominal petiole without a
ventral tooth, and the ovipositor not exserted. Males of Polynema (as with

most mymarids) are more difficult to separate; however, the narrow fore-
wing may be used to distinguish this species.

Remarks.—Polynema ema probably will be found to occur throughout
the range of its host, Megamelus davisi. In the continental United States,
M. davisi is known throughout the eastern states and west as far as Kansas
(Metcalf, 1943; Beamer, 1955). Van Duzee (1896: 18) was the first to asso-
ciate P. ema with Megamelus in Michigan. Wilson and McPherson (1981:
346) recently made brief mention of this parasite in relation to its host in
Illinois. According to Zimmerman (1948: 248), Polynema ema [reported as
ciliata (Say)] was ‘‘brought to Honolulu from Michigan in 1941 by Fullaway
and it quickly established itself on local leafhopper colonies’ of Megamelus
davisi (reported as angulatus Osborn, a synonym of davisi).

Etymology.—The species epithet is an euphonius combination of arbi-

| trary letters.

.

Polynema ciliatum Perkins
Polynema ciliata Perkins, 1910: 666.

As previously mentioned, we do not accept the placement of ciliatus Say
as a Polynema. Therefore, Perkins’ ciliatum is not a homonym, and having

534 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

seen the type, we consider it a valid species. It is known only from Oahu,
Hawaiian Islands.

ACKNOWLEDGMENTS

We thank S. W. Wilson (California State University, Chico) for providing
specimens of the new species, J. P. Kramer (Systematic Entomology Lab-
oratory, IIBII, USDA, Washington, D.C.) for providing information about
its host, G. M. Nishida (B. P. Bishop Museum, Honolulu, Hawaii) for loan-
ing us the type-specimen of Polynema ciliata Perkins, Linda Heath Law-
rence for the habitus drawing, and Phyllis W. Iglehart for typing the manu-
script.

LITERATURE CITED

Ashmead, W. H. 1887. Studies of the North American Proctotupidae with descriptions of new
species from Florida. Can. Entomol. 19: 192-198.

—. 1893. A monograph of North American Proctotrypidae. Bull. U.S. Natl. Mus. 45: I-
472.

Beamer, R. H. 1955. A revision of the genus Megamelus in America North of Mexico. J.
Kans. Entomol. Soc. 28: 29-46.

Cresson, E. T. 1887. Synopsis of the families and genera of the Hymenoptera of America
North of Mexico together with a catalogue of the described species and bibliography.
Trans. Am. Entomol. Soc. Suppl. 1887: 155-350.

Dalla Torre, C. G. 1898. Catalogus Hymenopterorum hucusque descriptorum systematicus et
synonymicus 5. Chalcididae et Proctotrupidae, Leipzig. 598 pp.

DeBauche, H. R. 1948. Etude sur les Mymarommidae et les Mymaridae de la Belgique (Hy-
menoptera:Chalcidoidea). Mem. Mus. Hist. Nat. Belg. 108: 1-248.

Kieffer, J. J. 1911. Hymenoptera Fam. Diapriidae. Jn Wytsman, P., ed., Genera Insectorum

124. 75 pp.

. 1916. Diapriidae. Das Tierreiche 44, 627 pp.

Metcalf, Z. P. 1943. General catalog of the Hemiptera. Fascicle IV. Fulgoroidea Pt. 3. Araeo-
pidae (Delphacidae). Smith College, Mass. 552 pp.

Peck, O. 1951. Chalcidoidea, pp. 410-594. In Muesebeck, C. F. W., et al., eds., Hymenoptera |
of America North of Mexico, Synoptic Catalog. U.S. Dep. Agric. Agric. Monogr. 2.

Perkins, R. C. L. 1910. Supplement (by R. C. L. Perkins) to Hymenoptera. Fauna Hawaiiensis |
Hym. Suppl. 2: 600-686.

Say. T. 1828. A description of some new species of Hymenoptera of the United States. Contrib.
Maclurian Lyceum Phil. 1: 67-83.

Van Duzee, E. P. 1896. Pests of House and Ornamental Plants. Mich. Agric. Exp. Stn., Spec.
Bull. No. 2, 43 pp.

Wilson, S. W. and J. E. McPherson. 1981. Life history of Megamelus davisi with descriptions
of immature stages. Ann. Entomol. Soc. Am. 74: 345-350.

Zimmerman, E. C. 1948. Insects of Hawaii. Vol. 4. University of Hawaii Press, Honolulu.
268 pp.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 535-538

CHROMOSOME STUDY IN MALES OF NEARCTIC SPECIES OF
GERRIS FABRICIUS AND LIMNOPORUS STAL
(HEMIPTERA: HETEROPTERA: GERRIDAE)

DIANE M. CALABRESE AND PETER TALLERICO

(DMC) Department of Biology and The Wildlife Sanctuary, Dickinson
College, Carlisle, Pennsylvania 17013; (PT) Nursing Department, Pennsyl-
vania State University, University Park, Pennsylvania 16802.

Abstract.—Chromosome study of males of some Nearctic species of Ger-
ridae of the genera Gerris and Limnoporus reveals that contrary to what
has been reported for Palearctic species, the mechanism of sex determina-
tion varies among subgroups. Consistently absent are m chromosomes, and
kinetochore activity is restricted to the terminal ends of chromosomes dur-
ing meiosis.

Although diploid and/or haploid numbers have been reported for 15
species of Gerridae (Hemiptera: Heteroptera) (see summary, Ueshima,
1979), results for only one Nearctic species have been reported. Montgom-
ery (1901) reported a haploid number of 10A + XO for Gerris marginatus
Say (as Limnotrechus marginatus). Unfortunately, in 1901 at least three
species were included under the binomen G. marginatus and the nomen-
clature was not clarified until 1934 by Drake and Harris.

We have undertaken a study of chromosome number and structure in the
Gerridae because we believe, as do Ueshima and Ashlock (1980), that chro-
mosome morphology may be useful in the phylogenetic reconstruction of
the families of aquatic and semiaquatic Heteroptera. We also anticipate that
chromosomal rearrangements may be correlated with change in wing form
within populations of wing polymorphic species.

We have looked at representatives of Nearctic Gerris (Gerris), G. (Aquar-
ius), and Limnoporus (until 1975, Andersen, G. (Limnoporus)) and we re-
port here our results for these groups. We compare the results to those that
have been reported for Palearctic species of the same genera and subgenera.

MATERIALS AND METHODS

Gerris (A.) remigis (Say) was collected at Aguirre Springs, N.M., G. (G.)
buenoi Kirkaldy and L. dissortis Drake and Harris were collected near

536 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Wy 3 | 5

Figs. 1-5. Metaphase I, photographed at ca. 6400. 1, Gerris (Aquarius) remigis, early |
metaphase I. 2, G. (Gerris) alacris. 3, G. (G.) comatus. 4, G. (G.) buenoi. 5, Limnoporus
dissortis.

VOLUME 84, NUMBER 3 537

Minneapolis, Minn., G. (G.) alacris Hussey and G. (G.) comatus Drake
and Hottes were collected at Presque Isle State Park, Pa., and G. (G.)
marginatus and G. (G.) insperatus Drake and Hottes were collected in
Washington, D.C. All species were lab-reared under a 14L:10D photoperiod
using techniques that have been described elsewhere (Calabrese, 1974,
1979). Spermatogenic (meiotic) tissue was obtained from males approxi-
mately 24 hours after they had molted to the fifth instar. Such males, we
discovered, showed the greatest number of metaphase I figures.

Testes were fixed (15 ml glacial acetic acid, 45 ml absolute ethanol, 5 ml
acetone) for 2-3 minutes. Each testis was divided and each follicle placed
on a slide with a drop of lacto-proprio orcein stain for at least 3—5 minutes.
A standard squash was made. Slides were sealed in temporary mounts and
scanned for figures at 40x. Numerous preparations were made and scanned

for each species. Metaphase I figures were photographed at about 6400
-under oil on a Zeiss compound microscope with a Wetzler large format
camera attachment.

RESULTS

_ All results were consistent for the minimum of five representatives ex-
amined per species.

— Gerris (Aquarius) remigis (Fig. 1) exhibits a diploid number of 22, and
the male is XO.

Four of the Gerris species examined exhibit an XY sex determination
mechanism in the male and a diploid number of 20 (Figs. 2, 3; G. insperatus
and G. marginatus not shown). However, G. (G.) buenoi, although it ex-
hibits a diploid number of 20, shows an XO sex determination mechanism
(Fig. 4).

Limnoporus dissortis exhibits a diploid number of 22 and what appears
to be a XO, sex determination mechanism in the male (Fig. 5).

No m chromosomes are visible in any of the species studied (Figs. I-5;
G. insperatus and G. marginatus not shown). Kinetochores are restricted
to the ends of meiotic chromosomes in all species studied (examples, Figs.
1-5).

DISCUSSION

Ueshima (1979) suggested that the Gerridae do not have m chromosomes
and that kinetochores are found only at the terminals of chromosomes dur-
ing meiosis. The results of our studies to date support those suggestions.
However, we do not agree with Ueshima (1979) that the XO system is the
universal mode of sex determination among the Gerridae. In four of the
Gerris (Gerris) species we studied the male is heterogametic, but it shows
an XY system. In one G. (Gerris) species (G. buenoi) and in G. (Aquarius)

=_——_—

538 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

remigis and Limnoporus dissortis the mode of sex determination appears
to be XO, male heterogametic. It is notable that in a reanalysis of the cy-
togenetics of G. (Aquarius) paludum, a Palearctic species, (Takenouchi and
Muramoto, 1968) it has been shown that the sex determination mechanism
is XY and not XO as had been reported earlier (Wilke, 1913).

The results of the study to date suggest that there is no consistency within
genera and subgenera in terms of sex determination mechanism. Differences
in chromosome morphology and the cytogenetic characters discussed herein
may prove to be useful characters in a further test of the existing phyloge-
netic reconstruction for the Gerridae (Calabrese, 1980), when more species
are studied (compare Figs. 1-5).

ACKNOWLEDGMENTS

We are very grateful to Donald Bennett, University of Connecticut, who
took the time to collect gerrid specimens for us in Minnesota and the care
to send them to us as live specimens.

LITERATURE CITED

Andersen, N. M. 1975. The Limnogonus and Neogerris of the Old World with character
analysis and a reclassification of the Gerridae (Hemiptera: Gerridae). Entomol. Scand.
Suppl. 7, 96 pp.

Calabrese, D. M. 1974. Keys to the adults and nymphs of the species of Gerris Fabricius
occurring in Connecticut. Mem. Conn. Entomol. Soc. 1974, pp. 227-266.

———. 1979. Pterygomorphism in ten Nearctic species of Gerris F. Am. Midl. Nat. 101: 61-

68.

. 1980. The zoogeography and a cladistic analysis of the Gerridae. Misc. Publ. Entomol.

Soc. Am. 11(5): 1-119.

Drake, C. J. and H. M. Harris. 1934. The Gerrinae of the Western Hemisphere (Hemiptera).
Ann. Carnegie Mus. 23: 179-240.

Montgomery, T. H. 1901. Further studies on the chromosomes of the Hemiptera-Heteroptera.
Proc. Acad. Nat. Sci. Phila. 53: 261-271.

Takenouchi, Y. and N. Muramoto. 1968. A survey of the chromosomes in twenty species of
Heteroptera insects. (in Japanese). J. Hokkaido Univ. Educ. Sect. II B 19: 1-19.
Ueshima, N. 1979. Animal Cytogenetics. vol. 3 Insecta 6. Hemiptera II, Heteroptera. Gebru-

der Borntraeger, Stuttgart. 117 pp.

Ueshima, N. and P. D. Ashlock. 1980. Cytotaxonomy of the Lygaeidae (Hemiptera-Heter-
optera). Univ. Kans. Sci. Bull. 51: 717-801.

Wilke, G. 1913. Chromatinreifung und Mitochondrienk6rper in der Spermatogenese von Hy-
drometra paludum Fab. Arch. Zellf. 10: 203-236.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 539-546

A NEW SPECIES OF BITING MIDGE OF THE GENUS
ALLUAUDOMYIA KIEFFER (DIPTERA: CERATOPOGONIDAE)
FROM THE SOUTHEASTERN UNITED STATES!

JAYSON I. GLICK AND GARY R. MULLEN

Department of Zoology-Entomology and Alabama Agricultural Experi-
ment Station, Auburn University, Auburn University, Alabama 36849;
Graduate Research Assistant and Associate Professor of Entomology, re-
spectively; (JIG) presently: Captain, Medical Service Corps, U.S. Army,
Department of Arboviral Entomology, U.S. Army Medical Research Insti-
tute of Infectious Diseases, Ft. Detrick, Frederick, Maryland 21701.

Abstract.—Alluaudomyia variegata, a new species of biting midge from
the southeastern United States, is described and illustrated, and the known
biology is discussed. This species is believed to be a member of the Mac-
ulipennis Group and is most closely related to the widespread Nearctic
species A. needhami Thomsen.

The worldwide genus Alluaudomyia Kieffer is comprised of relatively
small, often strikingly marked biting midges. Only eight species have thus
‘far been reported for the Nearctic Region (Wirth, 1952a; Williams, 1956).
During a study of the Ceratopogonidae of Alabama, males and females of
a previously undescribed species of Alluaudomyia were taken in light traps
at localities in central and northern Alabama. Additional specimens obtained
from the National Museum of Natural History, Smithsonian Institution,
Washington, D.C., extend the known distribution of this species to encom-
pass the Gulf Coast states of the United States.

For an explanation of the general morphology and terminology of the
ceratopogonids see Wirth (1952b). The descriptions are based on specimens
slide-mounted in balsam or Hoyer’s medium; measurements were obtained
with the aid of a calibrated ocular micrometer in the eyepiece of a Zeiss
phase-contrast microscope. The wing length is measured from the basal

.

: 1 This research was supported by Alabama Hatch Project AL-431. Part of a dissertation by
the senior author in partial fulfillment of the Ph.D. requirements of the Graduate School of
Auburn University.

540 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

arculus to the wing tip; costal length (costal ratio) is obtained by dividing
the length of the costa by the wing length. Antennal ratio is determined by
the combined lengths of the 5 distal flagellomeres divided by the combined
lengths of the preceding 8 flagellomeres. The hindtarsal ratio is obtained by
dividing the length of the hindbasitarsus by the length of the second tarso-
mere. Proportions given for antennal flagellomeres and for palpal segments
refer to the relative lengths as units in an ocular micrometer and not as
absolute measurements. Variation is given by the mean, followed by the
minimum-maximum values and sample size.

All types and other material examined are slide-mounted except where
noted. Types are deposited in the National Museum of Natural History,
Smithsonian Institution, Washington, D.C. (USNM); paratypes will be de-
posited in the British Museum (Natural History), London (BM); California
Academy of Sciences, San Francisco (CAS); Florida State Collection of
Arthropods, Gainesville (FSCA); and the Auburn University Entomological
Museum (AU). All other material examined is deposited in the National
Museum of Natural History, Washington, D.C.

Alluaudomyia variegata Glick and Mullen, NEw SPEcIEs
Figs. 1, 2

Alluaudomyia needhami Thomsen; Snow et al., 1957: 21 (in part, misiden-
tification; reared ¢).

Diagnosis.—A small brownish species; wing markings extensive with dark
streaks distally on the veins, membrane between the veins with a varying
amount of infuscation. Female with | large, ovoid spermatheca. Male 9th
tergum with long, slender apicolateral processes; aedeagus with stout, blunt
tip; distal portion of paramere laterally recurved terminating in a ventrally
bent, pointed apex.

Female.—Wing length 1.26 (0.98—1.52) mm (n = 15).

Head: Brown; occiput white with a midbasal brownish triangulate spot.
Eyes medially contiguous for a short distance, bare. Antenna (Fig. la) with
flagellomeres in proportion of 9-6-6-7-7-8-8-8-10-10-12-11-14; antennal ratio
0.94 (0.87-1.01, n = 14); proximal flagellomeres relatively short and some-
what tapering distally; distal flagellomeres slender; apices of flagellomeres
3-10 pale. Palpal segments (Fig. Ic) short, in proportion of 5-7-8-5-8; seg-
ment 3 with a small, round, distal sensory pit with several elongate hyaline
sensilla; Ist, 2nd, and base of 3rd segments pale. Mandible (Fig. 1j) with 18
(17-19, n = 10) teeth on inner margin, outer margin with 3—5 widely spaced
minute teeth.

Thorax: Brown; mesonotum with numerous, small, dark brown spots;
humeri and lateral margins of mesonotum whitish; prescutellum with a me-
dian pair of whitish spots and 2 long median setae; scutellum white with a

VOLUME 84, NUMBER 3 541

h

Fig. 1. Alluaudomyia variegata. a, c, e-h, j, Female. b, d, i, Male. a, b, Antenna. c, d,
Palpus. e, Genital sclerotization. f, Fifth tarsomeres and claws of (top to bottom) fore-, mid-,
and hindlegs. g, Femora and tibiae of (left to right) fore-, mid-, and hindlegs. h, Spermatheca.
i, Genitalia. j, Mandible.

542 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

thin median brown band and with 6 long setae; postscutellum and pleuron
brown.

Legs (Fig. |g): Brown, femora pale basally, with subapical pale rings;
tibiae with subbasal and subapical pale rings, broader on hindtibia; tarsi
pale, darker at apices, hindbasitarsus brownish; hindtibial comb with 6—7
spines; hindtarsal ratio 2.9 (2.7—3.3, n = 14). Tarsal claw (Fig. If) long and
slender, unequal on all legs.

Wing (Figs. 2a, b): With 6 conspicuous dark spots as follows: 1 large
irregular marking proximal to r-m crossvein; a smaller spot just caudad of
latter spot; | large spot just below apex of radial cell; | large elongate spot
near base of vein M2; a moderately small spot on anterior wing margin just
distad of junction of costa and R1; and a small spot at tip of anal vein. Wing
with prominent dark streaks over apices of veins RI and Rs, midportion
and apex of M1, apices of M2 and M3+4, middle of cubitus, and over medio-
cubital fork to apex of Cul; small amount of infuscation just distad of basal
arculus between bases of radius and cubitus, and on anterior wing margin
in costal cell; a varying amount of dark infuscation on the membrane be-
tween the veins in distal portion of cells R5 and M2, and extensively in M1,
M4, and anal cells (often weakly infuscated in larger individuals, Fig. 2b);
costa extending to 0.73 (0.68-0.77, n = 15) of wing length. Halter stem and
base of knob whitish, dark apically.

Abdomen: Brown; apices of terga with white bands, 7th and 8th terga
almost entirely whitish. Genital sclerotization (Fig. le) as figured. One large
ovoid spermatheca (Fig. Ih) 0.116 x 0.075 mm, with short, tapering, scler-
otized neck.

Male.—Similar to female with the usual sexual differences. Antenna (Fig.
lb) with apex of flagellomere 12 and all of 13-15 brownish; scutellum with
4 long setae; dark streaks on wing veins more prominent and extensive than
in female, with spot absent on anterior margin of wing distad of junction of
costa and RI. Genitalia (Fig. li) with sternum 9 relatively short, with a
broad, deep, caudomedian emargination, caudal membrane spiculate; ter-
gum 9 elongate with moderately tapering sides and long, slender apicolateral
processes. Basistyle moderately long, dististyle nearly straight with stout,
blunt apex. Aedeagus heavily sclerotized, with a deep basal arch; basal arms
slender and nearly straight, with a small flange at base; distomedian process _ |
short and stout with blunt, ventrally bent tip. Paramere with short basal
arm and a short, anterolaterally directed basal lobe; stem moderately slender
with a slight bend mesally, expanded distally and laterally recurved into a
short, slender appendage terminating in a ventrally bent, pointed apex.

Distribution.—Southeastern United States, from extreme eastern Texas
to Florida, and north to Tennessee.

Types.—Holotype ¢, allotype 2, Alabama: Lee Co., Chewacla State

VOLUME 84, NUMBER 3 543

| Fig. 2. Alluaudomyia variegata. a, b, Female wing, showing range of infuscation on the
_ membrane between the veins.
|

Park, near Chewacla Pond, J. I. Glick, light trap, 10-11 June 1977 (USNM
| type no. 76104). Paratypes, 35 6, 25 2, as follows. ALABAMA: Dallas

Co., Marion Junction, Alabama Agricultural Experiment Station (Black Belt

Substation), J. I. Glick, light trap, 21-23 May 1978, | ¢ (AU); same data,
126-27 May 1978, 1 2 (CAS); same data, 31 May-—2 June 1978, | 2 (AU);
' same data, 9-12 June 1978, 1 9 (AU); same data, 22-24 June 1978, 1 3
(USNM); same locality, M. Lame, reared, 2 May 1979, | 6 (AU); Lauder-
dale Co., Florence, Collier Slough, near Tennessee River, J. I. Glick, light
trap, 21-24 July 1978, | 6 (AU); Lee Co., Chewacla State Park, near Che-
wacla Pond, J. I. Glick, light trap, 18-19 April 1977, 2 2 (AU); same data,
11-17 May 1977, 1 2 (AU); same data, 2-8 June 1977,4 6,5 2 (46,4 9
in alcohol) (AU); same data, 9 June 1977, 2 6, 2 2 (FSCA); same data, 14
June 1977, 1 2 (AU); same data, 22 June 1977, 2 ¢ (AU); same data, 29
June 1977, 1 6 (AU); same data, 4 Aug 1977, | ¢ (AU); same data, 25 Aug
1977, 1 6 (AU), 1 2 (USNM); same data, 18-23 July 1978, | 2 (USNM);

:

544 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

same data, 21 Aug 1978, 1 ¢, 1 2 (BM); same data, 26 Sept—1 Oct 1978, 4
3 (AU); same data, 10-15 Oct 1978, 1 ¢ (CAS); same data, 24-29 Oct 1978,
93,2 2 (AU), 2 d (USNM); same data, 30-31 Oct 1978, 1 ¢ (AU); same
data, 1-6 Nov 1978, | 6 (AU); Loachapoka, Loachapoka Hunt Club, J. I.
Glick and B. Buxton, light trap, 27-29 May 1977, | 2 (AU); Morgan Co.,
Decatur, Wheeler National Wildlife Refuge, J. I. Glick, light trap, 8-10 May
1978, | 2 (USNM); same data, 11-14 May 1978, 1 2 (AU); same data, 19-
21 May 1978, 1 2 (AU); same data, 22-25 May 1978, 1 6, 1 2 (USNM);
same data, 26-29 May 1978, 1 2 (AU).

Other specimens examined.—42 ¢, 6 2 (USNM) as follows. FLORIDA:
Alachua Co., Gainesville, W. W. Smith, light trap, 10 April 1965, 1 d; same
locality, W. W. Wirth, light trap, 20 April 1967, | ¢; same locality, F. S.
Blanton, light trap, 8 May 1967, | ¢; Hawthorne, F. S. Blanton, light trap,
27 April 1968, 1 2; Baker Co., Glen St. Mary, F. S. Blanton, light trap,
May 1971, | 9; Gulf Co., Wewahitchka, State Board of Health, light trap,
May 1961, | 6; same data, July 1961, | d; same data, Aug 1961, 1 d; same
data, Oct 1961, | 6; Holmes Co., Bonifant, State Board of Health, light
trap, June 1961, | 6; same data, Sept 1961, | 2; same data, Oct 1961, 1
3; Jackson Co., Florida Caverns State Park, W. W. Wirth, light trap, 26
May 1973, | 2; Sneads, State Board of Health, light trap, 6 April 1954, 1
6; same data, March 1955, 1 6; same data, 31 May 1955, 1 6; Jefferson
Co., Monticello, W. H. Whitcomb, light trap, May 1969, | 6, 1 9; Leon
Co., 3 mi N. of Tallahassee, F. S. Blanton, light trap, May 1970, 1 9;
Liberty Co., Torreya State Park, F. S. Blanton, light trap, 27 April 1958,
1 6; same data, May 1971, | ¢6; same locality, H. V. Weems Jr., light trap,
20 May 1966, 2 6; same locality, W. W. Wirth, light trap, 22 April 1967, |
6. LOUISIANA: St. Tammany Parrish, Pearl River, K. T. Khalaf, light
trap, April 1966, 16 36; Slidell, K. T. Khalaf, light trap, July 1966, 1 ¢.
MISSISSIPPI: Hancock Co., Gainesville, K. T. Khalaf, light trap, April
1966, | ¢; Harrison Co., K. T. Khalaf, light trap, May 1966, 4 6. TEN-
NESSEE: Polk Co., Copperhill, Ocoee Reservoir, E. Pickard and W. Snow,
reared from pupa, 26 Aug 1954, 1 6. TEXAS: Jasper Co., Jasper, R. E.
Woodruff, light trap, 2 Aug 1968, 1 ¢.

Biology.—Snow et al. (1957) reported rearing a single pupa of A. varie-
gata to the adult (misidentified by Wirth as Alluaudomyia needhami Thom-
sen). The pupa was collected in late August from a small pool on an island
in Ocoee Reservoir, Copperhill, Tennessee; the pool was overgrown with
prickly stem smartweed (Polygonum sagittatum Linnaeus) and black willow
(Salix nigra Marsh). During the current study, an adult male was reared in
early May from a substrate sample of wet mud and decomposing leaves
taken from the margin of a small pasture pond at the Alabama Agricultural
Experiment Station (Black Belt Substation), Marion Junction, Alabama.
Other Nearctic species of Alluaudomyia including A. bella (Coquillett), A.

VOLUME 84, NUMBER 3 545

needhami, and A. paraspina Wirth share a similar larval habitat (Thomsen,
1937; Williams, 1953; Grogan and Messersmith, 1976).

Adults of A. variegata have been collected in light traps from March
through November, with the greatest numbers being taken during April and
May. In east-central Alabama two activity peaks were noted, the first from
May through June and a second in October.

Discussion.—Wirth and Delfinado (1964) proposed five ‘“‘natural’’ groups
for the genus Alluaudomyia in which they placed 79 of the 99 then known
species. We are placing A. variegata in the large and diverse Maculipennis
Group, the adults of which are characterized by the following combination
of group characters: (1) wing with at least two conspicuous spots, one
proximal to the r-m crossvein and one at the apex of the costa; (2) the distal
portions of the wing veins often with dark streaks, and often with prominent
spots present in the cells between the veins near the wing margin; (3) legs
usually dark with narrow pale rings; (4) antennae dark with narrow basal
pale rings on flagellomeres 3—10; (5) tarsal claws usually unequal on all legs;
and (6) one spermatheca present, without diverticulum. The Nearctic
species A. variegata, A. megaparamera Williams, and A. needhami are
members of the Maculipennis Group which lack distal spots in the wing
cells. Alluaudomyia variegata most closely resembles A. needhami, but can
be easily separated by its distinctive wing markings with infuscation on the
membrane between the veins, differences in the shape of the female genital
sclerotization, and by the unusual shape of the male parameres. Alluaudo-
myia megaparamera is a smaller species with more reduced wing markings,
a yellowish thorax and abdomen, and extensive pale markings on the legs.

ACKNOWLEDGMENTS

The authors thank Willis W. Wirth of the Systematic Entomology Lab-
oratory, USDA, Washington, D.C., for providing additional specimens for
study from the collections of the National Museum of Natural History, and
for critically reviewing the manuscript, and Eugene Pickard of the Tennes-
see Valley Authority, Water Quality and Ecology Branch, Muscle Shoals,
Alabama, for his kind donation of a reared specimen. We also thank Lt.
Colonel Harold W. Lupton and Major Charles L. Bailey, U.S. Army Med-
ical Research Institute of Infectious Diseases, Ft. Detrick, Frederick, Mary-
land, and Captain Michael E. Faran, Dept. of Entomology, Walter Reed
Army Institute of Research, Washington, D.C., for their helpful suggestions
and review of the manuscript.

LITERATURE CITED

Grogan, W. L., Jr. and D. H. Messersmith. 1976. The immature stages of Alluaudomyia
paraspina (Diptera: Ceratopogonidae) Wirth with notes on its biology. Ann. Entomol.
Soc. Am. 69: 687-690.

546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Snow, W. E., E. Pickard, and J. B. Moore. 1957. The Heleidae of the Tennessee River Basin.
J. Tenn. Acad. Sci. 32: 18-36.

Thomsen, L. C. 1937. Aquatic Diptera. Part V. Ceratopogonidae. Cornell Univ. Agric. Exp.
Stn. Mem. 210: 57-80.

Williams, R. W. 1953. Notes on the bionomics of the Alluaudomyia of Baker County, Georgia.
I. Observations and breeding habits of bella and needhami (Diptera, Heleidae). Proc.
Entomol. Soc. Wash. 55: 283-285.

——. 1956. Two new species of Alluaudomyia from Cheboygan County, Michigan, with a
note on the synonomy of parva and downesi (Diptera, Heleidae). Proc. Entomol. Soc.
Wash. 58: 327-331.

Wirth, W. W. 1952a. The genus Alluaudomyia Kieffer in North America (Diptera, Heleidae).

Ann. Entomol. Soc. Am. 45: 423-434.

. 1952b. The Heleidae of California. Univ. Calif. Publ. Entomol. 9: 95-266.

Wirth, W. W. and M. D. Delfinado. 1964. Revision of the Oriental species of Alluaudomyia
Kieffer (Diptera, Ceratopogonidae). Pac. Insects 6: 599-648.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 547-567

LYCHNUCHOIDES FRAPPENDA FROM CENTRAL MEXICO JOINS
LUNUS AND ZWEIFELI IN A LUNUS GROUP OF ATRYTONOPSIS
(LEPIDOPTERA: HESPERITDAE: HESPERIINAE)

JOHN M. BURNS

Department of Entomology, National Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.

Abstract.—Lychnuchoides frappenda, described by Dyar from two fe-
males in 1920, is actually a species of Atrytonopsis (new combination) un-
usual in having a broad semihyaline band that boldly disrupts the dark pri-
mary from midcosta to tornus. Despite its distinctive facies, frappenda

clusters tightly with A. Junus and A. zweifeli in what appears, from limited

locality data, to be a superspecies occurring discontinuously at middle el-
evations in woodland and scrub from southeastern Arizona to central Mex-
ico. This ‘‘/unus group’’ is defined on the basis of size (it includes the largest
species of Atrytonopsis), antenna (the apiculus is long and the nudum av-
erages 13 segments), male stigma (absent!), facies (primary always with a

strong cell spot and with the spot in space 1b under the spot in space 2;

secondary rarely with any dorsal spots), male and female genitalia (figured),
temporal distribution (the flight period of Junus, mostly from mid-July to
mid-August, is unique among species of Afrytonopsis in the United States),
and spatial distribution (mapped). Facies provides the best characters for
distinguishing species within the /unus group. This study was carried out in
the context of a review of the entire genus Afryfonopsis, with special atten-
tion to male and female genitalia and to variation in characters generally.

Dyar (1920: 187) so badly placed his new species frappenda, in Lychnu-
choides Godman (1901: 612), that its proper position, in Atrytonopsis God-
man (1900: 497) (NEW ComBINATION), has escaped notice for sixty years. Dyar

| missexed the type, though not the paratype, of frappenda—both are female.

He misspelled the name of the genus to which he banished frappenda—as
Lychnucoides rather than Lychnuchoides (a misspelling that some later
workers have repeated).

Dyar’s original description dealt only with expanse and appearance of the

| wings. It included no illustrations. Had Dyar looked critically and compar-

atively at facies or, better, beyond facies to some of the morphology God-

548 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

man used in defining Lychnuchoides, he could never have put frappenda
there. The excuse for doing so is crude similarity: a broad semihyaline band
cuts obliquely across the middle of a brown primary from midcosta almost
to the tornus both in true species of Lychnuchoides—saptine (Godman and
Salvin) and ozias (Hewitson)—and in frappenda (cf. Figs. 1-5 for this and
other aspects of facies discussed below). This conspicuous band, which is
rich yellow in Lychnuchoides, was described by Dyar as “‘light yellow”’ in
frappenda but is better dubbed creamy white. The spots that form it are not
shaped in frappenda as they are in true species of Lychnuchoides. More-
over, in frappenda, a spot in space 3 of the primary does not quite fill the
tip of the angle between the bases of veins 3 and 4 to become an integral
part of the band as it does in Lychnuchoides. Still other features of facies
fail to jibe: subapical spots occur in spaces 6, 7, and 8 of the primary in
frappenda, and in all other species of Atryfonopsis, but not in Lychnu-
choides:; complex ventral wing patterns in shades of gray and rust in Lychnu-
choides have no counterpart in frappenda, whose ventral secondary
closely resembles that of the other species of Atrytonopsis that usually lack
spots on the dorsal secondary, except for Joammi (Whitney). Most impor-
tant, frappenda fits smoothly into Atrytonopsis, but grossly distorts Lychnu-
choides, on many major morphological counts involving wing shape and
venation, antennae, palpi, vestiture and spination of legs, and genitalia.
Many of these differences were set down by Godman when he originally
characterized Atrytonopsis and Lychnuchoides two decades before Dyar
described frappenda.

With spots in spaces Ib, 2, and the cell enlarged and aligned to form a
conspicuous band bisecting the primary, frappenda at first looks bizarre in
Atrytonopsis. Semideparture from the Afrytonopsis image no doubt con-
spired with Dyar’s misclassification so that, over the years, neither the
segregated type of frappenda nor the paratype filed with true species of |
Lychnuchoides (both in USNM) registered correctly with skippermen who
saw them.

Hoffmann (1941: 282) listed Lychnuchoides frappenda for Mexico, and
Evans (1955: 253) included frappenda in his key to species of Lychnu-
choides (misspelled Lychnucoides but corrected in separate ‘‘Addenda and
Corrigenda’’); Evans never saw an example of it. Most recently, Okano
(1981: 31) retained Lychnuchoides (misspelled Lychnucoides) in his list of |
hesperiid genera occurring in Mexico. This he did strictly on account of |
frappenda, for L. saptine (long known from Guatemala) has yet to be re-
corded from Mexico. At least for the time being, Lychnuchoides should be
dropped from the Mexican list.

A third specimen of frappenda has surfaced, again a female. She was sent
to me because, in her unmounted state and with her pale band dividing her

VOLUME 84, NUMBER 3 549

Table 1. Length (mm) of right primary in Atrytonopsis lunus from southeastern Arizona.

Mean with Standard Coefficient
Sex N Range Standard Error Deviation of Variation
fo) 13 19.8-22.1 20.95 + 0.22 0.78 S)57/72
Q 44 19.6-23.6 21.80 + 0.13 0.87 3.99

brown primary, she suggested some sort of Autochton. When I took a hard
look at her, she said, *‘I belong to Atrytonopsis,’’ which provoked a review of
that genus and the writing of this paper.

At the end of his original description of A. zweifeli Freeman (1969: 277)
said, ‘‘The nearest related species to A. zweifeli is A. lunus (Edwards).”’
Actually, lunus, frappenda, and zweifeli all cluster tightly. It is sometimes
difficult to characterize this group: although /unus is represented by ade-
quate series of both sexes, frappenda is known only from three females and
zweifeli, from two males.

In what follows, I consider one sort of character at a time, first with
respect to its use in defining the /unus group within genus Afrytonopsis and

then, when pertinent, with respect to its further use in defining species
within the Junus group. Throughout I give special attention to problems
involving variation in characters, not only within the species of the lunus
group but also in the genus as a whole.

THE LUNUS GROUP

| Size.—The largest species of Afryftonopsis are in this group. Certainly
lunus itself and probably frappenda are markedly larger than most other
species of Atrytonopsis; however, the largest individuals of deva (Edwards)
and of ovinia (Hewitson) enter the size range of /unus and frappenda.

Using a vernier caliper calibrated to tenths of a millimeter, I measured
the primary of each available specimen of the /unus group on two different
days: 25% of my double readings agreed, and another 60% differed by only
0.1 or 0.2 mm. Differing measurements were averaged before calculation of
| the statistics in Table 1.

In Atrytonopsis, females average larger than males—a prevalent feature
of sexual dimorphism in skippers, as in many other animals. Because fe-
males of Junus are collected more often than males (the reverse is more

Sa among skippers), females afford the better sample (N = 44). With a
/mean primary length of 21.80 mm which exceeds that of males by nearly |
/mm (Table 1), females of /unus are significantly larger than males (tf = 3.16,
P < .005).

The wing lengths of the female type and paratype of frappenda are both
i mm, the same as the mean wing length of /unus females. The third

550 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Frequency of nudum variants in members of the /unus group.

Number of Nudum Segments (on base of club/on apiculus)

Species N 4/7 3/8 4/8 5/8 4/9 5/9 4/10 5/10
A. lunus 46 l 2 9 10 12 1 2 3
A. frappenda 3 | l l
A. zweifeli 2 2

known female of frappenda is smaller (20.4 mm) but well within the female
size range of Junus (Table 1). These scant data suggest that frappenda is
about as large as /unus. The still more limited data for zweifeli—male type
and paratype with wing lengths of 19.5 and 18.1 mm, respectively—fall below
the male size range of /unus (see Table 1, but note that N is only 13) and
suggest that zweifeli is substantially smaller than /unus and frappenda.

Nothing can be said about possible geographic variation in size in any of
these forms except that /unus appears undifferentiated over its discontin-
uous distribution in the mountain islands of southeastern Arizona.

Antenna.—On average, the apiculus is longer in members of the /unus
group than it is in any of the other species of Afrytonopsis that rarely or
never show spots on the dorsal secondary. Among these species, the short-
est apiculus relative to size occurs in deva and a somewhat longer apiculus,
in loammi, hianna (Scudder) plus turneri Freeman, and vierecki (Skinner).
Among those species that usually or always have spots on the dorsal sec-
ondary, the shortest apiculus (in fact, the shortest and stubbiest in genus
Atrytonopsis) occurs in python (Edwards) and margarita (Skinner) and the
longest, in ovinia and edwardsi Barnes and McDunnough. Intermediate
lengths occur in pittacus (Edwards) and cestus (Edwards). Like other con-
tinuously varying characters, the apiculus varies appreciably within species
and populations of Atrytonopsis, though not so much as to obliterate dif-
ferences between species or species groups.

In characterizing genus Atrytonopsis, Evans (1955: 384) pegs the nudum
of the antenna at ‘‘4/6,”’ that is, at a total of 10 segments, of which 4 are on
the base of the club and 6 are on the apiculus. However, in the /unus group,
the number of segments in the nudum is usually 12, 13, or 14; and their |
arrangement is usually 4/8, 5/8, 4/9, or 5/9 (Table 2). The terminal segment is
almost always long.

Scoring nudum segments is complicated by asymmetry in segmentation
between left and right antennae, incomplete sutures on a single antenna,
variation in the scaling that marks the start of the nudum on the club, and
difficulties in pinpointing the junction between the apiculus and the rest of
the club. Some of these problems have been discussed by MacNeill (1964:
51) and Burns (1964: 10). I scored each specimen at least twice.

VOLUME 84, NUMBER 3 551

Because I could detect neither sexual dimorphism nor geographic varia-
tion in the expression of the nudum within the sizable series of A. lunus
from southeastern Arizona, I pooled all data. Although the variate is dis-
continuous, the resulting frequency distribution of nudum variants looks
‘‘normal,’’ especially when the data appear by total number of segments,
regardless of their arrangement on the club:

Number of nudum segments

1] 12 13 14 l

in

Frequency in A. lunus 3 9 22 9 3

Distribution of number of nudum segments is similar, with a range of 5
_ segments and a pronounced peak, in homogeneous samples of Erynnis pro-
pertius (Scudder and Burgess) and E. meridianus Bell (Burns, 1964: 97).

Stigma.—Males belonging to the /unus group lack a stigma. Both this
assertion and the simultaneous assertion that the lack of a stigma helps
define the /unus group contradict important literature.

The end of Evans’ (1955: 384) characterization of genus Atrytonopsis,
‘*$ upf [upper forewing] with a narrow stigma from base vein 3 to mid vein
1, broken at vein 2 and mid space 1Ib,”’ reappears, slightly altered, at the
beginning of Freeman’s (1969: 276) original description of A. zweifeli:
‘*There is a very narrow indistinct stigma from base vein 3 to mid vein 1,
broken at vein 2 and mid space Ib.”’ It is not enough for Freeman to change
‘narrow stigma’’ to ‘“‘very narrow indistinct stigma’’; there is no stigma at
all in A. zweifeli, at least not in the two males from which it was described.

Evans incorrectly attributes a stigma to males of all species of Atryton-
opsis not only in his characterization of the genus, quoted above, but also
in his key to genera of the Lerodea Group where, given the choice “‘d upf
with a stigma or brands’’ versus ‘‘d upf no stigma or brands,’’ one must
choose the former to get to Atrytonopsis (Evans, 1955: 383). In defense of
Evans, recall that zweifeli was then unknown and frappenda, though de-
scribed, unseen by him and therefore left in Lychnuchoides; so the only
species he included in Atrytonopsis whose males never have a stigma was
lunus, and it was represented in the British Museum (Natural History) only
by females (Evans, 1955: 384)!

However, the situation in Atrytonopsis is more complex than all-or-none:
the degree of development of the stigma varies greatly both between and
within species. Most extreme is A. deva which runs a gamut from no stigma,
through many and various vestigial and reduced expressions, all the way to
the complete three-part structure. Of the 160 males of deva that I have
- microscopically examined (mostly from Arizona, but also from New Mexico

313). PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

and Mexico), 40% are virtually or quite without a stigma. Of those 60% that
have one, about half show at least some of each of the three parts; but half
show only a bit (to all) of only one or two parts—upper or middle or lower
or upper plus middle or middle plus lower or upper plus lower. In one other
species outside the /unus group, A. pittacus, a lone male wholly lacks the
stigma; and the stigma is inconspicuous in 30% of the 117 males I have
examined from Arizona, west Texas, and Mexico (and very weak in 8, and
exceedingly reduced in 2, of these). In males of all other species, the stigma,
though somewhat variable, is easy to find with a stereomicroscope; none-
theless, occasional individuals show marked reduction: 3 males (from Man-
itoba) out of 157 males of hianna from eastern North America; | out of 84
males of loammi from Florida; 2 males (from Texas) out of 77 males of
vierecki from New Mexico and west Texas; | male (from Arizona) out of
146 males of edwardsi from Arizona, west Texas, and Mexico; | male (from
Oaxaca) out of 25 males of ovinia from Mexico and Nicaragua; and 8 males
(from Arizona) out of 145 males of python from Arizona and New Mexico.
The stigma is ‘‘always’’ well-developed in margarita (43 males from New
Mexico), turneri (14 males from Kansas and Nebraska), and cestus (10
males from Arizona).

With one important exception, I support MacNeill’s (1975: 443-446) ob-
servations on stigmas in Atryftonopsis. Regarding the more aberrant species,
he writes, ‘‘there is no obvious male stigma’ in /unus (here, ‘‘obvious”’ can
be dropped) and ‘‘no well developed stigma’ in deva (which is a fair average
statement). And he correctly notes the routine occurrence of a stigma in
males of vierecki, pittacus, edwardsi, python, and cestus. However, he errs
in stating that ‘‘The male stigma is absent” in hianna (and, by implication,
in the closely related populations turneri and loammi); males of all three of
these forms always have a stigma.

So I reiterate that only the males of the /unus group never have a stigma.
One may object that no such statement can be made because the male of
frappenda is unknown and the sample of zweifeli is small. But judging from
the divers characters (including geographic distribution) that closely bind
the members of the /unus group, and knowing something of the ways of
evolution, I stick out my neck with confidence.

Facies.—Members of the /unus group always have a bold spot (often dual)
spanning the cell of the primary but rarely any spots on the dorsal secondary
(Figs. 3-8). In dorsal view, the spot in space |b of the primary, when present
(it usually is), is more distal than it is in other species of Atrytonopsis: look
for it under the spot in space 2, not more medial than the spot in space 2.

The fringe of the secondary is yellowish to whitish, at least around the
apex and sometimes all the way to the tornus. At and below the apex, short
scales of the fringe tend more to brown on the dorsal surface of the sec-

VOLUME 84, NUMBER 3 553

ondary but to white on the ventral surface, so that the fringe usually looks
a little wider and a little whiter ventrally than dorsally. Fringes of both wings
are uncheckered.

With data on quantitative and sexual characters still so limited and frag-
mentary, facies seems to provide the best means of distinguishing among
members of the /unus group. To appreciate many of the following points,
keep comparing Figs. 3-8.

The spots of the primary are white in Junus and zweifeli but creamy in
frappenda. The three subapical spots (in spaces 6, 7, and 8 of the primary)
are prominent in /unus and frappenda but small in zweifeli. (I have seen one
male of /unus from the Santa Rita Mountains of southern Arizona with
subapical spots reduced about halfway toward their very reduced level of
expression in zweifeli.) All three females of frappenda have a spot in space
4 of the primary; this spot is tiny in two of them (Figs. 3 and 5) and small
in the other (Fig. 4), which also has a small spot in space 5. By contrast,
both males of zweifeli lack spots in spaces 4 and 5, and Junus never has spots
in these spaces in either sex. Again, all three females of frappenda, but no
specimens of lunus or zweifeli, have pale scales dorsally in spaces 9, 10,
and || forming narrow opaque spots immediately anterior to the large se-
mihyaline cell spot and, in effect, extending the pale primary band to the
costa.

Of the remaining spots on the primary, not only the cell spot but also the
spots in spaces 2 and 3 are always present, whereas the spot in space Ib is
sometimes absent. These spots are smaller, well separated, and relatively
out of line in /unus but much enlarged, in broad contact, and aligned to
form a salient band across the middle of the wing in frappenda; zweifeli is
peculiarly intermediate with only the spots in the cell and space 2 somewhat
enlarged and imperfectly aligned to form a conspicuous patch.

Expression of the spot in space Ib varies greatly, and both sexual and
specific differences contribute to the variation. The spot occurs in one of
the two males of zweifeli, where it appears in the upper part of space Ib,
adjacent to the spot in space 2 (Fig. 8). It occurs in 50% of the males of
lunus, but appears only in the /ower half of space Ib. It occurs in 91% of
the females of /unus, appearing only in the lower half of space 1b in 67%
(Fig. 6) but in both halves in 24% (Fig. 9). It occurs, greatly hypertrophied,
in all three females of frappenda, where it appears in both halves of space
lb and looks like a fat chevron lying on its side and pointing toward the
base of the wing (Figs. 3-5).

Ignoring individual variation invites error. The original description of
zweifeli (Freeman, 1969: 276) specifies that, ‘“‘In space 3 there is a very
narrow elongated spot which curves outward at the lower end.”’ This is true
of the holotype (male) but not of the paratype (also male), in which this spot

Figs. |-4. Females of Lychnuchoides and Atrytonopsis in dorsal and ventral view (all
USNM, all x1). 1, L. saptine, Volcan Santa Maria, Guatemala, October. 2, L. ozias, Castro,
Parana, Brazil. 3, A. frappenda, holotype, Mexico City, Mexico, August 1917, R. Miiller. 4,
A. frappenda, paratype, near Mexico City, Mexico, July, R. Miiller.

VOLUME 84, NUMBER 3 555

8

Figs. 5-8. Atrytonopsis in dorsal and ventral view (all <1). 5, A. frappenda, 2, 7 km SE
Morelos Canada, Puebla, Mexico, 4 to 5 July 1974, J. Chemsak and J. Powell (UCB). 6, A.
lunus, 2, Herb Martyr Camp and Dam, Cave Creek, Chiricahua Mountains, 1770 m, Cochise
County, Arizona, 31 July 1974, J. M. and S. N. Burns (USNM). 7, A. zweifeli, 3, holotype,
1.6 km S Cedritos, Coahuila, Mexico, 23 June 1957, R. Zweifel (AMNH). 8, A. zweifeli, 3,
paratype, Durango-Villa Union highway, route 40, 1980 m, Sinaloa, Mexico, 29 April 1966, P.
Hubbell (AMNH).

556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 9. Dorsal and ventral view of female of Atrytonopsis lunus whose facies varies in the
direction of A. frappenda. Baboquivari Mountains, Arizona, 15 to 31 August (AMNH) (x1).

is quadrilateral and relatively large (cf. Figs. 7 and 8). Typologists might still
defend an original description based solely on the holotype. But there is no
excuse for the general assertion (Freeman, 1969: 277) that, ‘‘lunus has the
spot in space 3 oval and better defined than in zweifeli where this spot is
very linear and indistinct.’’ Comparison of the lone paratype of zweifeli with
males of /unus shows this spot to be as large and distinct in the former as
in the latter (and equally angular in both).

Of the 60 specimens of Junus I have examined, one female (Fig. 9) from
the Baboquivari Mountains of southern Arizona deviates remarkably from
the ‘‘/unus look’’ in the direction of frappenda. This deviant underscores
both the close genetic similarity of the two species and the potential one
species may have for abruptly approximating the superficially different fa- |
cies of the other. The deviant female is clearly /unus: her primary spots are
white rather than cream; dorsally her primary ‘“‘band’’ does not reach the
costa, nor is it as wide or as even along its proximal edge as it is in frap-
penda, and ventrally it is much less striking than it is in frappenda (cf. Figs.
3-5 with 9).

More important, the deviant female is like /unus and unlike frappenda
in a morphologic detail involving the spacing of veins 1, 2, and 3 where they
are nearly parallel in the distal third of the primary: the vertical distance
between veins | and 2 exceeds that between veins 2 and 3 by a lot in /unus
but by relatively little in frappenda. This difference, surprising in what are |
likely sister species, is obvious (once you have seen it): compare Figs. 3—5
(females of frappenda) with Figs. 6 and 9 (females of /unus). Having seen
it repeatedly, I quantified the difference by measuring the height of spaces
lb and 2 with a filar micrometer—its scale through the proximal end of the
spot in space 3, and line 60 directly over vein 2—on the left primary of every
female of Junus and frappenda at hand and then calculating the ratio of the
height of the shorter space to that of the taller. The frequency distribution
of this ratio is quite distinct for the two species (Fig. 10).

VOLUME 84, NUMBER 3 557

6
S 4
Zi
=)
o
m%

2

26055 362 j040 = OO = eoon <0) save) aia 276 S780 SO —820 084 eco mesos ea IO

RATIO OF HEIGHT OF PRIMARY SPACE 2 TO HEIGHT OF PRIMARY SPACE lb

Fig. 10. Frequency distribution of the ratio of the height of space 2 to the height of space
lb on the left primary in females of Atrytonopsis lunus (solid) and A. frappenda (open). The
mean ratio for A. /unus (N = 31) is indicated.

Genitalia.—Although male genitalia of Atrytonopsis have been examined
and figured, more or less inadequately, for nearly a century (Scudder, 1889:
pl. 37, fig. 36; Skinner and Williams, 1924: figs. 21-28; Williams and Bell,
1930: fig. 1; Lindsey et al., 1931: pl. 7, fig. 5; pl. 29, figs. 21-24; pl. 30, figs.
25-28; Evans, 1955: pl. 82, figs. N.1.1-N.1.8; Forbes, 1960: fig. 129), female
genitalia have been wholly ignored.

Both sexes of Atrytonopsis have genitalia so conservative in essential
form that they loudly proclaim their genus and so individually variable that
they may scarcely whisper which species. Facies, though also individually
variable, is generally easier to use in distinguishing differentiates. Because
interspecific genitalic differences may be subtle, or quantitative rather than
qualitative, series of dissections should be closely and repeatedly compared.

The taxonomically most useful genital element in males of Afrytonopsis
is the uncus, especially its form, but also its position in relation to the
gnathos. (Unexpectedly, the valvae are next to useless.) In females, the
most useful element is the lamella postvaginalis, which amounts to little
more than midventral sclerotization projecting caudad from the dorsal rim
of the ostium bursae (sometimes expanding laterad, as well) and bearing
many short fine setae. Variation in this simple device deserves all-out scru-
tiny.

The 200 Atrytonopsis genitalia dissected in connection with this study
spent 6 to 9 minutes in boiling 10% KOH and, after thorough cleaning, went
with their descaled abdominal skins into |-dram vials of 80% ethanol plus
3 drops of glycerin. They never wound up on slides because slides severely

558 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 11-13. Male genitalia of Atrytonopsis lunus from Apache Indian Reservation, White
Mountains, 1830-2440 m, Arizona, 20 to 27 July 1925, O. C. Poling (X-1082) (USNM). 11,
Uncus, gnathos, and tegumen in dorsal view and uncus, gnathos, tegumen, vinculum, and

saccus in left lateral view. 12, Left valva, aedeagus, and juxta in left lateral view. 13, Aedeagus
and juxta in dorsal view.

VOLUME 84, NUMBER 3 359

Figs. 14-16. Male genitalia of paratype of Atrytonopsis zweifeli from Durango-Villa Union
highway, route 40, 1980 m, Sinaloa, Mexico, 29 April 1966, P. Hubbell (X-1062) (AMNH). 14,
Uncus, gnathos, and tegumen in dorsal view and uncus, gnathos, tegumen, vinculum, and
saccus in left lateral view. 15, Left valva, aedeagus, and juxta in left lateral view. 16, Aedeagus
and juxta in dorsal view.

limit angles of observation, preclude direct side-by-side comparison of dif-
ferent individuals, and may distort genitalic form. Genitalia dissection num-
bers of drawn individuals appear in figure legends in parentheses.

Male genitalia: In the lunus group (Figs. 11-16) and those other species

560 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

21 22 23 24 25

Figs. 17-25. Female genitalia of Atrytonopsis lunus from southeastern Arizona (all USNM).
17, Bursa copulatrix and tip of abdomen in ventral and right lateral view. 18-25, Lamella
postvaginalis, ostium bursae, and ductus bursae in ventral view. 17, Tonto Basin, Gila County
(X-1079). 18-21, ‘‘Redington’’ (probably Santa Catalina Mountains), Pima County (X-1050,
1078, 1141, and 1142). 22, E Nogales, Santa Cruz County, 9 August 1963, A. R. Moldenke (X-
1083). 23-24, Huachuca Mountains, Cochise County, 16 to 23 August (X-1077 and 1084). 25,
Chiricahua Mountains, Cochise County, 24 to 31 July (X-1076).

VOLUME 84, NUMBER 3 561

Figs. 26-28. Female genitalia of Atrytonopsis frappenda from southcentral Mexico. 26,
Bursa copulatrix and tip of abdomen in ventral and right lateral view. 27-28, Lamella post-
vaginalis, ostium bursae, and ductus bursae in ventral view. 26, Holotype from Mexico City,
August 1917, R. Miller (X-1059) (USNM). 27, Paratype from near Mexico City, July, R. Miiller
(X-1058) (USNM). 28, 7 km SE Morelos Canada, Puebla, 4 to 5 July 1974, J. Chemsak and J.
Powell (X-1153) (UCB).

of Atrytonopsis that usually or invariably lack spots on the dorsal secondary
(i.e., deva, vierecki, loammi, and hianna including turneri), uncus and gna-
thos end in long paired prongs that are widely separated from one another
both horizontally and vertically (Figs. 11 and 14). The uncus prongs do not
(or barely) converge toward their tips. Though the horizontal gap between
uncus prongs usually exceeds the vertical gap between uncus and gnathos
prongs, the vertical gap is still large.

Uncus and gnathos prongs in the /unus group are a little shorter than they
are in the rest of the species that usually or always lack spots on the dorsal
secondary. Because the difference is not only small but also variable, it is
elusive. In the /unus group, uncus prongs are shorter than gnathos prongs
(see Figs. 11 and 14); though this difference is variable, too, it is often
conspicuous. Uncus prongs are usually shorter than gnathos prongs in deva,

562 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

as well; but the difference is a trifle less pronounced, on average (and, as
noted above, both pairs of prongs are slightly longer than they are in the
lunus group). By contrast, uncus prongs are usually as long as, or longer
than, gnathos prongs in vierecki and in loammi and hianna (plus turneri).

I cannot separate /unus and zweifeli genitalically on the basis of available
material. Differences between the drawn specimens of /unus and zweifeli
are apparently individual rather than specific. For example, the sclerotized
arm on the left side of the flared distal end of the aedeagus bears 3 teeth in
lunus (Figs. 12 and 13), 4in zweifeli (Figs. 15 and 16). But the other known spec-
imen of zweifeli (the holotype) has 3 teeth on this arm; and in the series of
lunus dissections I have examined, these teeth vary from 2 to 4, with a
mode of 3. Again, the right cornutus ends in a single point in the drawn
individual of /unus, in two points in that of zweifeli. But there is only one
point on the right cornutus in the holotype of zweifeli; and though there is
only one in the other dissections of /unus, two are likely to occur in occa-
sional individuals in view of the fact that the number of points always varies
by one or two in longer series of dissections of such related species as deva,
vierecki, and hianna. Finally, valval shape often varies more within species
of Atrytonopsis than it does between the drawn individuals of /unus and
zweifeli, and aedeagal length varies intraspecifically as well as interspecifi-
cally.

Freeman’s (1969: fig. 19) figure of the genitalia of the holotype of zweifeli
is all wrong. It shows an uncus prong longer than its corresponding gnathos
prong (the reverse is true) and misrepresents both the shape of the uncus
and the nature of its union with the tegumen. The vinculum is oversimpli-
fied; the valva, inaccurate (most seriously along its dorsal and distal mar-
gins); and the aedeagus, far too short. In Freeman’s genitalic dissection of
the holotype, vinculum and saccus are badly torn, and at least the anterior
half of the aedeagus is missing. This piece may still be in the type-specimen
because Freeman removed only the distal part of its abdomen (see Fig. 7),
even though the aedeagus has long been known to be notably long in Atry-
tonopsis. He must, then, have manufactured the anterior end of the aedea-
gus in his figure. In his text where he compares zweifeli with lunus, Freeman
(1969: 277) says only that “‘there are differences in the genitalia.’’ He neither
provides comparative figures nor analyzes and describes the ‘‘genitalic dif-
ferences’? he claims to see, and he does not check series of genitalia for
variation. Unfortunately, such broad statements as ‘‘the genitalia are dif-
ferent’? often appear, without amplification, in Freeman’s descriptions of
new species.

Female genitalia: In the lunus group (Figs. 17-28), as in females of Afry-
tonopsis generally, the ductus bursae is large in diameter, slightly bowed to
the left, and heavily sclerotized from end to end, whereas the corpus bursae

VOLUME 84, NUMBER 3 563

X e lunus

a zweifeli

s frappenda

MEXICO

100 200 300
————————
wines

|
L Ss
ma 112° 110" roa” 106° TT

Fig. 29. Geographic distribution of the Junus group of Afrytonopsis (based on material
examined).

is modest in size and fully membranous. In the /unus group and a number
of other species, the ductus bursae is not exceedingly long but is wide and
is conspicuously flared at its posterior end (i.e., at the ostium bursae), where
the sclerotized walls of the ductus bursae extend much farther caudad both
dorsally and ventrally than laterally (see particularly the lateral views in
Figs. 17 and 26).
Slight bilateral swelling of the sort shown in Figs. 17—28 near or at the
anterior end of the ductus bursae is peculiar to the /unus group.
In dorsal or, better, ventral view, the quietly distinctive lamella postva-
ginalis of Junus and frappenda suggests a toadstool in outline; its projecting
cap lies just anterior to the ovipositor lobes (=papillae anales) and its stalk,
- embracing the midline like a pair of parentheses in reverse, merges with the
ductus bursae dorsally (Figs. 17-28). The whole device, especially the cap,
is densely felted with fine setae.

Usually the cap is flatter and broader in frappenda (Figs. 26-28), more
sharply humped and less wide in /unus (Figs. 17-25).

Temporal distribution.—Among the nine or so species of Afrytonopsis in

564 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

the United States, /unus is peculiar because it flies in summer rather than
spring. The spring-flying species variously appear from March to June (es-
pecially in April, May, and June), and some reappear in latter summer and
fall. Every dated specimen of /unus that I know of (N = 188, including 148
courtesy K. Roever) is from July or August (save one from June and one
from September); and most are from a one-month span, 15 July to 15 Au-
gust. Adult emergence is almost certainly geared to summer rains.

Presumed misdated and so discounted is one male of /unus, in AMNH,
collected by O. C. Poling in the Baboquivari Mountains, Arizona, 15 to 30
March 1924. Williams and Bell (1930: 137) published these data without
questioning the date.

The fact that the three known specimens of frappenda are also from July
and August might suggest a distinctive flight period for the /unus group as
a whole. But zweifeli breaks the mold: although one specimen is from latter
June, the other is from late April. So few examples of Atrytonopsis have
been collected in Mexico that it is hard to generalize about flight periods
there except to say that they seem to depart from those evident in the United
States, becoming more spread out in time and more continuous.

Spatial distribution.—The /unus group occurs at middle elevations—1190
to 2240 m (3900-8000 ft), usually 1555 to 2010 m (5100-6600 ft)—from the
southwestern United States to southcentral Mexico (Fig. 29). The three
species appear to be limited in distribution and allopatric, with wide inter-
vening gaps: /unus is known from the Mogollon Rim country south through
the mountain islands of southeastern Arizona and extreme southwestern
New Mexico—see Fig. 30 for details (K. Roever, personal communication
-1981): zweifeli, from both the Sierra Madre Oriental (southern Coahuila)
and Occidental (southern Sinaloa); and frappenda, from the Distrito Federal
and Puebla. Undoubtedly, each is more widespread in Mexico than available
data show: /unus must occur at the very least in some montane parts of
Sonora and Chihuahua; zweifeli, in some mountains on the Central Plateau
as well as more extensively in the ranges that flank it; and frappenda, in
more of the Volcanic Cordillera and mountains to the south, plus perhaps
in mountains to the north. But whether any two of these species make
contact or overlap is anybody’s guess.

In broad ecologic terms, all three are skippers of woodland and scrub:
lunus is most often in oak (-pine-juniper) woodlands; the type of zweifeli
was in a dry pinyon-yucca association on rocky limestone (R. G. Zweifel,
personal communication 1981); and the Puebla female of frappenda was
near agave and thorn scrub on rocky limestone (J. A. Powell, personal
communication 1981).

Bell (1942: 465) reported that the collection of R. Muller contained one
male of /unus taken in Mexico, Distrito Federal, in August. Since R. Miller

VOLUME 84, NUMBER 3 565

Fig. 30. Detailed geographic distribution of Atrytonopsis lunus (based on personal records
of K. Roever). The map centers on Arizona and adjacent New Mexico; /unus occurs in ten
counties, outlined with dashes, from northwest to southeast, as follows: Gila, southern Navajo,
southern Apache, eastern Pinal, Graham, Greenlee, eastern Pima, Santa Cruz, Cochise, and
Hidalgo.

took both the type and paratype of frappenda in this very area in July and
August, since (as shown earlier in this paper) at least females of Junus and
frappenda can run close, and since Bell knew neither frappenda nor zwei-
feli, there is some question about the true identity of this male.

A female of Junus in AMNH bears a pair of phony labels (roman type
denotes machine-printed information; italic, information inked in by hand):

566 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

[1] Santa Cruz
IX -14-1923 Cal
ZjtO.1467Polme

I have long since marshalled enough evidence for an airtight case against
this recurring and treacherous pair of labels (Burns, 1964: 110). The data
have been listed for /unus (Williams and Bell, 1930: 137), but publication
does not make them valid.

DISCUSSION

Two-thirds of the North American species of Erynnis neatly cluster in
half a dozen essentially transcontinental superspecies each comprising 2 or
3 species (Burns, 1964, and unpublished). Atrytonopsis, too, shows signs of
considerable recent speciation (though most patterns of splitting differ from
those in Erynnis). At present, the /unus group—three closely related, allo-
patric species replacing one another from northwest to southeast—looks like
a superspecies (Fig. 29). Especially with zweifeli intermediate as regards
both development of the band on the primary and geography, it is tempting
to assume that lunus, zweifeli, and frappenda are still distributed much as
they were when they speciated. This assumption may be wrong because
lunus and frappenda, which occur at opposite ends of the /unus group’s
‘‘linear’’ distribution, share features suggesting that each is the other’s near-
est relative. Given such tiny samples for inferring relationship and total
range (two males of zweifeli, three females of frappenda), I will not speculate
further.

ACKNOWLEDGMENTS

Thank you: For specimens lent, Frederick H. Rindge and the American
Museum of Natural History, New York (AMNH) and Jerry A. Powell and
the Essig Museum of Entomology, University of California, Berkeley
(UCB); for data from personal notes, Jerry A. Powell, Richard G. Zweifel,
and especially Kilian Roever; for 200 exquisite dissections of male and fe-
male genitalia and for statistical calculations, Richard G. Robbins; for pho-
tographs, Victor E. Krantz; for drawings of genitalia plus a histogram,
George L. Venable; for skill with a word processor, Noreen M. Connell;
and for critical review of the manuscript, C. Don MacNeill and Robert K.
Robbins.

LITERATURE CITED

Bell, E. L. 1942. New records and new species of Hesperiidae from Mexico. An. Esc. Nac.
Cienc. Biol. Mex. 2: 455-468.

Burns, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Calif. Publ.
Entomol. 37, 216 pp.

VOLUME 84, NUMBER 3 567

Dyar, H. G. 1920. New Lepidoptera, chiefly from Mexico, with synonymic notes. Insecutor
Inscitiae Menstruus 8: 187-198.

Evans, W. H. 1955. A catalogue of the American Hesperiidae indicating the classification and
nomenclature adopted in the British Museum (Natural History). Part IV. Hesperiinae
and Megathyminae. British Museum, London. 499 pp., pls. 54-88.

Forbes, W. T. M. 1960. Lepidoptera of New York and neighboring states. Part IV. Cornell
Univ. Agric. Exp. Stn. Mem. 371, 188 pp.

Freeman, H. A. [1969] **1968.’" New species and records of Hesperiidae from Mexico (Lep-
idoptera). J. N.Y. Entomol. Soc. 76: 267—277.

Godman, F. D. 1900, 1901. Jn Godman, F. D. and O. Salvin, 1879-1901. Biologia Centrali-
Americana. Insecta. Lepidoptera-Rhopalocera. Vol. I, xlvi + 487 pp.; vol. II, 782 pp.;
vol. III, 113 pls.

Hoffmann, C. C. 1941. Catalogo sistematico y zoogeografico de los Lepidopteros Mexicanos.
Segunda Parte. Hesperioidea. An. Inst. Biol. Mex. 12: 237-294.

Lindsey, A. W., E. L. Bell, and R. C. Williams, Jr. 1931. The Hesperioidea of North America.
Denison Univ. Bull., J. Sci. Lab. 26, 142 pp.

MacNeill, C. D. 1964. The skippers of the genus Hesperia in western North America with

special reference to California (Lepidoptera: Hesperiidae). Univ. Calif. Publ. Entomol.

35, 230 pp.

. 1975. Family Hesperiidae, pp. 423-578. In Howe, W. H., ed., The Butterflies of North

America. Doubleday and Co., Inc., Garden City, N.Y. 633 pp., 97 pls.

Okano, K. 1981. Studies on the Mexican butterflies (1). Additions and revisions to . . . Hoff-
mann’s ‘‘Catalogo sistematico y zoogeografico de los Lepidopteros Mexicanos.”’ Part 1.
Hesperiidae (Lepidoptera: Rhopalocera). Tokurana (Acta Rhopalocera) No. 1: 23-139.

Scudder, S. H. 1889. The butterflies of the eastern United States and Canada with special
reference to New England. Publ. by the author, Cambridge. Vol. I, pp. i-xxiv, 1-766;
vol. II, pp. i-x, 767-1774; vol. II, pp. i-vi, 1775-1958, pls. 1-89, 3 maps.

Skinner, H. and R. C. Williams, Jr. 1924. On the male genitalia of the Hesperiidae of North
America. Paper IV. Trans. Am. Entomol. Soc. 50: 57-74.

Williams, R. C., Jr. and E. L. Bell. 1930. Short studies in American Hesperiidae (Lepidoptera:
Rhopalocera). Trans. Am. Entomol. Soc. 56: 133-138.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 568-585

THE CACAO-POLLINATING MIDGES OF THE
FORCIPOMYIA ARGENTEOLA GROUP
(DIPTERA: CERATOPOGONIDAE)

WILLIS W. WIRTH

Systematic Entomology Laboratory, IIBIII ,Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Diagnoses and keys are given for the Forcipomyia argenteola
Group of species of the subgenus Forcipomyia s. str. Included in this group
are F. argenteola Macfie from southern Brazil; guatei Wirth, ranging from
the southern United States to southern Brazil; calatheae, n. sp., from Brazil,
Colombia, and Dominica; and youngi, n. sp., from Costa Rica, Ecuador,
and Panama. Because of its close similarity to species of the argenteola
Group, diagnostic notes are also given for the widespread Neotropical
species, squamitibia Lutz. Larvae of species of the argenteola Group are
commonly found in rotting banana stems, cacao pods, bracts of Calathea
and Heliconia, and similar decaying vegetable matter; and the species are
often abundant in cacao plantations where the adults may serve as polli-
nators.

Beginning with the classic work of Saunders (1956, 1959), intensive stud-
ies have been made on the immature stages and classification of the For-
cipomyia midges associated with cacao culture. Although the smaller adults
of the subgenera Euprojoannisia Brethes, Thyridomyia Saunders, and
Warmkea Saunders are most important in cacao pollination, the very abun-
dance and ubiquity of the species of the subgenus Forcipomyia s. str. also
place these midges high on the list of important cacao pollinators. Previous
publications have reported on the taxonomy of all stages of Euprojoannisia
(Saunders, 1956; Soria and Bystrak, 1975; Bystrak and Wirth, 1978); Thyr-
idomyia (Saunders, 1956); and Warmkea (Wirth and Soria, 1980). Wirth and
Soria (1975) described all stages of Forcipomyia (Forcipomyia) genualis
(Loew) and a closely related species, F. harpegonata Wirth and Soria.
Wirth (1976) described all stages of F. (F.) pictoni Macfie and two new
closely related species from Florida, F. seminole Wirth and F. beckae
Wirth.

VOLUME 84, NUMBER 3 569

The larval habitats of these Forcipomyia midges are somewhat correlated
with their taxonomic group; the habitats of Euprojoannisia species are usu-
ally semiaquatic situations such as algae-covered rocks or mud, wet moss,
or leaves, mats of decaying aquatic vegetation, and leaf axils of water-hold-
ing plants. Species of Thyridomyia usually are found in association with
moss or algae in rather damp habitats. Immature stages of Warmkea species
are frequently found in leaf axils of Pandanus, aroids and epiphytic and
terrestrial bromeliads, and less often in rotting cacao leaves on the ground,
in banana stems, and in bracts of Heliconia. Species of the subgenus For-
cipomyia are less aquatic and are more commonly associated with rotting
plant material, which in cacao plantations and their environs often involves
heaps of cacao pods, old banana stems, cacao leaves, coconut debris, bracts
of Heliconia and Calathea, and rotting fruits of coconut, calabash, palm
nuts, etc.

Over the past few years I have received numerous samples of Forcipo-
myia midges closely resembling F. argenteola Macfie. Dr. Allen M. Young,
working in Costa Rica, found them especially abundant in decaying sections
of banana stems he set out in cacao plantations to attract ovipositing female
midges and thus to sample the relative abundance of various ceratopogonid
species. Although the adults of these species were almost indistinguishable
from each other, larval and pupal characters were found to separate two

distinct species in Costa Rica, neither of which proved to be argenteola.

——

On closer study, adult characters were also discovered. A review of the
material in the National Museum of Natural History, Smithsonian Institu-
tion, disclosed that there are at least four species in this group, closely
related to argenteola. One species, quatei Wirth (1952), ranges from Florida
and California to southern Brazil; argenteola apparently is restricted to
southern Brazil; a third species, calatheae n. sp., is recorded from Brazil,
Colombia and Dominica; and the fourth, youngi n. sp., is found in Costa
Rica, Ecuador, and Panama. In this paper I am presenting descriptions or
diagnoses, figures, and a key for the separation of these species. Although
it is not a member of the argenteola Group, Forcipomyia squamitibia Lutz
adults are so similar to members of that group that a diagnosis is also pre-
sented for squamitibia.

A discussion of the taxonomic characters used in this paper can be found
in the publications by Saunders (1924, 1956), Wirth (1952), Chan and
LeRoux (1965), and Bystrak and Wirth (1978). Holotypes of the new species
are deposited in the National Museum of Natural History, Smithsonian In-
stitution, Washington, D.C.; paratypes will be deposited in the Museu de
Zoologia, Universidade de Sao Paulo, Brazil, the British Museum (Natural
History), London; the Muséum National d'Histoire Naturelle, Paris; the
California Academy of Sciences, San Francisco; and the Milwaukee Public
Museum, Milwaukee, Wisconsin.

570 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

I am greatly indebted to the following individuals and institutions for the
submission of material and for their interest and efforts in collecting and
rearing all stages of the most common Forcipomyia midges in American
cacao plantations: The late Leslie G. Saunders, working in Costa Rica,
Mexico, Puerto Rico, and Trinidad, with financial assistance from the
American Cocoa Research Institute; Saulo de J. Soria, Centro de Pesquisas
do Cacau, Itabuna, Bahia, Brazil; Eduarto J. Ureta S., Sanidad Agrope-
cuaria, Dept. Antioquia, Medellin, Colombia; and Allen M. Young, Mil-
waukee Public Museum, Milwaukee, Wisconsin, working in Costa Rica un-
der grants from the American Cocoa Research Institute.

Forcipomyia (Forcipomyia) argenteola Group

Diagnosis.—Legs yellowish, at most tarsi and apex of hindfemur infus-
cated; hindtarsal ratio 0.50—0.76; female wing brownish with one small yel-
lowish spot at end of costa, base of costa sometimes yellowish; male wing
usually extensively pale; male abdomen banded and genitalia yellowish with
pattern of brownish infuscation; 3rd palpal segment swollen at base and
bearing a deep round sensory organ; female tibiae without hastate spines;
spermathecae ovoid, rather elongate, tapering to slender opening without
elongate neck; male aedeagus poorly sclerotized, in form of an elongated
shield, more than 1.5 as long as basal breadth, with a submedian pair of
faint longitudinal ridges; parameres with bases broadly fused, caudal pro-
cesses moderately stout, each tapering to a sharp tip.

Larva hypognathous, typical of subgenus Forcipomyia; body yellowish
white, without cuticular armature, head capsule brownish to blackish. Pro-
thoracic pseudopods short and cleft a short distance, with 6-8 pairs of black-
ish hooks; posterior pseudopod with a double row of 16-20 brownish hooks.
Head with p and q hairs long and curved or bent, slender or with tips slightly
expanded. Body hairs b and c arising from a large conical common prom-
inence; a hairs moderately long and distinctly hastate, their bases often
connected across midline by a narrow blackish pigmented line; b, c, and d
hairs elongate, usually microscopically fringed; e and f hairs moderately
short to long, simple, a, b, and e hairs pale; c, d, and f hairs pigmented.

Pupa moderately stout; integument without conspicuous armature except
for the usual 6 pairs of dorsal thoracic tubercles; abdomen sometimes with
| or 2 pairs of elongate lateral processes and some inconspicuous spinose
tubercles in transverse dorsal row on each segment. Prothoracic respiratory
horn small to moderate in size, with short basal petiole, distal portion var-
iably swollen in a rounded or globular knob bearing a row of 12-35 spiracular
openings along apex and down dorsal side. Last abdominal segment drawn
out in a pair of long tapering processes, at least twice as long as basal

VOLUME 84, NUMBER 3 571

breadth of segment. Last instar larval exuviae remain attached to posterior
end of pupal abdomen.

i)

to

KEY TO SPECIES OF THE FORCIPOMYIA ARGENTEOLA GROUP

. Female tibiae with row of hastate spines on extensor surface; pale

wing spot at end of costa larger, including most of 2nd radial cell;
hindtarsal ratio 1.05; male parameres with slender common
ASCO COINS tits Aen Sink ai ee ee squamitibia Lutz
Female tibiae without hastate spines; pale wing spot smaller, not
covering end of 2nd radial cell; hindtarsal ratio 0.50-0.76; male par-
ameres with bases joined in a broad plate (argenteola Group) ...... 2

. Abdomen without broad dark brown scales; femora and tibiae with-

out broad scales, entirely yellowish; halter slightly infuscated; hind-
tarsal ratio 0.72; female mandibular teeth vestigial; male parameres
fused on proximal 14; larval head entirely dark brown............

MET A: Se AIRE he EE ae Ba ak calatheae, new species
Abdomen with broad dark brown scales; femora and tibiae with
some broad scales; hindtarsal ratio 0.42—0.56; apex of hindfemur
pale or dark; mandible, male parameres, and larval head various
(SESIBElOM DIRS hs QR Bh es bic ans ide Sts See ee 3
Hindfemur brownish on approximately distal 4, this portion bearing
broad brownish scales; mandibular teeth vestigial; 3rd palpal seg-
ment rather slender, swollen on proximal %; halter infuscated

PASE ele oR CE PERS Ye, eee argenteola Macfie
Femora and tibiae entirely yellowish (hindfemur brown at tip in
some quatei, in which case broad scales are pale); mandible with
melndeveloped tecthahalter pale On danrk 2320 .ay0s +l bee eeeeen ee 4

. Halter brownish; female mandible with about 20 small teeth; larval

Het wuMCOlOrOUSipAlesDEOW iil) 26, 33 brs ole ek, ona laye ee ae quatei Wirth
Halter pale; female mandible with 12-15 strong teeth; larval head
veLyedark DIOWwn. toward MOUtD, «. 0%. c025. + 2c youngi, new species

KEY TO KNOWN LARVAE

. Head hairs p and q swollen distally, not much longer than antenna;

head pale brownish, dark only along oral margin; bases of dorsal
DOG halcsS NOt SILOMELY DIBMENTEG ae, scan spp aye ieee yt quatei Wirth
Head hairs p and gq slender distally, more than twice as long as
antenna; head dark brown, especially toward oral margin; bases of
dorsal Body hairs sironely plemented . 24.2,.5).5 3, ase pec Sas: chee bie tena ew See 2
Head strongly pigmented, blackish on anterior ¥%2 toward mouth; e
and f hairs shorter than c and d hairs; a hair of last segment slender,
PATRAS TAR CUES TALI 6. oUt aac dhe Athena betia aa paca youngi, new species

572 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

— Head unicolorous dark brown; e and f hairs much longer than c and
d hairs; a hair of last segment hastate ....... calatheae, new species

KEY TO KNOWN PUPAE

1. Dorsal tubercles of thorax short and inconspicuous, not longer than
basal breadth; respiratory horn with petiole stouter, bearing 30-35
spiracular openings in a much convoluted row; each abdominal seg-
ment with 2 pairs of lateral processes ........ calatheae, new species

— Dorsal tubercles of thorax in form of elongate process at least twice
as long as basal breadth; respiratory horn with slender petiole, bear-
ing 12-25 spiracular openings; each body segment with | pair of
lateral’ processes: or<with none); esen ce Satin. Le ee Ae 2

2. Respiratory horn slender, the slightly swollen apex with 12-15 spi-
racular openings; abdominal segments without conspicuous lateral
processes; thoracic processes torchlike, the pair just behind level of
respiratory horn with seta less than 4 as long as process.........

JERR SAAS BS We a Se ie a ol Fa quatei Wirth

— Respiratory horn globular, with 20-25 spiracular openings; abdom-
inal segments each with | pair of elongate lateral processes; dorsal
thoracic processes between respiratory horns with apical seta about
1 aS lonp aS processys as ah Rae youngi, new species

Forcipomyia argenteola Macfie

Forcipomyia argenteola Macfie, 1939: 146 (2; Brazil; fig. palpus, antenna,
hindleg, radial cells, spermatheca); Lane, 1947: 163 (¢, larva, pupa; Bra-
zil; fig. d genitalia).

Female diagnosis.—Wing length 1.1—-1.8 mm. Wing with a small yellow
spot at end of costa; costal ratio 0.45. Halter pale brownish. Thorax brown,
scutellum yellowish brown. Legs yellowish, hindfemur infuscated at apex,
sometimes infuscation continued across knee to base of tibia; fore- and mid-
femora sometimes slightly darker at tip, but this apparently due to more
numerous, broad, striated, brownish scales there than elsewhere on legs;
tarsi pale brownish; hindtarsal ratio 0.50-0.55. Palpus more slender than in
related species; 3rd segment swollen on proximal 1%; palpal ratio 3.0. Pro-
boscis longer than in related species; mandible with vestigial teeth. Abdo-
men with numerous dark brown, appressed, broad scales; penultimate seg-
ment yellowish; spermathecae ovoid, tapering to slender opening, without
distinct necks.

Male diagnosis.—Wing length 1.5 mm. Coloration similar to that of fe-
male, dark area on hindfemur broader; wing pale brownish, veins slightly
darker, dark brown on radial field, yellow spot at end of costa prominent;
costal ratio 0.40. Hindtarsal ratio 0.42. Abdomen yellowish with prominent

VOLUME 84, NUMBER 3 573

dark brown segmental bands; genitalia similar to those of F. youngi, brown-
ish except at base of 9th segment and on bases of basistyles, which are
yellow.

Distribution.—Brazil.

Types.—Brazil, Santa Catarina, Nova Teutonia, 15.iii.1937, and
1.vill. 1938, F. Plaumann, 2 2 syntypes (British Museum (Nat. Hist.)).

Specimens examined.—BRAZIL: Santa Catarina, Nova Teutonia, var-
ious dates, 1965-1971, F. Plaumann, 4 6,7 2 (USNM).

Immature Stages.—I have not seen the immature stages of this species.
Lane’s (1947) descriptions of the larvae and pupae are brief and typical of
the group. The pupal description as translated below is most diagnostic:

‘‘Pupa. Respiratory tube constricted at the base, expanded and rounded
toward the apex where it terminates in approximately 10 spiracles.

‘“*Cephalothorax with the setae (anterior, dorsal, and marginal) small,
smooth, and inserted on small mammillae; dorso-lateral tubercle conical,
smooth and with a small smooth seta on the apex; dorsal tubercles conical,
spiculose, and with blunt apices.

‘‘Abdomen: Segments I to VII glabrous, VIII and IX spiculose, the spic-
ules in the form of minute scales. Segments I to IV with the postero-marginal
tubercles elongated, spiculose and from whose apex go off small smooth
setae inserted on the middle of the lateral margin; in addition to these pro-
cesses there are on the median portion other much smaller ones, likewise
with smooth setae inserted on mammillae. Segment V like the preceding
except for the postero-marginal tubercle which is defective. Segments VI
and VII with the postero-marginal setae inserted on mammillae which are
distinct, elongated, and uniform at the base, the apex transformed into a
pennant, the internal much smaller than the external. Besides these setae
there are discrete spicules. Segments VIII with only 2 setae like pennants.
Last segment with apex blunt and the terminal filaments longer than the
body of the segment.”’

Forcipomyia calatheae Wirth, NEw SPECIES
Fig. |

A brownish species with pale yellow legs; wing dark brown with a small
yellow spot at end of costa, male wing yellowish with dark brown macro-
trichia over radial veins and infuscation over vein Cu,; halter brown.

Female Holotype.—Wing length |.18 mm; breadth 0.51 mm.

Head: Brown, with abundant long bristly brownish setae. Antenna (Fig.
la) short, proximal segments moniliform; flagellar segments with lengths in
proportion of 16-13-13-13-13-13-13-13-20-20-20-20-25, antennal ratio 1.00.
Palpus (Fig. 1b) with lengths of segments in proportion of 9-11-30-13-12; 3rd
segment moderately swollen on proximal 2, bearing a deep sensory pit

574 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Forcipomyia calatheae. a—d, Female. e-g, Male. h-l, p-r, Larva. m—o, Pupa. a,
Antenna. b, Palpus. c,e, Wing. d, Spermathecae. f, Parameres. g, Genitalia, parameres re-
moved. h-l, a, b, d, c, e, f hairs respectively, of 2nd abdominal segment. m, Lateral processes
of abdomen. n, Respiratory horn. 0, Thoracic tubercles. p, Head, lateral view. q, Second
abdominal segment, lateral view. r, Two posterior abdominal segments, lateral view.

VOLUME 84, NUMBER 3 575

opening by a slightly smaller pore; palpal ratio 2.1. Mandibular teeth ves-

tigial.

Thorax: Brown; mesonotum and scutellum with long bristly hairs and
shorter finer setae. Legs unicolorus yellowish with brownish tinge due to
numerous flattened, narrow appressed scales; tarsi concolorous with tibiae;
femora and tibiae stout, tibiae with longest bristly hairs on extensor surface
about 3x diameter of tibia; hindtibial spur short and inconspicuous, pointed,

_ about % as long as diameter of tibia. Tarsi with ventral palisade setae prom-
-inent on tarsomeres | and 2 of fore- and midlegs, poorly developed on
hindleg; some stronger dark ventral spines scattered on tarsomeres | and

2 of hindlegs, apical only on remaining tarsomeres 1-4. Tarsal ratios 1.00
on foreleg, 0.70 on midleg, and 0.76 on hindleg. Wing (Fig. Ic) brown,

darker anteriorly, with dense, long, brown macrotrichia, a small yellow spot

at end of costa; costal ratio 0.44. Halter brown.

_ Abdomen: Brown, last 2 segments and cerci paler; vestiture of rather
short brown hairs, those on pleura less conspicuous; terga with some narrow
brown scales. Spermathecae (Fig. 1d) 2, dark brown, elongate ovoid, ta-
_ pering gradually to slender opening to duct; subequal, each measuring 0.087
by 0.043 mm.

Male allotype.—Wing length |.37 mm; breadth 0.40 mm. Similar to female
with usual sexual differences. General color as in female, but wing (Fig. le)
pale yellowish brown, slightly darker anteriorly and a line along vein Cu,,
costal and radial veins dark brown due to prominent long slender, scalelike
macrotrichia. Abdomen with broad brown segmental bands bearing long
yellowish to brown, bristly setae. Costal ratio 0.42; tarsal ratios 0.82 on
foreleg, 0.65 on midleg, and 0.72 on hindleg.

Genitalia (Fig. 1g): Basistyle and distal part of 9th segment brown, dis-
tistyle and midportion of 9th sternum pale. Aedeagus elongate, nearly tri-
angular, about 2x as long as basal breadth, poorly sclerotized, apex nar-
rowly pointed. Parameres (Fig. If) poorly sclerotized, yellowish bases fused
for % the total length of parameres, the caudomedian emargination forming
an acute angle; caudal processes slender with sharp-pointed apices.
Pupa.—Length 2.5-3.0 mm. Color brownish, abdomen paler. Respiratory
horn (Fig. In) on distinct stalk, the distal portion 1.5-2 as broad as stalk
depending on angle viewed, and about as long as stalked portion; bearing
about 30-35 spiracular openings in a single, much convoluted row around
apex and partway down dorsal side of swollen portion. Thoracic tubercles
(Fig. lo) low and conical, bearing a fine seta or fine spicules as figured.
'Each abdominal segment (Fig. 1m) with 2 pairs of spiculate lateral process-
es, their apices with several short spines.

Larva.—Length 4—5 mm when mature. Color yellowish white; head cap-
sule (Fig. Ip) uniformly brownish; dorsal body hairs borne on prominent

576 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

dark brown conical prominences as figured (Fig. 1q). Hairs p and q of head
long and slender, of uniform width to tip, pale yellowish. Body hairs a, b,
and e pale, c, d and f brownish (Figs. |h-l); b and d hairs arising from a
common tubercle. Last 2 segments as in Fig. Ir. In mature larvae and larval
exuviae, the a hairs connected across midline by a slender black sclerotized
line; a hair on last segment not so broadened subapically, with margins
microscopically and sparsely serrate. Cuticular armature absent. Prothora-
cic pseudopod cleft apically, each lobe with 6-7 brown hooks; posterior
pseudopod with a row of 16-20 brown hooks; anal blood gills bilobed.

Distribution.—Brazil, Colombia, Dominica.

Types.—Holotype 2, allotype ¢, Dominica, Calabishie, swamp at mouth
of Hodges River, 27.11.1965, W.W. Wirth, reared from bracts of Calathea
lutea (Type no. 76582, USNM). Paratypes, 22 3, 24 2, as follows: DO-
MINICA: Same data as types, 8 ¢, 6 9, with associated larval and pupal
exuviae, 3 larvae. BRAZIL: Santa Catarina, Nova Teutonia, various dates
1961-1970, F. Plaumann, 14 6, 18 9.

Other material examined.—BRAZIL: Bahia, Ilheus, Itabuna, 24.x.1977,
S. J. Soria, reared from sugar cane leaf axils, | 6 with pupal exuviae, 4
larvae. COLOMBIA: Antioquia, San Jeronimo, 17.1.1974, E. Urueta, reared
from decaying papaya stem, 2 6, 3 2, | pupa, 2 larvae.

Discussion.—The species takes its name from the plant from which the
type-series was reared in Dominica. Forcipomyia calatheae is distinguished
from the other species in this group by rather negative characters: The
vestigial mandibular teeth; lack of hastate spines on the female tibiae; nar-
row rather than broad scales on the abdominal terga and legs; the uniformly
dark brown larval head; the large pupal respiratory horns with long, more |}
or less convoluted, line of spiracular openings; and the short dorsal tubercles
on the pupal cephalothorax.

Forcipomyia quatei Wirth
Fig. 2

Forcipomyia quatei Wirth, 1952: 142 (¢, 2; Calif.; fig. d genitalia).

A pale brown species with yellowish legs; wing dark brown with a small
yellow spot at end of costa; abdomen with conspicuous clumps of broad |
striated blackish scales on pleural segments; halter brown. |

Female.—Wing length 1.15 mm; breadth 0.48 mm.

Head: Brown including antenna and palpus. Antenna (Fig. 2a) with
lengths of flagellar segments in proportion of 15-11-11-12-12-11-11-11-24-25-
27-28-33; antennal ratio 1.45. Palpus (Fig. 2b) with lengths of segments in
proportion of 7-13-29-10-10; 3rd segment broad and evenly swollen on prox-
imal 42, abruptly tapering to slender tip past the sensory pit, palpal ratio
2.0; sensory pit 2X as deep as diameter of pore opening, which is slightly

VOLUME 84, NUMBER 3 577

I ea ere Pe a ee

Fig. 2. Forcipomyia quatei. a~e, Female. f—g, Male. h, k-q, Larva. i-j, Pupa. a, Antenna.
b, Palpus. c, Wing. d, Mandible. e, Spermathecae. f, Parameres. g, Genitalia, parameres re-
moved. h, Second abdominal segment, lateral view. i, Thoracic tubercles. j, Respiratory horn.
k, Head, lateral view. I-p, a, b, d, c, e, f hairs respectively, of 2nd abdominal segment. q,
Two posterior abdominal segments, dorsal view.

- than diameter of pit. Mandible (Fig. 2d) with 21 small teeth, their size
slightly increasing proximad in series.

Thorax: Brown; mesonotum and scutellum dark brown, with abundant
mixture of long bristly hairs and shorter, fine, somewhat flattened hairs.
Legs bright yellow, tarsi and narrow apex of hindfemur brownish; knees,

578 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

foretibia, and all of tarsi appearing shaggy due to abundant vestiture of
broad, striated brown scales; tibiae with long extensor bristles, as long as
4x diameter of tibia on hindleg. Hindtibial spur yellowish, inconspicuous,
length less than apical breadth of tibia; tarsomeres | and 2 with ventral row
of palisade setae, prominent only on foreleg; some long, rather slender ven-
tral spines apically and subapically on tarsomeres 1-4 on mid- and hindlegs,
poorly developed on foreleg. Tarsal ratios 0.54 on foreleg, 0.49 on midleg,
and 0.56 on hindleg. Claws slender and curved, empodium well developed.
Wing (Fig. 2c) brownish, with abundant brownish long decumbent macro-
trichia, more prominent along veins, a small yellowish spot formed by yel-
lowish scales at end of costa; veins in radial field appearing much darker
due to abundant flattened broad striated scales; costal ratio 0.44. Halter
brownish.

Abdomen: Brown; covered with broad flattened brown scales, pleura ap-
pearing blackish due to prominent clumps of such scales; cerci yellowish.
Spermathecae (Fig. 2e) 2, dark brown, ovoid, tapering gradually to slender
opening to duct; subequal, each measuring 0.081 by 0.032 mm.

Male.—Wing length 1.39 mm; breadth 0.45 mm. Similar to female with
usual sexual differences. General color as in female, but wing appearing |
slightly paler brown between veins due to sparser vestiture of longer nar-
rower macrotrichia; abdomen yellow with segmental banding of brown in- |
tegumental color plus abundant broad, flattened, dark brown, appressed
scales. Antennal segments 12-15 with lengths in proportion of 42-45-50-50;
3rd palpal segment not as broadly swollen on proximal %, palpal ratio 2.5;
costal ratio 0.44; tarsal ratios 0.45 on foreleg, 0.53 on midleg, and 0.55 on |
hindleg.

Genitalia (Fig. 2g): Short and broad; dark brown, yellowish on proximal
halves of basistyles, proximal halves of dististyles, and proximal portion of
9th segment; with abundant vestiture of long bristly yellowish hairs and
broad, flattened, moderately long, striated, dark brown scales. Aedeagus
shield-shaped as usual in the subgenus, slightly longer than basal breadth,
without basal arch, tip with a slender, pointed process, and with a pair of
faint, linear, submedian internal ribs. Parameres (Fig. 2f) with bases joined
in a broad basal portion extending to 4 of total length, the caudal processes
rather stout and swordlike with pointed apices. |

Pupa.—Length 2.7—2.8 mm. Color of exuviae pale yellowish. Respiratory |
horn (Fig. 2j) slender, with distinct petiole on proximal %2; apex rounded |
with 12-15 spiracular openings in a row at apex and down posterior side,
the anterior margin of distal portion carinate and meeting proximal end of
Ist spiracle as figured. Thoracic tubercles (Fig. 2i) elongated, stalklike, the
anterior 2 pairs each bearing a short slender seta, the other 3 pairs bearing
numerous spinules on distal halves. Abdomen without conspicuous lateral
processes.

VOLUME 84, NUMBER 3 579

Larva.—Length 4-5 mm when mature. Color yellowish white; head cap-
sule pale brownish, somewhat darker around oral margin; prominences at
bases of body hairs not strongly pigmented. Hairs p and q of head (Fig. 2k)
slightly bent forward, thickened and microscopically spiculose on distal
halves. Body hairs a-/ as in Figs. 2h and 2I—p. Last 2 body segments as in
Fig. 2q, the a hair of last segment hastate and only slightly longer than those
on preceding segments; anal blood gills short, bilobed. Cuticular armature

_absent. Prothoracic pseudopod cleft apically, each lobe with 7-8 slender
black hooks; posterior pseudopod with a row of 16-20 brown hooks.

Distribution.—USA (California, Louisiana, Florida), Dominica, Belize,
Costa Rica, Panama, Ecuador, Brazil.

Types.—Holotype d¢, Bakersfield, Kern Co., California, vii.1946, B.
Brookmam (Type no. 60929, USNM). Allotype and 2 paratypes, Kern Co.,

California.

_ Specimens examined.—All from light traps unless otherwise noted. BE-
-LIZE: Hummingbird Hershey, viii.1981, A. M. Young, from cacao pods,
| 10 36,9 2. BRAZIL: Bahia, Itabuna, iv.1971, J. Winder 1 ¢6; same, 1973,
reared from cacao pods, | 6; from emergence traps, 6 6, 6 2. COSTA
/RICA: Heredia Prov., La Virgen, Finca La Tigra, 19.ix.1979, A. Young, ex
banana stem, 4 larvae, 10 pupae. Limon Prov., Finca La Lola near Si-
‘quirres, ex banana stems, Young, 11.1981, 2 6, 4 2, 9 larvae, 8 pupae;
vili.1981, 3 ¢, 3 2, 2 larvae, 4 pupae. DOMINICA: Clarke Hall, 28.i.1965,
W. W. Wirth, reared from banana stem, 7 ¢, | 2, 4 larvae, 4 pupal exuviae
(specimens figured). ECUADOR: Pichilingue (INIAP), iv.1978, reared from
‘rotting log, J. Mendoza no. 13, 8 6, | 2. PANAMA: Canal Zone near
_Arraijan, monsoon forest, Spondias tree canopy, 23.vii.1979, E. Broadhead,
1 6. USA: Florida: Alachua Co., Gainesville, iv.1967, F. S. Blanton, |
36; 18.xi.1969, F. W. Mead, | ¢. Collier Co., Collier Seminole St. Park,
17.v.1973, Wirth, 1 6, 4 2. Escambia Co., Molina, 12.viii.1969, Blanton,
2 6; Walnut Hill, vi.1969, Blanton, | ¢. Jackson Co., Florida Caverns St.
Park, 26.v.1973, Wirth, 5 3. Jefferson Co., Monticello ix.1969, W. H. Whit-
eomb, 5 6, 4.2. Lee Co.,, Sanibel Island, 11.v.1973, Wirth; 16,3. 9. Leon
Co., 3 mi. N. Tallahassee, v.1970, Blanton, 2 2. Liberty Co., Torreya St.
Park, 15.v.1971, G. B. Fairchild, 1 6: Monroe Co., Big Pine Key,
10.iv.1970, Wirth, 13 6, 15 2. Orange Co., Lake Magnolia Park, 6.viii.1970,
= rons, 1 -o...Putnam Co.,, Lon’s, Lake, y.1971,,Blanton;,3.¢.,, 18.2 «Sar-
asota Co., Myakka River St. Park, 21.v.1973, Wirth, | 9. Louisiana: E.
Baton Rouge Parish, Baton Rouge, University Campus, v.1947, Wirth,

'

Forcipomyia (Forcipomyia) youngi Wirth, NEw SPECIES

A brown species with banded abdomen and pale yellow legs; wing in both
sexes dark brown with a small yellow spot at end of costa and a clump of

580 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

yellow scales at base of costa; abdomen with segmental bands of broad
striated blackish scales and pleura with additional mixture of golden scales;
tibia with broad brown scales; halter pale.

Holotype female.—Wing length 1.48 mm; breadth 0.60 mm.

Head (Fig. 3i): Brown, with abundant blackish, long, bristly setae on
vertex, clypeus, and first 2 antennal segments. Antenna (Fig. 3a) with seg-
ments 3-10 yellow, 11-15 pale brown; lengths of flagellar segments in pro-
portion of 16-12-13-13-13-13-13-13-33-33-33-32-44; antennal ratio 1.50. Pal-
pus (Fig. 3b) dark brown; lengths of segments in proportion of 10-14-30-14-
13; 3rd segment swollen on proximal %, bearing a deep sensory pit opening
by a slightly smaller pore; palpal ratio 1.9. Mandible with 11 coarse teeth,
the proximal teeth larger.

Thorax: Mesonotum, scutellum and postscutellum dark brown, pleuron
yellowish with some pale brown areas; mesonotum with long bristly hairs
and shorter finer setae; scutellum with numerous long bristly setae. Legs
bright yellow, tarsi pale brownish; tibiae with vestiture of short, broad,
appressed, striated brown scales; tarsi with similar scales more numerous
but more slender, scales at narrow apices of tarsomeres golden; tibiae with
numerous long yellowish extensor bristles, the longest on hindtibia as long
as 5x diameter of tibia; hindtibial spur yellowish, slender, pointed, about
as long as apical breadth of tibia. Tarsi with ventral palisade setae in one
row, prominent and spinelike on tarsomeres | and 2 of foreleg, poorly de-
veloped on mid- and hindlegs; some stronger dark ventral spines scattered
on tarsomeres | and 2 of mid- and hindlegs, apical only on tarsomeres | and
2 of foreleg and on tarsomeres 3 and 4 of all legs. Tarsal ratios 0.58 on
foreleg, 0.52 on midleg, and 0.54 on hindleg. Wing (Fig. 3c) brown, darker
anteriorly and along veins due to denser vestiture of long black macrotrichia;
a small yellow spot at end of costa and a clump of yellow scales on costa
at wing base; long, narrow, dark scales on costa and over radial field shaped
as in Fig. 3g; costal ratio 0.48. Halter pale.

Abdomen: Dark brown, last 2 segments and cerci yellowish; densely cov-
ered dorsally and ventrally with moderately long and broad, flattened, striat-
ed, dark brown scales; pleura with similar scales but golden yellow in color;
last tergum with long bristly yellow hairs. Spermathecae (Fig. 3h) 2, dark
brown, elongate ovoid, tapering gradually to slender opening to duct; sub-
equal, each measuring 0.104 by 0.054 mm.

Male allotype.—Wing length 1.44 mm; breadth 0.50 mm. Similar to female
with usual sexual differences. General color as in female, but wing with
costa yellowish and bearing yellow scales on proximal 42; abdomen yellow
below and brown above, each tergum with a line of brown infuscation fol-
lowing the bases of the long bristly hairs across midlength of tergum. An-
tennal segments 12-15 with lengths in proportion of 63-48-40-47; 3rd palpal

VOLUME 84, NUMBER 3 581

_ a eam ey aenlat ams Wr GD GO Gp t=

‘a
Ct ge
7h b

TRETES

Fig. 3. Forcipomyia youngi. a—d, g-i, Female. e-f, Male. j-n, r-t, Larva. o—q, Pupa. a,
‘Antenna. b, Palpus. c, Wing. d, Mandible. e, Genitalia, parameres removed. f, Parameres. g,
Scalelike macrotrichia from wing. h, Spermathecae. i, Head, anterior view. j-n, a, b, d, c, e,
f hairs, respectively, of 2nd abdominal segment. 0, Lateral processes of abdomen. p, Respi-
ratory horn. q, Thoracic tubercles. r, Head, lateral view. s, Second abdominal segment, lateral
view. t, Two posterior abdominal segments, dorsal view.

582 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

segment not as swollen as in female, palpal ratio 2.3. Costal ratio 0.45; tarsal
ratios 0.57 on foreleg, 0.47 on midleg, and 0.56 on hindleg.

Genitalia (Fig. 3e): Slightly longer than broad; moderately brown infus-
cated except at bases of 9th segment, basistyles, and dististyles. Aedeagus
almost hyaline, with rather narrow base, about 1.5x as long as basal
breadth, tip with slender process. Parameres (Fig. 3f) poorly sclerotized,
yellowish, basal portion fused for about % of total length, the caudomedian
emargination narrow and pointed, distal processes rather stout and sword-
like with pointed apices.

Pupa.—Length 2.8-3.0 mm. Color pale brownish. Respiratory horn (Fig.
3p) short and globular, with short slender petiole; swollen distal portion
with a row of 20-25 spiracular openings around apex and a slightly undu-
lating row down posterior side of swollen portion. Thoracic tubercles (Fig.
3q) greatly elongated and slender, the anterior 2 pairs each bearing a long
slender seta about as long as tubercle itself, the posterior 3 pairs with apices
finely and densely spinulose. Each abdominal segment (Fig. 30) with a pair
of elongate lateral processes, each process with spiculose tip and a subapical
seta.

Larva.—Length 5-6 mm when mature. Color yellowish white; head cap-
sule (Fig. 3r) brown, becoming conspicuously blackish toward oral margin; |
conical prominences at bases of dorsal body hairs moderately brownish
pigmented; a hairs connected across dorsum of each body segment by a
conspicuous narrow line of blackish pigment. Hairs p and q of head (Fig.
3r) long and slender, pale, of uniform width to tip, and sparsely fringed with
microscopic spinules. Body hairs a, b, and e hyaline, c, d, and f dark brown;
hairs b, c, and d with fringe of short, coarse spinules as figured. Last 2 |
body segments as in Fig. 3t, the 4 a and b hairs on last segment arising from
a common sclerotized plate, the a hairs long and slender, only slightly flat-
tened and expanded, and bearing fringing spinules; anal blood gills com-
paratively slender, apparently undivided. Cuticular armature absent. Pro-
thoracic pseudopod short and only slightly cleft, each lobe with 6-8 blackish
hooks; posterior pseudopod with a row of 18-20 blackish hooks.

Distribution.—Costa Rica, Ecuador, Panama.

Types.—Holotype 2, allotype 6 , Panama, Canal Zone, Gamboa, Pipeline
Road, vii.1967, W. and M. Wirth, reared from rotting bracts of Heliconia |
mariae (Type no. 76583, USNM). Paratypes, 29 3, 28 2, 20 larvae, 24
pupae, as follows: COSTA RICA: Limon Prov., Finca La Lola, 11.1981, 7 |
3,10 2,6 larvae, 10 pupae; same, but viii.81, 3 ¢, 6 2, 3 larvae, 3 pupae;
same, but 9-11.xi.1980, ex rotten cacao pods, | 6. Siquirres, 29.v.1956, L.
G. Saunders, reared from cacao pods and tree bark, | 6, 2 2, 3 larvae; |
same but 6.vi.1956, reared from cacao pods, 5 6,5 ¢@, 3 larvae. Cartago
Prov., Turrialba, CATIE, 24.vi.1980, A. Young, | ¢ and pupal exuviae.

VOLUME 84, NUMBER 3 583

Heredia Prov., La Virgen, Finca El Uno, 19.ix.1979, A. Young, ex banana
stem, | larva, 6 pupae. ECUADOR: Quevado, Pichilingue, INIAP, iv.1978,
J. Mendoza, reared from rotting vegetation, 8 ¢, 1 2. PANAMA: Same
data as types, 3 6, 4 2, 4 larvae, 5 pupae.

Discussion.—This species is named for Dr. Allen Young, Milwaukee Pub-
lic Museum, in appreciation of his interest and his studies on the biology of
Costa Rican ceratopogonid cacao pollinators.

| Forcipomyia youngi is easily recognized by its enlarged female mandib-
ular teeth, and by the very broad, short, flattened, blackish scales on the
tibiae. The larva has the head pale posteriorly and dark brown towards the
mouth, and the pupa is distinguished by the elongate thoracic tubercles with
long apical setae.

Because of its close external similarity with species of the argenteola
Group, the following diagnosis and discussion are presented for Forcipo-

myia squamitibia Lutz.

| Forcipomyia squamitibia Lutz

Forcipomyia squamitibia Lutz, 1914: 88 (2; Brazil; fig. wing, leg); Macfie,
1939: 145 (2 redescribed; Brazil; fig. antenna, palpus, tibial spine; radial

cells, spermatheca); Macfie, 1949: 111 (¢ described; Mexico; fig. para-

meres).

Female (measurements from a female from Nova Teutonia, Brazil):—
Wing length 1.30 mm; breadth 0.80 mm. A brown species with yellowish
legs; antenna pale brown; last 2 abdominal segments and cerci yellowish.
Wing dark brown, darker along anterior margin and along veins; a rather
large yellow spot covering most of 2nd radial cell and slightly distad on
mere wing margin; costal ratio 0.44. Halter slightly infuscated. Antennal
segments short and vasiform, antennal ratio 0.80. Palpus with 3rd segment
‘moderately swollen on proximal 2, with a small deep sensory pit; palpal
ratio 2.2. Mandibular teeth vestigial. Body, legs, and abdomen without
broad flattened scales, long narrow scales on abdominal terga. Tibiae with
slender, pointed hastate spines in extensor series. Tarsal ratios 1.05 on
foreleg, 1.06 on midleg, and 1.08 on hindleg. Spermathecae 2, oval to ovoid,
slightly tapering to short slender neck; slightly unequal, measuring 0.101 by
0.058 mm and 0.087 by 0.057 mm.

Male.—Similar to female with usual sexual differences. Tibiae without
hastate spines. Wing yellowish, anterior margin with large yellow spot over
tip of 2nd radial cell, preceded by a strip of dark brown scalelike macrotri-
chia over costal and radial veins and followed by a prominent quadrate
/patch of dark brown scales; a dark brown line along vein Cu,. Abdomen
with prominent brown segmental bands giving rise to long golden to brown-
ish, bristly hairs; no broad flattened scales. Genitalia with 9th segment

584 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

brown, basistyles and dististyles yellowish; aedeagus shield-shaped, slightly
longer than basal breadth, caudal apex pointed; parameres a pair of long
processes, rodlike at bases and tapering to filamentous tips, their bases
fused only a short distance in a rather narrow common base (as figured by
Macfie, 1949).

Distribution.—Brazil to Mexico.

Types.—An unspecified number of syntypes, Manguinhos, Brazil, Lutz
collection, common at lights with other marine species (Inst. Oswaldo Cruz,
Rio de Janeiro).

Specimens examined.—BRAZIL: Santa Catarina, Nova Teutonia, var-
ious dates 1961-1970, F. Plaumann, 40 6, 45 2 (USNM). COSTA RICA:
Limon Prov., Finca La Lola near Siquirres, 11.1981, A. M. Young, reared
ex banana stems, 2 6, 2 2, 2 larvae, 2 pupae; same but vili.1981, 1 pupa.

Discussion.—This species is set apart and readily distinguished from the
other species discussed in this paper by the following characters: The yellow
wing spot is much larger, covering the distal portion of the second radial
cell; the female tibiae bear an extensor series of prominent pale brownish,
slender, pointed, hastate spines; the hindtarsal ratio is 1.0 or slightly more;
and the male parameres are much more slender, filiform distally, and have ©
their common base much narrower and fused only a short distance.

The immature stages of the series from Costa Rica will be described later
in a paper dealing with other groups of Forcipomyia (Forcipomyia).

LITERATURE CITED

Bystrak, P. G. and W. W. Wirth. 1978. The North American species of Forcipomyia, subgenus |
Euprojoannisia (Diptera: Ceratopogonidae). U.S. Dep. Agric. Tech. Bull. 1591, 51 pp.

Chan, K. L. and E. J. LeRoux. 1965. Description of Forcipomyia (Neoforcipomyia) sp. n.
and redescription of Forcipomyia (Neoforcipomyia) eques (Johannsen) (Diptera: Cera-
topogonidae), with an account of the digestive and reproductive systems. Phytoprotec-
tion 46: 74-104.

Lane, J. 1947. A biologia e taxonomia de algunas espécies dos grupos Forcipomyia e Culi-
coides (Diptera, Ceratopogonidae) (Heleidae). Arg. Fac. Hig. Saude Publica Univ. Sao
Paulo 1: 159-170. |

Lutz, A. 1914. Contribuigao para o conhecimento das Ceratopogoninas do Brazil. Terceira
memoria. Aditamento terceiro e descrigao de espécies que nao sugam sangue. Mem.
Inst. Oswaldo Cruz, Rio de J. 6: 81-99, 2 plates.

Macfie, J. W. S. 1939. A report on a collection of Brazilian Ceratopogonidae (Dipt.). Rev. |

Entomol. 10: 137-219.

. 1949. Notes on Ceratopogonidae. Proc. R. Entomol. Soc. Lond. (B) 18: 109-115.

Saunders, L. G. 1924. On the life history and the anatomy of the early stages of Forcipomyia
(Diptera, Nemat., Ceratopogoninae). Parasitology 16: 164-213, 3 pls.

—. 1956. Revision of the genus Forcipomyia based on characters of all stages (Diptera,
Ceratopogonidae). Can. J. Zool. 34: 657-705.

—. 1959. Methods for studying Forcipomyia midges, with special reference to cacao-
pollinating species (Diptera, Ceratopogonidae). Can. J. Zool. 37: 33-S1.

VOLUME 84, NUMBER 3 58

n

Soria, S. J. de and P. G. Bystrak. 1975. A new species of Forcipomyia (Diptera, Ceratopo-
gonidae) described in all stages, with an account of its role as a cacao pollinator. Rev.
Theobroma 5(2): 3-11.

Wirth, W. W. 1952. The Heleidae of California. Univ. Calif. Publ. Entomol. 9: 95-266.

. 1976. Forcipomyia pictoni Macfie and descriptions of two new related species from

Florida (Diptera: Ceratopogonidae). Fla. Entomol. 59: 77-84.

Wirth, W. W. and S. J. de Soria. 1975. A new Neotropical Forcipomyia midge closely related

to F. (F.) genualis (Loew) (Diptera: Ceratopogonidae). Rev. Theobroma 5(2): 19-27.

. 1980(1979). Studies on the genus Forcipomyia VI. The Neotropical species of the

subgenus Warmkea (Diptera: Ceratopogonidae). Rev. Theobroma 9: 137-161.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 586-593

ANTENNAL SENSILLA AND SETAE OF ANOPLIUS TENEBROSUS
(CRESSON) (HYMENOPTERA: POMPILIDAE)

STEVEN R. ALM AND FRANK E. KURCZEWSKI

(SRA) New York State Agricultural Experiment Station, P.O. Box 727,
Hudson Valley Laboratory, Highland, New York 12528; (FEK) Department
of Environmental and Forest Biology, SUNY College of Environmental
Science and Forestry, Syracuse, New York 13210.

Abstract.—The antennal sensilla and setae of both sexes of Anoplius te-
nebrosus (Cresson) (Pompilidae) were examined with a scanning electron
microscope. Sensilla placodea, corrugated conical sensilla, pit organs, sen-
silla campaniformia, sensilla trichodea Al, BI, Cl, sensilla basiconica, and
several types of setae were found, and are described and illustrated. The
sensilla are rather similar in both sexes, except that females have numerous
corrugated conical sensilla which males lack. Sensilla trichodea Cl are |
found only in the males. While most of the setae are similar in the two
sexes, the shape and sculpturing of some distinguish males from females.

Scanning electron microscopy studies of Hymenoptera to date have pri-
marily involved parasitoids, ants, social wasps, and bees. No such studies
have been reported for the Pompilidae, or spider wasps. In the following
examination of Anoplius tenebrosus antennae, the external morphology of |
various sensilla are illustrated and described. Morphological similarities of |
A. tenebrosus sensilla to those of other Hymenoptera are noted, and the
possibilities of functional analogies are discussed.

MATERIALS AND METHODS

Air dried antennae of male and female A. tenebrosus were mounted on
specimen studs with silver conducting paint. Specimens were coated in
vacuo with a gold/palladium mixture. The microscope used was an ETEC |
autoscan, model RI, at accelerating voltages of 10-20 KV. |

The nomenclature follows that of Agren (1977), who combined the clas-
sifications of Lacher (1964) and Esslen and Kaissling (1976), except for the
corrugated conical sensillum and sensilla trichodea designated Al, B1, Cl
for Pompilidae. Flagellar segments are designated fl.1, 2, etc. from proximal
to distal.

VOLUME 84, NUMBER 3 587

Figs. 1-5. Anoplius tenebrosus, female antenna. 1, Underside of antenna showing patches
| of sensilla trichodea; numerous openings on outer edge of antenna are corrugated conical
sensilla. 2, Enlargement of flagellar segment 5. 3, Sensilla trichodea, flagellar segment 10. 4, -
Enlargement of flagellar segment 4. 5, Enlargement of flagellar segment 7. Al = sensilla tri-
chodea; B! = slender sensilla trichodea; ccs = corrugated conical sensilla; P = pit organ; sb =
sensilla basiconica; sp = sensilla placodea; SS = sabre-shaped seta.

| RESULTS

Specimens of Pompilidae have filiform antennae. Females of A. tenebro-
sus have 12 segments, males 13 including the scape and pedicel. The sensilla
and setae are as follows:

Sensilla placodea (Figs. 2, 5, 10, 11, 16, 17, 20).—These elongate sensilla
are found on fi.2—11 in males and are distributed around the entire flagellum.
\In females they are found on fi.1-10 and are dispersed in an area encom-
passing slightly more than half of each flagellum. Females possess a region
-of corrugated conical sensilla in which s. placodea are not found. Sensilla
fplacodea consist of pits averaging 3.5 x 18.4 uw (females), 3.7 x 18.3 uw
(males) which contain elongate sensilla.

Corrugated conical sensilla (Figs. 1, 2, 5, 6, 9).—These sensilla are found
only in females on fl.1-10. They are one of the more numerous types of

588 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 6-11. Anoplius tenebrosus, female antenna. 6, Corrugated conical sensilla, flagellar |
segment 2. 7, Enlargement of flagellar segment |. 8, Long setae on scape. 9, Enlargement of |
flagellar segment 2. 10, Enlargement of flagellar segment 6. 11, Enlargement of flagellar segment

2. Bl = sensilla trichodea; ccs = corrugated conical sensilla; P = pit organ (Fig. 9, probably
s. ampullacea; Fig. 10, probably s. coeloconica); sb = sensilla basiconica; sp = sensilla pla- |
codea; S2 = long seta; S4 = seta.

sensilla A. tenebrosus females possess (see Fig. 1). The oval pit surrounding
the sensillum is approximately 7.6 x 9.6 w and contains a conical shaped
peg which is corrugated and has the apex recessed.

Pit organs (Figs. 5, 9-11, 16, 18, 21).—Pit organ openings of A. tenebrosus
range from 1.1 to 5.7 w. Some pit organs contain pegs that are not exposed
above the rim around the opening.

Sensilla campaniformia (Figs. 12, 20).—Sensilla campaniformia are found
on fl.1, 3-6, and 9 in males and I-10 in females. They consist of a slightly

VOLUME 84, NUMBER 3 589

Figs. 12-16. Anoplius tenebrosus, antenna. 12-13, Female. 14-16, Male. 12, Enlargement
of flagellar segment |. 13, Enlargement of scape, 14, Flagellar segments 5-8 showing differential
setae on underside of antenna (arrow); region composed of stout setae, sensilla trichodea, and
sensilla placodea. 15, Enlargement of flagellar segment 7. 16, Enlargement of flagellar segment
2. Al = sensilla trichodea; B1 = sensilla trichodea; P = pit organs; sc = sensilla campanifor-
mia; sp = sensilla placodea; S| = smooth setae; S7 = stout setae.

depressed oval region ca. 6.3 x 9.6 uw (females) and 5.5 x 6.9 « (males)
with a centrally located node.

Sensilla trichodea Al (Figs. 1-3, 15, 17).—These receptors are found on
the undersides of fi. 1—10 in females and I-11 in males. They occur in patches
in females; in males they are more numerous but are not found in patches.
This sensillum is smooth, sabre-shaped and appears to have a terminal pa-
pilla.

Sensilla trichodea B1 (Figs. 5, 7, 16).—Sensilla trichodea B1 are found on
all flagellar segments and pedicels of both males and females. They are
slender structures that lay close and nearly parallel to the cuticle.

Sensilla trichodea Cl (Figs. 17, 19).—These sensilla are found mainly on
the undersides of fl. 1-11 in males. They are deeply ridged and are nearly
perpendicular to the flagellar segment.

590 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Sensilla basiconica (Figs. 4, 5, 9, 18).—These receptors are found in both
sexes on all flagella. They have blunt tips, are found in pits and are relatively
common (see Fig. 5). They are ridged but not nearly as much as s. trichodea
Cl. There are two sizes of this sensillum (see Fig. 4).

Setae S1-S7 (Figs. 4, 5, 7, 8, 13, 15, 17, 18, 21, 22).—These appear to be
modifications of one basic type and the sculpturing and thickness vary con-
siderably from smooth on the scape (S1) to stout with deep sculpturing (S7,
males only) on the flagellum.

DISCUSSION

Although the functions of the various sensilla in Pompilidae and other
morphologically related Hymenoptera are mostly unknown, structural sim-
ilarities imply physiological similarities. Sensilla placodea of A. tenebrosus
are similar to those found in the Vespidae (Spradbery, 1973; Callahan, 1975),
Braconidae (Norton and Vinson, 1974a, 1974b; Borden et al., 1978b), and
Ichneumonidae (Borden et al., 1973, 1978a; Norton and Vinson, 1974b).
Oval s. placodea of Apis mellifera L. have been shown to be odor receptors
(Lacher and Schneider, 1963 [cited by Agren, 1977]). Slifer (1970) suggested
that the s. placodea of Nasonia vitripennis (Walker) (Pteromalidae) have an
olfactory function. Richerson et al. (1972) and Richerson and Borden (1972)
postulated that the s. placodea of Coeloides brunneri Viereck (Braconidae)
function in host finding by detection of metabolic heat produced by a host
beetle larva beneath bark.

Corrugated conical sensilla are found only in females of A. tenebrosus.
The apex of this sensillum was found to be recessed in air dried specimens.
Spradbery (1973) called similar structures on Vespidae, s. campaniformia;
however, it seems unlikely that so many s. campaniformia would be found
on the antenna. Callahan (1975) named a similar sensillum of Polistes me-
tricus Say (Vespidae) a corrugated pyramidal sensillum, and his nomencla-
ture has been modified for the conical shaped peg of A. tenebrosus. Kais-
sling (personal communication) stated that this receptor is reminiscent of a
taste bristle, s. chaeticum. Ulla Klein, working in Kaissling’s laboratory,
found similar structures to be taste bristles in crickets.

Pit organs are thought to function as CO,, temperature, and/or humidity
receptors in A. mellifera (Lacher, 1964 [cited by Agren, 1977)).. Int Agape=
nebrosus they may function similarly. These organs can be separated into
Ss. ampullacea or s. coeloconica by the structure of the sensory peg within
the pit (Dietz and Humphreys, 1971). Scanning electron microscopy of A.
tenebrosus antennae does not reveal enough of the peg to differentiate be-
tween these organs.

Sensilla campaniformia are probably mechanoreceptors in A. mellifera
(Esslen and Kaissling, 1976 [cited by Agren, 1977]). Dietz and Humphreys

VOLUME 84, NUMBER 3 591

Figs. 17-22. Anoplius tenebrosus, male antenna. 17, Enlargement of flagellar segment 2.
18, Enlargement of flagellar segment 8. 19, Enlargement of flagellar segment 3. 20, Enlargement
of flagellar segment 9. 21, Enlargement of flagellar segment 9. 22, Enlargement of pedicel.
Al = sensilla trichodea; Cl = sensilla trichodea; P = pit organ (probably sensilla coeloconica
in Fig. 21); sb = sensilla basiconica; sc = sensilla campaniformia; sp = sensilla placodea;
S3 = seta; S6 = seta; S7 = stout seta.

(1971) suggested that mechanical depression of the node of s. campaniformia
in A. mellifera may result in a neural response but added that the numerous
hairs on the antennae would probably prevent that mode of stimulation from
taking place. S. campaniformia have been found in several species of bees,
but not in all wasps.

Sensilla trichodea Al of A. tenebrosus bear resemblance to the small
trichoid of Cheiloneurus noxius Compere (Encyrtidae) (Weseloh, 1972),
curved non-fluted sensilla of Cardiochiles nigriceps Viereck (Braconidae)
(Norton and Vinson, 1974a), s. chaetica, in A. mellifera (which also has a
terminal papilla) (Whitehead and Larsen, 1976), and the s. trichodea A of

592 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Colletidae (Agren, 1977) and Andrenidae (Agren, 1978). The morphology of
a similar sensillum studied by Slifer and Sekhon (1961) in A. mellifera in-
dicated an olfactory function.

Sensilla trichodea BI of A. tenebrosus resemble the s. trichodea B2 of A.
mellifera (Esslen and Kaissling, 1976 [cited by Agren, 1977]) and the s.
trichodea B of Colletidae (Agren, 1977) and Andrenidae (Agren, 1978).
Lacher (1964 [cited by Agren, 1977]) indicated they are responsive to me-
chanical stimuli in A. mellifera.

Sensilla trichodea Cl of A. tenebrosus look like the fluted basiconic sen-
silla of Cardiochiles nigriceps (Norton and Vinson, 1974a). However, the
tip is more pointed and the sculpturing more pronounced. Norton and Vin-
son (1974b) suggested that the fluted basiconic sensilla of three parasitoid
species are contact receptors.

Sensilla basiconica are known to be olfactory receptors in many insects
(Schneider and Steinbrecht, 1968), although attempts to determine a func-
tion for these sensilla in A. mellifera have been unsuccessful.

The nomenclature applied to various hymenopteran sensilla is unstable
at the present time. Borden et al. (1973) described a ‘“‘new”’ sensillum (trun-
cate cuticular peg) on Itoplectis conquisitor (Say) (Ichneumonidae) which
resembles the small basiconicum b of Cheiloneurus noxius (Weseloh, 1972).
The small, subterminal basiconicum of C. noxius may be homologous to the
basiconic capitate peg of Nasonia vitripennis (Miller, 1972). Other similar-
ities among sensilla of various groups have been discussed. Only from
knowledge of the neuro- and electrophysiology of each sensillum will this
instability be resolved.

ACKNOWLEDGMENTS

We thank G. N. Lanier and R. A. Norton, SUNY College of Environ-
mental Science and Forestry, for reviewing the manuscript; and H. E. Ev-
ans, Colorado State University, for identifying Anoplius tenebrosus. W. A.
Cote and A. C. Day instructed in the use of the scanning electron micro-
scope. Karl-Ernst Kaissling, Max Planck Institut fir Verhaltenphysiologie,
J. H. Borden, Simon Fraser University, and P. S. Callahan, USDA, Gaines-
ville, Florida, aided in identifying sensilla.

LITERATURE CITED

Agren, L. 1977. Flagellar sensilla of some Colletidae (Hymenoptera: Apoidea). Int. J. Insect
Morphol. Embryol. 6: 137-146.

——. 1978. Flagellar sensilla of two species of Andrena (Hymenoptera: Andrenidae). Int.
J. Insect Morphol. Embryol. 7: 73-79.

Borden, J. H., G. E. Miller, and J. V. Richerson. 1973. A possible new sensillum on the
antennae of Itoplectis conquisitor (Hymenoptera: Ichneumonidae). Can. Entomol. 105:
1363-1367.

VOLUME 84, NUMBER 3 593

Borden, J. H., L. Chong, and A. Rose. 1978a. Morphology of the elongate placoid sensillum
on the antennae of Jtoplectis conquisitor. Ann. Entomol. Soc. Am. 71: 223-227.
Borden, J. H., A. Rose, and R. J. Chorney. 1978b. Morphology of the elongate sensillum
placodeum on the antennae of Aphidius smithi (Hymenoptera: Aphidiidae). Can. J.

Zool. 56: 519-525.

Callahan, P. S. 1975. Insect antennae with special reference to the mechanism of scent detec-
tion and the evolution of sensilla. Int. J. Insect Morph. Embryol. 4: 381-430.

Dietz, A. and W. J. Humphreys. 1971. Scanning electron microscopic studies of antennal
receptors of the worker honey bee, including sensilla campaniformia. Ann. Entomol.
Soc. Am. 64: 919-925.

Esslen, J. and K.-E. Kaissling. 1976. Zahl und verteilung antennaler sensillen bei der honig-
biene (Apis mellifera L.). Zoomorphologie (n.s.) 83: 227-251.

Lacher, V. 1964. Elektrophysiologische unterschungen an einzelnen rezeptoren fiir geruch,
kohlendioxyd, luftfeuchtigkeit und temperatur auf den antennen der arbeitsbiene und
der drohne (Apis mellifica L.). Z. VGl. Physiol. (n.s.) 48: 587-623.

Lacher, V. and D. Schneider. 1963. Elektrophysiologischer nachweis der riechfunktion von
porenplatten (sensilla placodea) auf den antennen der drohne und der arbeitsbiene (Apis
mellifica L.). Z. VGI. Physiol. (n.s.) 47: 274-278.

Miller, M. C. 1972. Scanning electron microscopic studies of the flagellar sense receptors of
Peridesmia discus and Nasonia vitripennis (Hymenoptera: Pteromalidae). Ann. Ento-
mol. Soc. Am. 65: 1119-1124.

Norton, W. N. and S. B. Vinson. 1974a. A comparative ultrastructural and behavioral study
of the antennal sensory sensilla of the parasitoid Cardiochiles nigriceps (Hymenoptera:
Braconidae). J. Morphol. 142: 329-350.

—. 1974b. Antennal sensilla of three parasitic Hymenoptera. Int. J. Insect Morphol. Em-
bryol. 3: 305-316.

Richerson, J. V. and J. H. Borden. 1972. Host finding by heat perception in Coeloides brunneri
(Hymenoptera: Braconidae). Can. Entomol. 104: 1877-1881.

Richerson, J. V., J. H. Borden, and J. Hollingdale. 1972. Morphology of a unique sensillum
placodeum on the antennae of Coeloides brunneri (Hymenoptera: Braconidae). Can. J.
Zool. 50: 909-913.

Schneider, D. and R. A. Steinbrecht. 1968. Checklist of insect olfactory sensilla. Symp. Zool.
Soc. Lond. No. 23, pp. 279-297.

Slifer, E. H. 1970. The structure of arthropod chemoreceptors. Annu. Rev. Entomol. 15: 121-
142.

Slifer, E. H. and S. S. Sekhon. 1961. Fine structure of the sense organs on the antennal
flagellum of the honeybee, Apis mellifera L. J. Morphol. 109: 351-381.

Spradbery, J. P. 1973. Wasps—an account of the biology and natural history of solitary and
social wasps. Sidgwick and Jackson, London. 408 pp.

Weseloh, R. M. 1972. Sense organs of the hyperparasite Cheiloneurus noxius (Hymenoptera:
Encyrtidae) important in host selection processes. Ann. Entomol. Soc. Am. 65: 41-46.

Whitehead, A. T. and J. L. Larsen. 1976. Ultrastructure of the contact chemoreceptors of
Apis mellifera L. (Hymenoptera: Apidae). Int. J. Insect Morphol. Embryol. 5: 301-315.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 594-602

NOTES ON GRETCHENA: A NEW SPECIES AND THE SYNONYMY
OF GWENDOLINA (LEPIDOPTERA: TORTRICIDAE)

RICHARD L. BROWN

Department of Entomology, Drawer EM, Mississippi State, Mississippi
39762.

Abstract.—Morphological characteristics of Gretchena are discussed and
compared with those of the related Pseudexentera. Gretchena obsoletana,
a new species occurring in California, is described and compared with Epi-
notia biangulana (Walsingham), with which it has previously been confused.
Gwendolina is synonymized with Gretchena.

Gretchena was described by Heinrich (1923) to include eight species of
Nearctic Eucosmini that were characterized chiefly by distinctive male gen-
italia. References to Gretchena have been scant since 1923 but include the
description of an additional species by McDunnough (1925) and the occa-
sional mention of the pecan bud moth, G. bolliana (Slingerland) (Payne et
al., 1979). The larvae of G. bolliana and G. dulciana Heinrich have been
described by MacKay (1959). Tedders and Osburn (1970) have described
the morphology of the female reproductive system of G. bolliana.

Adults of Gretchena have a gray to grayish-brown ground color. In most
species, the basal patch is distinct and is usually more distinct on the inner
margin than on the costa of the forewing. Median and apical streaks are
present on the forewing of most species and are often confluent with each
other. Gretchena and the related Pseudexentera Heinrich are superficially
similar; some species are often mixed in collections. Heinrich distinguished
Gretchena from the latter by the presence of raised scales on the forewing
and by characters of the genitalia. As the raised scales are often difficult to
detect, characters of the genitalia should be used to separate the two genera.

In males of Gretchena, the uncus is absent; the elongately triangular socii
are separate and are joined with the tegumen dorsally; the aedeagus is long
and is supported by a long caulis of the juxta; and the cucullus of the valva
has a projecting anal area with developed anal and ventral marginal spines.
In Pseudexentera, the males have socii that are fused basally and anal and
ventral marginal spines of the valvae are absent. The valvae are more setose

VOLUME 84, NUMBER 3

n
\o
an

Fig. 1. Pseudexentera spoliana, 2, 7th sternum and lamella postvaginalis; Ithaca, N.Y.;
R. L. Brown genitalia slide 252. Scale line = | mm.

in Pseudexentera than in Gretchena. Male genitalia of the two genera have
been figured by Heinrich (1923).

The females of Gretchena have a seventh sternum that is not emarginate
around the ostium bursa; the posterior margin of the sternum is inflected
greatly; the lamella postvaginalis is reduced; the ductus bursa has a long,
sclerotized band; and the globular corpus bursa has two signa. The females
of Pseudexentera are distinct from Gretchena in having a seventh sternum
that is deeply emarginate around the ostium and a well-developed lamella
postvaginalis (Fig. 1). Females of these two genera can be separated easily,
without dissections, by the scale patterns on the seventh sternum that in-
dicate the presence or absence of an emargination.

Of the eight species included by Heinrich in Gretchena, seven occur in
eastern United States and one occurs in California. The California species
was misidentified by Heinrich as Steganoptycha biangulana Walsingham.
Examinations of type-specimens of Eucosmini in the British Museum (Nat-
ural History) have revealed that Walsingham’s species, described from Or-
egon, is a polymorphic species belonging to Epinotia Hubner. Although

596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Gretchena obsoletana, paratype ¢; Mt. Washington District, Los Angeles, Calif.;
approximately 4x natural size.

Heinrich figured the California Gretchena species and differentiated it from
related species, the following description is given to make a name available.

Gretchena obsoletana Brown, NEW SPECIES
Figs. 24

Description.—Head: Frons and labial palpus dark grayish brown, vertex
dark grayish brown, tips of scales white.

Thorax: Mesonotum dark grayish brown to brown, tegula dark grayish
brown basally, grayish brown to brown apically.

Forewing (Fig. 2): 8.5—9.0 mm long; ¢ costal fold absent; termen weakly
convex; ground color grayish brown, most scales with white tips; basal
patch dark grayish brown, basal inner margin suffused with light grayish
brown in some specimens; median fascia weakly defined in some specimens,
dark grayish brown; outer marginal scales unicolorous with or slightly dark-
er than ground color.

Hindwing and abdomen: Light grayish brown, without contrasting colors
dorsally or ventrally.

Male genitalia (Fig. 3): Tegumen narrow, expanded below socii; socius
with dorsal margin irregularly serrate, apex directed dorsally, setose from
near base to apex; gnathos arising from ventral bases of socii, lightly scler-
otized; caulis without lateral flanges; anellus not closely surrounding aedea-
gus ventrally; aedeagus with 20—22 cornuti; cucullus with 1-5 ventral mar-
ginal spines. Six preparations examined.

Female genitalia (Fig. 4): Sternum VII with posterior margin inflected
inward, posterior lateral areas sclerotized, not in same plane as posterior
median area. Tergum VIII without lateral extensions, without scales or se-

VOLUME 84, NUMBER 3 597

Fig. 3. Gretchena obsoletana, paratype, 3 genitalia; Los Angeles, Calif.; USNM genitalia
slide 17785. Scale line = 0.5 mm.

tae. Papillae anales with shallow, posterior cleft, setae moderately sparse,
numbering less than 7 across middle of one ventral face; sterigma forming
ringlike antrum cephalad to posterior margin of sternum VII; poststerigmal
membrane covered with small papillae between ostium and papillae anales;
inception of ductus seminalis cephalad to sclerotized band of ductus bursae;
signa subequal in size, equidistant from neck. Three preparations examined.
Holotype.—?, Los Angeles, California, L. A. Co., elev. 840 ft., Mt.
Washington Dist., HI-30-1976; at light, J. Powell, Collector; genitalia slide
970, R. L. Brown; in California Academy of Sciences, San Francisco, on
indefinite loan from University of California, Berkeley.
Paratypes.—California: Same data as holotype, 6 ¢, JAP genitalia 4083,
in University of California, Berkeley; same data as holotype, | 6, R. L.
Brown genitalia slide 969, R. L. Brown photo 157, in R. L. Brown Collec-
tion; Ventura Co., N end of Casitas Res., III-16-67, Collectors P. A. Opler,
J. Powell, 1 6, JAP genitalia 2225, in University of California, Berkeley; |
~9,in R. L. Brown Collection; Saticoy, 3-6-17, S. E. Flanders, Collector,
1 3, AB genitalia slide Mar. 20, 1927, in the National Museum of Natural
History, Smithsonian Institution. The following 7 specimens are labeled as
collected by Coquillett in California and probably represent one lot; all have

598 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Gretchena obsoletana, paratype, ? genitalia, unmated; Los Angeles, Calif.; USNM
genitalia slide 17783. Scale line = | mm.

VOLUME 84, NUMBER 3 599

Figs. 5-7. Imagos, approximately 4x natural size. 5, Epinotia biangulana, 3; Siskiyou
Co., Calif. 6, E. biangulana, 3; Marin Co., Calif. 7, Gretchena concitatricana, 2°; Cave Creek
Canyon, Chiricahua Mountains, Ariz.

‘62° on the first label, with 2 specimens also bearing **‘164°’ on a second
label. Three specimens are labeled y bes Angeles’’; 3 other specimens la-
beled ‘‘through C. V. Riley 1888’: 3 d (one without abdomen), USNM
genitalia slide 17785, CH genitalia slide 19 Nov. 1917; 3 2, USNM genitalia
slides 17783, 17784, CH genitalia slide 21 Feb. 1940, #7, in the National
Museum of Natural History, Smithsonian Institution; | d, in University of
California, Berkeley. [Two pupal exuviae are included with this series, la-
beled—*‘62, Cal., Coquillett, Collector, through C. V. Riley 1888”’.]
Remarks.—Gretchena obsoletana can be separated from other species in
the genus by the uniform, grayish-brown color of the apical half of the

forewing, lacking median and apical streaks. This species can be superfi-

cially distinguished from Epinotia biangulana, also occurring in California,

600

Fig. 8. Gretchena concitatricana, ° genitalia, mated; Madera Canyon, Santa Rita Moun-_
tains, Ariz., USNM genitalia slide 17785. Scale line = 1 mm.

VOLUME 84, NUMBER 3 601

by the shape of the forewing termen, slightly convex in G. obseletana and
emarginate in E. biangulana. The latter species also has hindwings that are
mottled rather than being uniform in coloration (Figs. 5, 6).

Gretchena Heinrich
Gwendolina Heinrich, 1923. New Synonymy.

__ Gwendolina was described by Heinrich (1923) to include a single species,
concitatricana Heinrich (Fig. 7). Heinrich distinguished this species from
_Gretchena by wing venation, male genitalia, and secondary sexual charac-
ters. According to his description, the hindwing has veins M, and Cu, con-
nate or very short stalked; the male genitalia (figured by Heinrich) are sim-
ilar to Gretchena except that the socii are not strongly chitinized nor
elongately triangular but are as broad as long and almost circular; the male
) has a forewing costal fold and heavy black sex scaling on the upper surface
of the abdomen, on the upper surface of the inner margin of the hindwing,
and along the upper and lower margins of the cells on the undersides of
both wings.

The acquisition of additional material, and the reassessment of characters
used for defining genera provide the basis for synonymizing Gwendolina
and Gretchena. Ina series of concitatricana collected by Ronald W. Hodges
in Arizona at Madera Canyon, Santa Rita Mountains, veins M, and Cu,
vary from nearly connate to stalked over one-third the length. The differ-
ences in the male genitalia between Gwendolina and Gretchena are only of
species level significance. The female genitalia of concitatricana are similar
to other Gretchena species, differing in having short projections on the
posterior lateral angles of the seventh sternum (Fig. 8). The absence of
secondary sexual characters are not of generic significance. These charac-
ters, such as the costal fold and sex scales, are reduced or lost independently
in many groups, e.g. Epinotia and Rhopobota Lederer.

ACKNOWLEDGMENTS

| thank John G. Franclemont and L. L. Pechuman, Cornell University,
‘Ithaca, N.Y., Don Davis, Department of Entomology, Smithsonian Insti-
tution, Washington, D.C., and Jerry A. Powell, University of California,
Berkeley, for the loan of specimens. Figures 1, 4, and 8 were drawn by
Amy Trabka; Fig. 3 was drawn by Steve Sierigk. Photographs of the imagos
were made by Howard Lyons. This research was supported in part by Na-
tional Science Foundation Grant DEB 77-15808 and a Grant-in-Aid of Re-
search from Sigma Xi, the Scientific Research Society of North America.

LITERATURE CITED

Heinrich, C. 1923. Revision of the North American moths of the subfamily Eucosminae of the
family Olethreutidae. U.S. Natl. Mus. Bull. 123, 1V+ 298 pp.

602 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

MacKay, M. R. 1959. Larvae of the North American Olethreutidae (Lepidoptera). Can. Ento-
mol. Suppl. 10, 338 pp.

McDunnough, J. H. 1925. New Canadian Eucosminae (Lepidoptera). Can. Entomol. 57: 115—
116.

Payne, J. A., H. L. Malstrom, and G. E. KenKnight. 1979. Insect pests and diseases of the
pecan. U.S. Dep. Agric. Agric. Rev. and Man. S-5, 43 pp.

Tedders, W. L. and M. Osburn. 1970. Morphology of the reproductive system of Gretchena
bolliana, the pecan bud moth. Ann. Entomol. Soc. Am. 63: 786-789.

NOTE

Change of Name of a North American Ypsolopha
(Lepidoptera: Plutellidae)

Cataloging of the world microlepidoptera fauna has uncovered an over-
looked homonym caused by the inclusion of two species of the same name
in the plutellid genus Ypsolopha. The species homonymy was previously
not existent due to the prevalent usage of the genus name Cerostoma
Latreille. Ypsolopha Latreille, 1796, and Cerostoma Latreille, 1802, are
now considered congeneric. The following correction is therefore recorded:

Ypsolopha buscki Heppner, New NAME
Cerostoma manella Busck, 1903, J. N.Y. Entomol. Soc. 11: 51 (type-
locality: Williams, Arizona), preocc. in Ypsolopha by Ypsolophus
[sic] manella Moschler, 1890, Abh. Senckenberg Nat. Ges. 15:
344 (type-locality: Puerto Rico).

John B. Heppner, Department of Entomology, Smithsonian Institution,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 603-609

LARVAL HABITATS AND MATE-SEEKING SITES OF FLOWER
FLIES (DIPTERA: SYRPHIDAE, ERISTALINAE)

CHRIS T. MAIER

Department of Entomology, The Connecticut Agricultural Experiment
Station, P.O. Box 1106, New Haven, Connecticut 06504.

Abstract.—Larval habitats of 26 species and mate-seeking locations of 30
species of Syrphidae (subfamily Eristalinae) are described. Males search for
mates near flowering plants and/or larval habitats. These observations con-
firm and extend previous findings, which indicate that males seek mates
near resources utilized by females.

Male syrphids search for mates near resources that attract females.
Species of Eristalinae (=Milesiinae) studied by Maier and Waldbauer
(1979a) use flowering plants, larval habitats, or both as sites for finding
mates. Females visit blossoms to feed upon pollen and sometimes nectar,
the former being necessary for rapid ovarian development (Maier, 1978;
Schneider, 1948). Maier and Waldbauer (1979a, b), Morse (1981), and many
others have investigated sexual and foraging behavior at flowers. Maier
(1978) and Maier and Waldbauer (1979a) have examined reproductive be-
havior of native species near larval habitats. Individual males defend terri-
tories centered around larval habitats and thus gain access to females arriv-
ing to lay eggs (Maier and Waldbauer, 1979a).

Metcalf (1913, 1916) used his own studies and those of Verrall (1901),
Williston (1886), and others to categorize syrphid larval habitats, mate-seek-
ing sites for male Eristalinae (Maier and Waldbauer, 1979a). Valuable com-
pilations by Coe (1953), Dixon (1960), Hartley (1961), Heiss (1938), Johann-
sen (1935), and others supplemented this early work. Nevertheless, larval
habitats of most of the species of Eristalinae are unknown or poorly de-
scribed. In this paper, I describe the larval habitats of 26 species, eight for
the first time, and extend Maier and Waldbauer’s (1979a) and Maier’s (1980)
observations on male mate-seeking behavior.

MATERIALS AND METHODS

Third-instar larvae were collected from rotting areas of cacti and trees
and from sewage settling and woodland ponds between November and May

604 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

1974-1981 and were then released in uncovered Petri dishes containing their
apparent food. Petri dishes were enclosed in plastic vegetable crispers that
were partially filled with sand, a suitable material for pupariation. Rearing
was done in growth chambers adjusted to 24 + 2°C and to a 16 h light: 8 h
dark photoperiod. Less than one month after collection, larvae pupariated
in the sand. Adults eclosed 2 weeks or less after pupariation.

Male mate-seeking behavior was studied in Arizona, Connecticut, Illinois,
and North Carolina between April and July 1972-1981. Individual males
were observed for 30 min or until their departure from a mate-seeking lo-
cation. Each observation (Table 1) represented a male that pursued con-
specific flies, mated, or engaged in both activities at a mate-seeking site.
Representatives of all species mentioned here are deposited in my syrphid
collection.

RESULTS AND DISCUSSION

LARVAL HABITATS

The following list gives the larval habitats of Syrphidae in the subfamily
Eristalinae. The species is followed by the locality, number reared (in pa-
rentheses), and description of larval habitat.

Brachyopini

Brachyopa vacua Osten Sacken.—Mason Co., Illinois. (6). Sap-soaked de-
caying wood and associated plant growth in live Fagus grandifolia Ehrh.
and Quercus spp.

Myolepta nigra Loew.—Mason Co., Illinois. (4). Moist detritus in treehole
of live Quercus marilandica Muenchh. and Q. velutina Lam.

Myolepta varipes Loew.—Gibson Co., Indiana. (2). Moist detritus in tree-
hole of live Acer saccharum Marsh.

Cerioidini
Ceriana abbreviata (Loew).—Champaign Co., Illinois. (3). Sap-soaked de-

tritus in treehole of live Liquidambar styraciflua L. (treehole with Cam-
ponotus sp. colony).

Eristalini

Eristalinus aeneus (Scopoli).—Champaign Co., Illinois. (200). Sewage set-
tling pond with standing water.

Helophilus fasciatus Walker.—Champaign Co., Illinois. (5). Small, leaf-
filled pools in mesic forest dominated by Acer, Fagus, and Quercus spp.

Mallota bautias (Walker).—New Haven Co., Connecticut. (2). Wet detritus

VOLUME 84, NUMBER 3 605

in decayed center of live Pyrus malus L. (opening to outside less than 2
cm in diameter).

Mallota posticata (Fabricius)—New Haven Co., Connecticut. (4). Mason
Co., Illinois. (200). Gibson Co., Indiana. (1). Montgomery Co., Virginia.
(3). Wet detritus in large treeholes or in water-filled crotches of many
species of live deciduous trees (treeholes usually with entrances of 10-20
cm diameter).

Milesiini

Blera umbratilis (Williston).—Mason Co., Illinois. (1). Moist detritus in tree-
hole of live Quercus velutina Lam.

Chalcosyrphus chalybeus (Wiedemann).—New Haven Co., Connecticut.
(3). Champaign Co., Illinois. (2). Moist to wet detritus in rot pockets of
fallen Acer saccharum Marsh. logs.

Chalcosyrphus metallicus (Wiedemann).—Union Co., Illinois. (6). Pulpy
wood of partially submerged Acer sp. log in small stream.

Chalcosyrphus nemorum (Fabricius).—Litchfield Co., Connecticut. (47).
Champaign Co., Illinois. (12). Piatt Co., Illinois. (5). Union Co., Illinois.
(4). Spaces (partly filled with detritus and insect frass) in firm, moist
decaying wood near surface of fallen logs of Acer and Ulmus spp. in
marshes and along streams. Actual site of larval development may be
elsewhere in logs as these larvae were collected after feeding was com-
pleted.

Lejota aurea (Loew).—Mason Co., Illinois. (10). Friable, slightly moist de-
caying wood at top of Quercus sp. stump.

Milesia virginiensis (Drury).—Mason Co., Illinois. (2). Gibson Co., Indiana.
(1). Wet detritus in water-filled center of decaying stumps.

Somula decora Macquart.—New Haven Co., Connecticut. (3). Mason Co.,
Illinois. (50). Moist detritus in treeholes of live Quercus spp. (treeholes
with elongate vertical openings).

Spilomyia alcimus (Walker).—Washtenaw Co., Michigan. (5). Wet detritus
in treehole of live Quercus alba L.

Spilomyia longicornis Loew.—Mason Co., Illinois. (3). Wet detritus along
wall of treeholes in live Quercus velutina Lam.

Temnostoma alternans Loew.—Middlesex Co., Connecticut. (2). Firm,
moist decaying wood in fallen Betula alleghaniensis Brit. log in marshy
area (wood-boring larvae).

Temnostoma barberi Shannon.—Middlesex Co., Connecticut. (10). Same
as previous species.

Temnostoma vespiforme (Linnaeus).—Middlesex Co., Connecticut. (1).
Same as previous two species.

606 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Volucellini

Copestylum apiciferum (Townsend).—Pima Co., Arizona. (20). Soupy, de-
caying material in rot pockets in trunk of live and dead Ferocactus wis-
lizenii (Engelm.) Brit. & Rose.

Copestylum florida (Hull).—Marion Co., Florida. (35). Monroe Co., Flori-
da. (23). Rotting areas in joints of live Opuntia spp.

Copestylum isabellinum (Williston).—Pima Co., Arizona. (35). Soupy, de-
caying material in rot pockets in upright and fallen trunks of live and dead
Carnegiea gigantea Brit. & Rose.

Copestylum marginatum (Say).—Pima Co., Arizona. (7). Rotting areas in
thin, non-basal joints of live Opuntia phaeacantha Engelm.

Copestylum mexicanum (Macquart).—Pima Co., Arizona. (5). Rotting areas
in large, basal joints of live Opuntia phaeacantha Engelm.

Copestylum vittatum Thompson.—Marion Co., Florida. (69). Mason Co.,
Illinois. (8). Rotting areas in joints of live Opuntia sp. (Florida) and O.
compressa (Salisb.) Macbr. (Illinois).

Larvae of the 26 reared species inhabited many different types of decaying
material. Except for Eristalinus aeneus larvae collected from sewage set-
tling ponds, all lived in natural habitats, mostly decaying parts of living and
dead cacti or deciduous trees. My descriptions of larval habitats of Ceriana
abbreviata, Chalcosyrphus chalybeus, C. metallicus, Helophilus fasciatus,
Spilomyia alcimus, S. longicornis, Temnostoma alternans, and T. barberi
are the first.

Habitats listed here are similar to those described by earlier workers (e.g.
Dixon, 1960; Hartley, 1961; Heiss, 1938) for the same or congeneric species.
My descriptions augment previous ones by giving data on plant identities,
moisture conditions, and/or other features around the larval habitats. This
new information contributes to a better understanding of the range of larval
habitats used by genera and tribes of Eristalinae.

Two or three species sometimes coexisted in the same log (Temnostoma
alternans, T. barberi, T. vespiforme), treehole (Mallota posticata, Somula
decora, Spilomyia longicornis), or cactus (Copestylum marginatum, C.
mexicanum). A decaying area in a particular log, rot pocket, or cactus was
usually inhabited by a single species. For example, Copestylum marginatum
and C. mexicanum larvae were spatially separated by about | m in the same
cactus. Copestylum marginatum larvae (n = 7) lived in thin, non-basal
joints of Opuntia phaeacantha Engelm.; and C. mexicanum larvae (n = 5),
which are larger, inhabited thicker, basal joints.

MATE-SEEKING SITES

Males searched for females near flowering plants, potential larval habitats
(oviposition sites), or both (Table 1). Although male mate-seeking was

VOLUME 84, NUMBER 3 607

Table 1. Mate-seeking locations of male Syrphidae in the subfamily Eristalinae. Evidence:
1 = initiation of copulation observed near flowers; 2 = initiation of copulation observed near
larval habitat; 3 = mating pair(s) observed near flowers; 4 = mating pair(s) observed near
larval habitat; and 5 = males chased conspecific flies and other flying insects of similar size
and grappled with conspecifics (territorial defense).

Tribe and Species Number Observed Evidence

A. Flowering plants

Brachyopini
Neoascia distincta Williston 4 5
N. globosa (Walker) >40 15Sie5
Sphegina flavimana Malloch 3 5
S. keeniana Williston 13 Te
S. lobata Loew 5 5
S. rufiventris Loew 21 5
Eristalini
Lejops albiceps (Macquart) 12 5
L. anausis (Walker) 3 5
L. bilinearis (Williston) 32 IBS)
L. chrysostomus (Wiedemann) 15 Is)
L. stipatus (Walker) 9 5
Parhelophilus anniae (Brimley) 6 5
Milesiini
Blera analis (Macquart) 4 5
Chalcosyrphus chalybeus (Wiedemann) 2 il.
Criorhina verbosa (Walker) 11 iN,
Tropidia albistylum Macquart 2) il.
Xylota angustiventris Loew 3 5
Rhingiini
Rhingia nasica Say >30 5
Volucellini
Copestylum mexicanum (Macquart) 15 5
Volucella bombylans (Linnaeus) 2 5
B. Larval habitat
Milesiini
Chalcosyrphus metalliferus (Bigot) 4 25
C. nemorum (Fabricius) >40 255
C. plesia Curran 2 5
C. Both A and B
Brachyopini
Sphegina campanulata Robertson > 100 Ie2s' 399
Merodontini
Eumerus tuberculatus (Fabricius) 23 4,5
Merodon equestris Rondani 4 5

gS

608 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Continued.

Tribe and Species Number Observed Evidence
Milesiini
Brachypalpus oarus (Walker) 9 5. 255)
Temnostoma alternans Loew 16 1: 23a
T. balyras Walker >80 TENS)
Xylota bicolor Loew 7 5

concentrated near these resources, its behavioral characteristics varied
among species. As described by Maier and Waldbauer (1979a), males of
most species alternately rested on flowering plants and patrolled blossoms
to locate potential mates at flowers. Hovering near blossoms sometimes
accompanied patrolling or occurred by itself. For example, on July after-
noons between 1200 and 1500 EST, groups of 2-20 Rhingia nasica Say
males hovered directly above patches of flowering Jmpatiens biflora Walt.
These males were equally spaced except when they pursued flying insects
or conspecific males that entered or passed by their aerial territories. After
pursuits males returned to their original hovering spot, suggesting they have
visual spatial memory (Collett and Land, 1975).

Within | m of larval habitats, males defended territories. They spent 80—
90% of their time resting in places that afforded an unobstructed view of
females arriving to oviposit. Representative examples of mate-seeking be-
havior at flowering plants and larval habitats are more fully described by
Gruhl (1924) and Maier and Waldbauer (1979a).

Fifteen species of Eristalinae, eight listed by Maier and Waldbauer (1979a,
Table 4) and seven recorded here (Table 1), employ dual mate-seeking strat-
egies. As larval habitats of more species become known, this number will
surely grow. The present evidence, however, strongly suggests that dual
mate-seeking strategies are common among species of Eristalinae. Compre-
hensive studies are necessary to determine the entire range of mate-seeking
strategies utilized in the other two subfamilies, the Microdontinae and the
Syrphinae.

ACKNOWLEDGMENTS

I sincerely thank Mark S. McClure and Ronald M. Weseloh (Department
of Entomology, The Connecticut Agricultural Experiment Station, New
Haven) for comments on an earlier version of this manuscript. Helmut Maier
and John A. Maier directed me to collecting sites in Florida and Arizona,
respectively. Joel R. Coats made a special contribution by collecting the
Ceriana abbreviata larvae. Stephen W. Bullington reared the Mallota pos-
ticata larvae from Virginia.

eee OO EEO

VOLUME 84, NUMBER 3 609

LITERATURE CITED

Coe, R. L. 1953. Handbooks for the identification of British insects. Vol. X(1). Diptera:
Syrphidae. R. Entomol. Soc. Lond. 98 pp.

Collett, T. S. and M. F. Land. 1975. Visual spatial memory in a hoverfly. J. Comp. Physiol.
100(A): 59-84.

Dixon, T. J. 1960. Key to and descriptions of the third instar larvae of some species of
Syrphidae (Diptera) occurring in Britain. Trans. R. Entomol. Soc. Lond. 112: 345-379.

Gruhl, K. 1924. Paarungsgewohnheiten der Dipteren. Z. Wiss. Zool. 122: 205-280.

Hartley, J. C. 1961. A taxonomic account of the larvae of some British Syrphidae. Proc. Zool.
Soc. Lond. 136: 505-573.

Heiss, E. M. 1938. A classification of the larvae and puparia of the Syrphidae of Illinois
exclusive of aquatic forms. Ill. Biol. Mongr. 16, 142 pp.

Johannsen, O. A. 1935. Aquatic Diptera. Part Il. Orthorrhapha-Brachycera and Cyclorrhapha.
Mem. Cornell Univ. Agric. Exp. Stn. 177, 62 pp.

Maier, C. T. 1978. The immature stages and biology of Mallota posticata (Fabricius) (Diptera:

Syrphidae). Proc. Entomol. Soc. Wash. 80: 424-440.

. 1980. Mate-seeking strategies in male flower flies (Diptera: Syrphidae). J. N.Y. Ento-

mol. Soc. 87: 59-60.

Maier, C. T. and G. P. Waldbauer. 1979a. Dual mate-seeking strategies in male syrphid flies

(Diptera: Syrphidae). Ann. Entomol. Soc. Am. 72: 54-61.

. 1979b. Diurnal activity patterns of flower flies (Diptera: Syrphidae) in an Illinois sand

area. Ann. Entomol. Soc. Am. 72: 237-245.

Metcalf, C. L. 1913. Syrphidae of Ohio. Ohio Biol. Surv. Bull. 1: 7-123.

. 1916. Syrphidae of Maine. Maine Agric. Exp. Stn. Bull. 253: 193-264.

Morse, D. H. 1981. Interactions among syrphid flies and bumblebees on flowers. Ecology 62:
81-88.

Schneider, F. 1948. Beitrag zur Kenntnis der Generationsverhaltnisse und Diapause rauber-
ischer Schwebfliegen (Syrphidae, Dipt.). Mitt. Schweiz. Entomol. Ges. 21: 249-285.

Verrall, G. H. 1901. British flies. Vol. VIII. Platypezidae, Pipunculidae, and Syrphidae of
Great Britain. Gurney and Jackson, London. 691 pp.

Williston, S. W. 1886. Synopsis of North American Syrphidae. Bull. U.S. Natl. Mus. 31: I-
83555

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 610-616

SOME NEW NEOTROPICAL LEAFHOPPERS OF THE SUBFAMILIES
IASSINAE AND DELTOCEPHALINAE
(HOMOPTERA: CICADELLIDAE)

DwiGHT M. DELONG

Department of Entomology, Ohio State University, Columbus, Ohio
43210.

Abstract.—Six new species of Neotropical leafhoppers, Krisna aesta
(Puerto Rico), Scaroidana blockeri (Bolivia), Tenucephalus hamatus (Boliv-
ia, Argentina), Mendozellus serratus (Bolivia), M. tarandus (Bolivia), and
M. incisus (Bolivia) are described. Bolarga nigriloba Linnavuori is placed
in the genus Daltonia.

The genus Krisna was described by Baker (1919). Osborn (1938) described
Scaroidana, and Kramer (1963) treated the known species of the genus. The
genus Tenucephalus was described by DeLong (1944). Linnavuori (1959)
described Mendozellus as a subgenus of Amplicephalus.

A new species of each Krisna, Scaroidana, and Tenucephalus, and three
new species of Mendozellus are described in this paper. Bolarga nigriloba
Linnavuori is placed in the genus Daltonia. All holotypes are in the DeLong
collection, Ohio State University, Columbus.

Krisna aesta DeLong, NEw SPEcIES
Figs. 1-5

Description.—Length of male 10 mm, female unknown. Crown thin, fo-
liaceous, twice as wide at base, between eyes, as long at middle. Crown,
pronotum and scutellum dull yellow. Forewing yellow with slight tint of
green, without contrasting color markings.

Male genital plates slender, elongate, 9x as long as wide at middle, apices
narrow, rounded. Style bearing a ventral spine at more than 4/s its length,
apical '/s curved dorsally, apex pointed. Aedeagus short and broad in lateral
view, with apical portion divided and curved dorsobasally, extending twice
width of shaft. Pygofer with rounded, blunt projection apically.

Holotype.—d , Manayes, Puerto Rico, May 25, 1924, G. N. Wolcott coll.

Remarks.—Krisna desta is related to K. insularis Oman and can be sep-
arated from it by the absence of small spots on the forewing, by the shorter,

VOLUME 84, NUMBER 3 611

broader aedeagus, and by the wider, more foot-shaped apical portion of the
style.

This may prove to be the male of K. montana Caldwell (in Caldwell and
Martorell 1952: 19) which was described from female specimens. There are
no color markings on either male or female to assist in the identification.
For illustrations of the structural features of the other Puerto Rican members
of the genus see Caldwell and Martorell (1952: 21).

Scaroidana blockeri DeLong, NEw SPECIES
Figs. 6-10

Description.—Length of male 11 mm, female 11.5 mm. Crown broadly
rounded, short and broad, more than 4x as broad between eyes at base as
long at middle. Crown, pronotum, and scutellum dull, sordid yellowish
green with a brownish tint; reddish color of abdomen often showing through
wings. Apical portion of wing including all apical cells, dark brown, smoky.

Female 7th sternum with posterior margin broadly, slightly, concavely
rounded.

Male plates more than 3x as long as wide at middle, apices rounded.
Style elongate, slender, apical 2 gradually tapered to a narrow, pointed,
dorsally bent apex. Ventral margin of apical position serrate. Aedeagus
narrow dorsoventrally, rather broad with apical portion of shaft curved dor-
sally and bluntly pointed in dorsal view. Shaft broadened at middle with a
thin spinelike portion extending dorsally. Pygofer rounded apically and bear-
ing prominent ventral spine which curves dorsally on inner margin of py-
gofer at apex, and bears enlarged portion at % length of spine.

Types.—Holotype 6d, Bolivia, Santa Cruz, Saavedra, 250 m, 2-IV-1980,
D. R. Foster coll. Paratypes: | ¢ same as holotype; 1 ¢, 13 2 same except
12-[V-1979; 1 2, Santa Cruz, 2-III-79, Foster coll.

Remarks.—Scaroidana blockeri is related to §. fulvula Osborn (see Kra-
mer 1963: 42) and can be separated from it by the narrower basal portion

of the aedeagus, the enlarged process on the median portion of the pygofer

spine, and the dorsally curved pointed apex of the style.

I take pleasure in naming this species for Dr. H. Derrick Blocker, Kansas
State University, who assisted me with the generic placement of this
species.

Tenucephalus hamatus DeLong, NEw SPECIEs
Figs. 11-15

Description.—Length of male 8 mm, female unknown. Crown broadly
rounded, more than twice as wide at base between eyes as long at middle.
Crown dull smoky greenish, with black marginal band between eyes. Pro-
notum and scutellum same color as crown. Forewing dull white, opaque,

612 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. |-5. Krisna aesta. 1, Aedeagus laterally. 2, Aedeagus ventrally. 3, Pygofer laterally,

apical portion. 4, Plate ventrally. 5, Style laterally. Figs. 6-10. Scaroidana blockeri. 6, Plate
ventrally. 7, Aedeagus ventrally. 8, Style laterally. 9, Aedeagus laterally. 10, Pygofer with

spine. Figs. 11-15. Tenucephalus hamatus. 11, Aedeagus ventrally. 12, Style laterally. 13,
Pygofer laterally, apical portion. 14, Plate ventrally. 15, Aedeagus laterally.

a_i

— = —— ——

VOLUME 84, NUMBER 3 613

veins brown. Fuscous blotch on costal area at more than % its length. Apical
portion smoky. Veins of apical 43 broadly margined with brown.

Male genital plates more than twice as long as wide at middle, apices
broadly rounded. Style rather short and broad, apophysis curved outwardly
with apex rounded. Aedeagal shaft narrowed near apex and terminating in
a slender dorsobasally curved portion, widened at middle, slender, sharply
pointed apically, extending % length of shaft. Shaft bearing a short ventral
tooth at point of curvature and with a broad platelike process on ventral
margin subapically. Pygofer bearing sharply pointed tooth on ventro-caudal
margin.

Types.—Holotype ¢, Bolivia, Santa Cruz, 36 miles S, 26-IV-1978, 500 m,
C. E. Ward coll. at light, Paratypes: 5 2, 1 6, Tucuman, Argentina, Jan.
1947, Fernandez coll.; 1 ¢, 1 2 Urundel, Salta, Argentina, 1-31-1950, R.
Gelbach coll.

Remarks.—Tenucephalus hamata is related to T. sagittarius Linnavuori
and DeLong (1976: 29) and can be separated from it by the slender dorso-
basally curved apical portion of the aedeagal shaft.

Mendozellus Linnavuori

The genus Mendozellus contains a group of closely related species found
in Bolivia, Argentina, and Peru. Color patterns of a few of these species are
quite similar and the color pattern of each is variable. In general, the species
can usually be separated by the male or female genital structures. Mendoz-
ellus lineiceps (Osborn) was described from female specimens and cannot
be separated by genital structures. Mendozellus incisus, n. sp., which is
being described from a female specimen in the following pages, has an
incised seventh sternum and can easily be separated from its congeners on
this basis.

Mendozellus serratus DeLong, NEW SPECIES
Figs. 16-21

Description.—Length of male 3.5 mm, female 3.7 mm. Crown produced
and broadly angled, longer at middle than wide at base between eyes. Crown
whitish with round black spot on each side of apex, elongate black spot just
above margin between round black spot and eye, and angled black spot on
each side between eyes at % length of crown. Longitudinal orange stripe
extends just behind round black apical spot, on each side, along eyes and
continues across pronotum. Pronotum white with 2 longitudinal orange
stripes behind each eye. Scutellum orange, basal angles black, central por-
tion and apical angle white. Forewing smoky, veins white, margined with
brown.

Female 7th sternum with posterior margin broadly, shallowly, concavely
rounded.

614 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

J

21

= 7

26

Figs. 16-21. Mendozellus serratus. 16, Aedeagus and connective ventrally. 17, Aedeagus
and connective laterally. 18, Head, pronotum and scutellum. 19, Pygofer laterally. 20, Style |
laterally. 21, Plate ventrally. Figs. 22-27. M. tarandus. 22, Aedeagus ventrally. 23, Connective |
ventrally. 24, Aedeagus laterally. 25, Aedeagus apically. 26, Style laterally. 27, Plate ventrally. |
Fig. 28, M. incisus, 7th sternum.

Male genital plates 22x as long as wide at middle, apices bluntly pointed.
Style elongate, apophysis with apex rounded. Aedeagal shaft short, nar-
rowed and curved dorsally on apical 4%. A pair of slender, apically pointed
processes arise near base of shaft on ventral margin and extend caudally

VOLUME 84, NUMBER 3 615

along and beyond shaft to length of caudally directed portion. Pygofer with
ventral margin bearing row of irregular, sharply pointed “‘teeth.”’

Types.—Holotype ¢, El Gaite, Bolivia, 18-III-1980, on Maiz, Don Foster
coll. Paratypes: 5 2 same data as holotype; 2 ¢d Santa Cruz, Bol., 19-VIII-
°80; 1 ¢, Buena Vista, Bol., 14-V-°80, DeLong coll.; 2 ¢ Portachuela, Bol.,
14-V-’80, DeLong coll.

Remarks.—Mendozellus serratus is related to M. devius Linnavuori and
DeLong (1977: 251) but differs in color pattern as described, by the long
ventral processes of the aedeagus, and the serrate ventral margin of the
pygofer.

Mendozellus tarandus DeLong, NEw SPECIES
Figs. 22-27

Description.—Length of male 3 mm, female unknown. Crown angularly
produced, apex blunt, rounded, slightly longer at middle than width at base
between eyes. Crown orange brown with longitudinal white stripe extending
from apex to apex of scutellum, and commissure of forewing white. Elon-
gate dark brown spot on each side of white stripe at apex of crown, and
similar pair at “% and %4 length of crown. Brown line parallel to margin back
of each ocellus extending to eyes. Pronotum brown with darker brown lon-
gitudinal stripe on each side of median white stripe, and 6 narrow white

_ longitudinal stripes. Scutellum brown with dark brown longitudinal stripe

———— Se

——__

bordering white stripe, on each side. Forewing brownish subhyaline, veins
white.

Male genital plates almost twice as long as wide at middle, apices bluntly
pointed. Style with straight, finger-like apophysis. Aedeagus divided at %
its length, forming 2 curved processes which extend dorsally and curve
laterally, with apices proximal. Connective elongate with proximal apices.
Pygofer bluntly pointed apically.

Types.—Holotype ¢, Bolivia, Santa Cruz, 21-VII-°80, D. R. Foster coll.
Paratypes: 3 ¢ same as holotype; | ¢ same except 17-IX-’80; | d same
except 27-V-’80; | ¢ same except 3-VIII-’80.

Remarks.—Mendozellus tarandus is closely related to M. isis Linnavuori
(1959: 117) and can be separated from it by the slender, dorsally curved
apical processes of the aedeagus and their slender pointed tips.

Mendozellus incisus DeLong, NEw SPECIES
Fig. 28
Description.—Length of female 4 mm, male unknown. Crown angularly
produced, as long at middle as wide between eyes at base. Crown roundly
angled at apex. Color dull yellowish, pair of rounded, slightly elongated,
brown spots at apex. Two pairs of similar brownish spots, more elongate,

616 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

between eyes, % and % length of crown. Pronotum mostly white with 2
brownish longitudinal stripes on each side of median white stripe, and faint
brownish longitudinal stripe behind each eye. Scutellum white with brown
basal angles. Forewing pale brownish subhyaline, veins white.

Female 7th sternum with posterior margin broadly convexly rounded each
side of a narrow incision half way to base of segment which is broadly
embrowned.

Holotype.—@ , Bolivia, Santa Cruz, 20-IV-’80, Don Foster coll.

Remarks.—Mendozellus incisus can be separated from all other described
species of Mendozellus by the deeply incised seventh female segment.

Daltonia nigriloba (Linnavuori), NEw COMBINATION

Linnavuori described D. nigriloba and placed it in Bolarga. A series of
specimens from Bolivia and Argentina, one of which was identified by Lin-
navuori as Bolarga nigriloba, are very similar in form, coloration, and ap-
pearance to specimens of Daltonia estacada (Ball), which is a common
species in the southern United States and in the lower gulf coastal area of
Mexico. The forewings of D. nigriloba are a little more elongate and the
apical cells are longer than those of D. estacada. The aedeagus of D. ni-
griloba is similar to that of D. estacada, but it is not notched apically, and
the apophysis of the style is more elevated and pointed. Bolarga nigriloba
is considered a good species and is placed in Daltonia.

LITERATURE CITED

Baker, C. F. 1919. The genus Krisna (Jassidae). Philipp. J. Sci. 15: 209-226.

Caldwell, J. S. and L. F. Martorell. 1952. Review of the Auchenorhynchous Homoptera of
Puerto Rico, Part I, Cicadellidae. J. Agric. Univ. P. R. 34: 1-132.

DeLong, D. M. 1944. A New Genus (Tenucephalus) and species of Mexican leafhopper related
to Parabolocratus. Ohio J. Sci. 44: 236-237.

Kramer, J. P. 1963. A key to the new world species of Jassinae with reviews of Scaroidana
and Pachyopsis (Homoptera: Cicadellidae) Bull. Brooklyn Entomol. Soc. 58: 37-50.

Linnavuori, R. 1959. Revision of the Neotropical Deltocephalinae and some related subfamilies
(Homoptera). Ann. Zool. Soc. ‘Vanamo’ 20(1): 1-370.

Linnavuori, R. and D. M. DeLong. 1976. New Neotropical leafhoppers from Peru and Bolivia
(Homoptera: Cicadellidae). Rev. Peru. Entomol. 19(1): 29-36.

—. 1977. The leafhoppers (Homoptera: Cicadellidae) known from Chile. Brenesia 12/13:
163-267.

Osborn, H. 1938. Neotropical leafhoppers of the Carnegie Museum, Part 7, report on the
species of the subfamily Gyponinae. Ann. Carnegie Mus. 27: 11-62.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 617-627

ETHOLOGY OF NEOITAMUS VITTIPES (DIPTERA: ASILIDAE)
IN SOUTH AUSTRALIA’?

ROBERT J. LAVIGNE

Entomology Section, University of Wyoming, Box 3354, University Sta-
tion, Laramie, Wyoming 82071.

Abstract.—A field study of the predatory and mating behavior of Neoita-
mus vittipes (Macquart) is reported. Prey taken by N. vittipes represented
five orders, the majority being in the order Diptera. Mating, without prior
courtship, took place in a tail-to-tail position, mostly in the afternoon. In-
cidental data is included concerning distribution and predatory behavior of
Neoitamus armatus (Macquart) and N. margites (Walker).

With the exception of the recent paper by Daniels (1976) on the predatory
and courtship behavior of Promachus interponens Walker, nothing has been
written of the behavior of Australian robber flies. Thus far, the majority of
behavioral studies have been done on species in North America, Russia,
and Europe.

Neoitamus vittipes (Macquart)

Published records of the occurrence of Neoitamus vittipes (Macquart) are
few. The original type-series was collected in Tasmania, but the synonymy
established by Hardy (1920) extended its distribution to continental Austra-
lia, with records from Victoria and New South Wales (January to March).
While on sabbatical at the Waite Agricultural Research Institute (November
1978—May 1979), I collected N. vittipes at the following locations in South
Australia: 2.6 km N of Port Wakefield (23.xi.78), 6 km SW of Virginia
(23.xi.78), 4.5 km E of Keyneton (30.xi.78), 16 km N of Kingston (18-
19.xii.78), ca. 12 km SE of Milang (20.xii.78, 1.11.79), 3 km E of Milang
(8.1.79, 2—3.ii1.79), 8 km E of Milang (19.xii.78, 12.11.79, 1.11.79), 10 km S

' Published with the approval of the Director, Wyoming Agricultural Experiment Station, as
Journal Article No. JA 1111.

? This research was supported in part by National Science Foundation Travel Grant INT 78-
17131, in part by the Waite Agricultural Research Institute, Adelaide, Australia, and in part by
the University of Wyoming.

618 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of Milang (20.xii.78), 11.6 km NW of Morgan (22-27.11.79), 2.7 km S of
Kapunda, bank of Light River (22.111.79). Additionally, specimens in student
collections provided records for Adelaide (early February 1979) and Port
Gawler (16.ii.77, *‘flying around Melaleuca’’).

Typical habitats from which N. vittipes was collected included rocky
hillside pastures dominated by Avena barbata Pott ex Link, dryland lucerne
(alfalfa) fields, borders of irrigated lucerne fields, weedy borders of crop-
land, a weedy horse paddock and open rangeland dominated by Maireana
sedifolia (F. Muell.) Wilson (bluebush) and Danthonia sp. (wallaby grass).

Most behavioral observations were made on a population inhabiting a
field of irrigated lucerne 3 km east of Milang, S.A. This population was
largely concentrated within the first 5 m of lucerne along the borders of the
field, except where incursions of grass created clearings. The asilids would
then be found foraging from individual plants bordering the clearings. The
species was studied during the period March 5 to 13, 1979 at this site; a few
additional observations were made December 18-19 in a dryland lucerne
field opposite Coorong National Park, 16 km N of Kingston.

METHODS

Approximately 100 localities in South Australia were surveyed during
spring and summer of 1978-79 to ascertain the distribution and abundance
of species of robber flies. An hour or two of walking through a particular
habitat was usually sufficient to establish what species were present and
their density. In disturbed habitats in South Australia, it was not uncommon
to find only one species present, whereas many undisturbed areas, e.g.,
conservation parks, supported several species concurrently.

Once a suitably dense population of an asilid species was encountered,
one or two observers visited the area on several successive days. In general
two approaches were employed, (1) single flies were continuously observed
for one to two hours each, and (2) the observer slowly traversed the area
noting the activities of many different flies.

Constant and extended surveillance of individual flies yielded a mass of
information concerned primarily with feraging and feeding, whereas sur-
veying the area provided greater opportunity for observation of mating pairs
and for locating ovipositing females.

Observations were recorded in notebooks on the study site. These in-
cluded detailed records of foraging, feeding, mating, and oviposition, when-
ever observed. Times and when possible, durations of these activities were
noted. Temperatures were taken from the soil surface or the height on the
vegetation where the behavior was occurring. Permanent photographic rec-
ords of each behavior were obtained using a Nikon 35 mm, single lens reflex
camera with a Micro-Nikkor-P.C. Auto 1:3.5, 55 mm lens.

VOLUME 84, NUMBER 3 619

Recorded behavior patterns involving flight were defined as follows: (1)
Foraging flights were those in which the individual flew towards a potential
prey, regardless of whether it was successful; (2) investigatory flights where
the asilid would fly towards a potential prey, but turn back without attempt-
ing capture (presumably, size or structural characteristics functioned to
make the insect unsuitable); (3) manipulation flights (rotation flights of Scar-
brough and Norden, 1977) in which an individual would fly briefly from a
feeding site, manipulate prey, and return to the same or nearby perch; (4)

_ orientation flights in which the asilid would move to a new location as a

result of being shaded, disturbed, or not making visual contact with potential
prey over a period of time (these flights were made slowly and usually
covered only a short distance); (5) agonistic flights which involved aggres-
sive encounters between males; and (6) searching flights which consisted of
long, rapid, sometimes undulating flights taken by males in search of recep-
tive females.

Prey was collected whenever possible. Either the robber flies observed

_ feeding on prey were netted, the prey was collected, and the asilid was
_ released unharmed, or the predator was observed until completion of feed-

ing and the prey was recovered where it was dropped. The latter method,
while supplying information on feeding times and manipulation behavior,
sometimes resulted in the loss of prey, since asilids often dart after new
prey, releasing the exhausted meal in flight. Occasionally, an insect net was

_ placed carefully over the feeding asilid and held in place until completion

of feeding. However, this method lacks desirability since a puff of wind
striking the net may disturb the asilid, causing it to drop prey, which if tiny,
may disappear among the surface debris.

Collected prey were placed in a 2 dram vial and subsequently measured
for total body length to the nearest 0.5 mm, after which it was either mount-
ed by pinning or placed in 70% ETOH. Similar measurements were made
of the predator, i.e., 14 specimens of each sex. In this way, the predator to
prey size ratio was obtained. An additional procedure, although not used in
this study, is that of Scarbrough (1978), who oven-dried prey of selected
lengths to obtain predator to prey weight ratios.

Prey were identified to order-family and then submitted to specialists for
further identification (see Acknowledgments). The prey collection is housed
at Waite Agricultural Research Institute in Adelaide, S.A.

FORAGING AND FEEDING BEHAVIOR

Perch sites, from which N. vittipes launched attacks on potential prey,
varied with the character of the habitat. In the population which inhabited
the lucerne field, the majority of flies foraged from vegetation (lucerne, 62%;
weeds, 16%; irrigation pipe, 6%; soil, 10% and grass stalks, 6%) throughout

620 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Neoitamus vittipes male feeding on the syrphid, Simosyrphus grandicornis, while
holding onto lucerne leaves.

the day. However, in rocky pastures and dryland alfalfa fields, the majority
of asilids launched attacks from the soil surface or from small rocks.

Exclusive of soil the heights on plants, where foraging individuals were
observed, varied from 5 to 38 cm (mean 19.3 cm). When on soil, individuals
were either resting broadside to the sun or facing directly into it, a behavior
which is directly related to soil surface temperature in other asilid species
(Lavigne and Holland, 1969; Dennis and Lavigne, 1975, 1979). These other
asilids studied maintain temperature control through positional changes,
utilizing soil in early morning, thereby increasing heat absorption, and vege-
tation around midday when soil temperatures are excessive. Presumably the
Kingston population functioned in a similar manner.

While resting on foraging sites, the flies were largely quiescent, except
when potential prey flew within their field of vision. The asilid would then
turn its whole body to face the organism. One can assume that such postural
changes increase range of vision and place the asilid in a suitable position
to make a direct forage flight (Dennis and Lavigne, 1975).

All forage flights were directed at insects that were air borne. Distances
covered in forage flights ranged from 7.6 to 71 cm (mean 25.6 cm). Flights
which resulted in prey capture ranged in length from 7.6 to 45.7 cm (mean

a _—

a

VOLUME 84, NUMBER 3 621

Table 1. Relation between length of Neoitamus vittipes and that of its prey.

Predator Length (mm)! Prey Length (mm) Mean

Ratio of

No. Prey Predator:
Sex Min. Max. Mean Min. Max. Mean Measured Prey
Male 10.0 13.0 11.6 1.0 9.0 3u/ 12 3.1
Female ES 14.5 12.6 0.5 10.0 4.4 24 2.9
Combined 10.0 14.5 12.1 0.5 10.0 4.2 36 29

! 14 predators of each sex were measured from the Milang population.

21.4 cm), suggesting that as distance increases between predator and po-
tential prey, the asilids decrease in capture efficiency.

Most flights initiated in pursuit of prey were unsuccessful. Based on a
limited number of observations, prey capture efficiency for N. vittipes is
poor, i.e., 16%. On two occasions (one male, one female) flights were di-
rected towards cruising honeybees. In both instances, the asilid turned away
and returned to the foraging site, after covering 4% to % of the distance.
These investigatory flights undoubtedly maximize net energy gain, 1.e., low-
er pursuit costs since the asilid does not chase prey that is likely to escape.

Once prey were captured, the asilid hovered briefly, manipulated the prey
using all six tarsi and impaled it prior to landing.

During the feeding sequence, most prey were manipulated one or more
times. The prey were manipulated with all six tarsi, during a short flight I-
2 cm away from the feeding site. Manipulation, while hovering is typical of
the behavior of Asilinae in North America (Dennis and Lavigne, 1975, 1979;
Lavigne, 1979; Lavigne et al., 1976; Lavigne and Dennis, 1980).

Once feeding is completed, the chitinous exoskeleton of the prey is
pushed off the proboscis by the asilid’s foretarsi on the feeding site.

Only twice were complete feedings monitored. In one instance a female
fed upon a bushfly, Musca vetustissima Walker, for 25 minutes. A second
case involved a male, which fed upon the syrphid, Simosyrphus grandicor-
nis (Macquart), for a period of 2 hours and 10 minutes (Fig. 1).

One case of cannibalism was observed in the population 16 km N of
Kingston. In this instance a male attacked and killed a second smaller male
(Fig. 2). During the initial struggle, the larger male lay on its side on the
ground manipulating the other male with all six tarsi, a technique apparently
forced on it by the large size of the prey.

Based on 36 measured prey of N. vittipes, the ‘‘preferred’’ prey length
was 4.3 mm, although the size of prey taken varied from 0.5 to 10.0 mm
(Table 1). Females were slightly larger than males (12.6 vs. 11.6 mm) and
tended to capture slightly larger prey (4.4 vs. 3.7 mm). Males concentrated

622 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Comparison of prey items taken by three species of Neoitamus in various parts
of the world.

N. vittipes N. angusticornis N. cyanurus
(Australia) (Japan)! (England)?
Order No. % No. % No. %
Araneae 3 2.8
Coleoptera iI5) 14.2 9 14.0
Diptera 28 73.8 58 54.8 38 5}2).3)
Ephemeroptera 3 2.8
Heteroptera 2 So 3 4.7
Homoptera 4 10.6 4 3.8
Hymenoptera 2 Soe 12 11.3 | 1.6
Isoptera I 0.9
Lepidoptera 2 So 9 8.5 11 Ng/ 7
Neuroptera I 0.9 | 1.6
Odonata 1 1.6
Totals 38 100 106 100 64 100

' Records of prey collected by Iwata and Nagatomi (1962).
2 Compilation of prey records collected by Hobby (1931, 1933, 1934), and Poulton (1907).

on Diptera (90%) and only incidentally took Hemiptera (10%). Females,
while concentrating on Diptera (65%), fed on representatives of four other
orders, Hemiptera (3.9%), Homoptera (15.5%), Hymenoptera (7.8%) and
Lepidoptera (7.8%). The mean predator to prey length ratio for this species
was 2.9 (Table 1).

Various authors (Bromley, 1945; Crowhurst, 1969; Fattig, 1945; Melin,
1923; Myers, 1928; Valentine, 1967) have recorded occasional instances of
predation by members of the genus Neoitamus. However, for only two
species, N. angusticornis Loew and N. cyanurus Loew, are there sufficient
data for a prey selection Comparison. As can be seen in Table 2, the ten-
dency of N. vittipes to selectively choose Diptera over other available in-
sects in the habitat is shared by these two other species in widely separated
parts of the world.

In addition to the records for N. cyanurus presented in Table 2, Melin
(1923) noted Diptera and Lepidoptera as prey in Sweden. Diptera also were
recorded as the most common prey in Romania (Ionescu and Weinberg,
1960) and in England by Parmenter (1952). Attwood (1937), although not
collecting prey, noted that N. cyanurus took large numbers of Tortrix viri-
dana \.. (Lepidoptera: Tortricidae.)

Herein is a list of prey taken by N. vittipes. Specific identifications were
made, where possible, but because of the state of knowledge in some
groups, only genus and/or family are included for some specimens. The

VOLUME 84, NUMBER 3 623

Fig. 2. A case of cannibalism in which one Neoitamus vittipes male has captured and is
feeding on another male of the same species.

number of records and sex of the predator are indicated in parenthesis
following the prey record.

DIPTERA, Anthomyiidae: Hylemyia deceptiva Malloch, 7.i1i.79 (2); Asil-
idae: Neoitamus vittipes (Macquart), 18.xii.78 (¢); Chironomidae: Penta-
neurc levidensis (Skuse), 6.i1i1.79 (3); Procladius paludicola Skuse, 6.11.79
(3), 7.iii.79 (3); Choloropidae: indet. A, 6.iii.79 (2); Lioscinella sp., 6.11.79
(2); Ephydridae: indet., 8.iii.79 (2); Lauxaniidae: Poecilohetaerus schineri
Hendel, 9.iii.79 (2 2); Muscidae: Atherigona sp. A, 7.iii.79 (2 2); Coenosia
acuticornis Stein, 7.iii.79 (2), 9.iii.79 (3); Musca vetustissima Walker,
19.xii.78 (3, 2 2), 6.iii.79 (2); Phoridae: Megaselia sp., 6.i11.79 (2); Sep-
sidae: Lasionemapoda hirsuta (de Meij.), 9.iii.79 (2), Syrphidae: Melan-
gyna collata (Walker), 5.iii.79 (2), 7.11.79 (2), 9.iii1.79 (2); Simosyrphus
grandicornis (Macquart), 6.iii.79 (¢). HEMIPTERA-HETEROPTERA, Ly-
gaeidae: Nysius vinitor Bergr., 5.11.79 (3), Miridae: Campylomma lividum
Reuter, 8.iii.79. HEMIPTERA-HOMOPTERA, Aphididae: indet. winged re-
productive, 7.iii.79 ( 2); Cicadellidae: indet., 7.1ii.79(d). HYMENOPTERA,
Braconidae: Aphidiinae, indet., 8.iii.79 (¢); Ichneumonidae: Diplaxon sp.,
7.iii.79 (2). LEPIDOPTERA, Lycaenidae: Zizina labradus labradus (Go-

624 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

dart), 7.iii.79 (2 2). An additional two Diptera and three Homoptera remain
unidentified.

MATING

As with other species of Asilinae studied, there is no courtship by males.
It is probable that encounters between males and females resulted from
searching flights of males, although such data were not collected. This strat-
egy is used by many species of Asilini.

Upon observing a female, the male launches itself towards her, catching
her in flight. Copulation takes place prior to landing, in the male atop female
position. Approximately five seconds after landing, the pair take a tail-to-
tail position which is maintained throughout the copulation period (Fig. 3).
The male rotates his abdomen 70—90° at intermittent intervals which causes
the female’s abdomen to be extended, curved and pulled towards the male.
Rotations occur at a rate of ca. I/s in a short series, followed by periods of
rest. As the mating progresses, the rate of rotation increases to ca. 2/s. The
male’s wings remain folded over his dorsum throughout mating, being
flicked open occasionally, while the female’s wings remain slightly open.
Separation occurs when the male releases his claspers. Both either fly off
immediately or rest for short periods of time, and then fly.

Mated pairs were observed on vegetation (grass stalks and lucerne) at
heights ranging from 2.5 to 35 cm (mean 22 cm). Temperatures at these
heights ranged from 22 to 34°C (mean 26.9°C). Additionally, three mated
pairs were encountered resting on soil (surface temperature 26—28°C) at the
site north of Kingston. Pairs were observed in copula as early as 9:37 AM
and as late as 3:58 PM, with all but one occurring after 12 noon. Total time
in copula for two observed complete matings (i.e., initial contact to sepa-
ration) was 21 and 33 minutes.

OVIPOSITION

According to Melin (1923), female Neoitamus cyanurus deposits its eggs
in ‘‘closed top buds,” in ‘‘small round strobiles,’’ and in the apices of
boughs of alder from which the species also forages. The elongate, laterally
compressed ovipositor is highly suitable for this task.

The ovipositor of N. vittipes is similarly laterally compressed suggesting
that this species also oviposits in suitable locations on vegetation. On one
occasion, the author observed a female probing with its ovipositor, in the
manner described for Machimus callidus (Williston) (Dennis and Lavigne,
1975). The ovipositor was moved from side to side at the tips of lucerne
leaves and inserted within folded unopened leaves at heights of 20 to 36 cm.
No eggs were found, however, when these sites were examined. Whether
or not such sites are utilized is probably immaterial since the lucerne was
cut two days later. It is much more probable, since the species has survived
in this field as well as along the margins of several other lucerne fields in

VOLUME 84, NUMBER 3 625

Fig. 3. Mating pair of Neoitamus vittipes resting on vegetation in a field of lucerne.

the vicinity, that oviposition sites are grasses or weeds. Certainly, such sites
suffice for some other Asilini (Dennis and Lavigne, 1979).

Neoitamus armatus (Macquart)

Occasional specimens of this species were collected at two locations in
South Australia in areas containing native vegetation, Aldinga Beach and
The Ferries-McDonald Conservation Park, south of Monarto. The species
was collected during the period December I1th to January 3lst. Hardy
(1920) recorded this species in New South Wales and Tasmania, from Oc-
tober to January and April.

Specimens were observed foraging from branches of malee at heights up
to 1.7 m. It was also observed resting on fallen branches while feeding on
prey. Prey of one male in the Aldinga Beach scrub community was an
unidentified tiphiid wasp (Hymenoptera), while a female captured a speci-
men of Anthrenocerus australis Hope (Coleoptera: Dermestidae). A male
at Ferries-McDonald Conservation Park was photographed while feeding on
Nysius vinitor Bergr. (Hemiptera: Lygaeidae).

Neoitamus margites (Walker)

South Australian collection records for this species are as follows: Mort-
lock Experiment Station, Auburn (17.i.79, 24.i.79), Ferries-McDonald Con-
servation Park, S of Monarto (11.xii.78), 3.3 km S of Echunga (1.11.79), and

626 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

8 km ENE of Callington (21.x1.78). Neoitamus margites had previously
been recorded from Tasmania and New South Wales by Hardy (1920). Hab-
itats, in which this species was found were dominated by Eucalpytus sp.
with an understory of Avena barbata or native vegetation, e.g., horse pad-
docks and conservation areas.

Foraging flights were initiated from the soil surface, from fallen Eucalyp-
tus branches and from the trunks of gum trees. Flights were at heights of
25-50 mm over distances of 0.15 to 0.3 m. The only prey observed being
taken were four specimens of unidentified winged reproductive ants at Mort-
lock Experiment Station.

ACKNOWLEDGMENTS

I acknowledge, with great appreciation, the taxonomists at CSIRO, Di-
vision of Entomology, who identified the prey insects: D.H. Colless (Dip-
tera), 1. D. Naumann (Hymenoptera), and E. D. Edwards (Lepidoptera). G.
F. Gross is thanked for his identification of Hemiptera-Heteroptera and E.
G. Matthews, for his Coleoptera identification, both of the Entomology
Department, South Australian Museum, Adelaide. I express my apprecia-
tion to Gregory Daniels, Department of Entomology, University of Queens-
land, St. Lucia, for his identification of Neoitamus vittipes as well as other
identifications he is providing. E. D. Symon, Agronomy Department, Waite
Agricultural Research Institute, University of Adelaide, was most helpful
in identifying botanical specimens that characterized the environments in
which N. vittipes was collected. I additionally thank Peter Miles and staff
at Waite Institute for providing me with laboratory space, supplies, and
transportation. I especially thank my wife, Judy, for her time and effort as
an unpaid research associate.

LITERATURE CITED

Attwood, R. W. 1937. ‘“‘Robber flies’’ attacking Tortrix viridana. Entomol. Rec. J. Var. 49:
104-105.

Bromley, S. W. 1945. Insect enemies of the house fly, Musca domestica L. J. N. Y. Entomol.
Soc. 53: 145-152.

Crowhurst, P. S. 1969. Notes on Neoitamus bulbus (Walker) (Diptera: Asilidae). N. Z. Ento-
mol. 4(3): 121-124.

Daniels, G. 1976. Notes on the behavior of the robber fly Promachus interponens Walker
(Diptera: Asilidae). Aust. Entomol. Mag. 2(6): 124-125.

Dennis, D. S. and R. J. Lavigne. 1975. Comparative behavior of Wyoming robber flies II
(Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr. No. 30, 68 pp.

—. 1979. Ethology of Machimus callidus with incidental observations on M. occidentalis
in Wyoming (Diptera: Asilidae). Pan-Pac. Entomol. 55: 208-221.

Fattig, P. W. 1945. The Asilidae or robber flies of Georgia. Emery Univ. Mus. Bull. No. 3,
33 pp.

Hardy, G. H. 1920. Descriptions of Australian flies of the family Asilidae, (Diptera) with
synonyms and notes. Proc. Linn. Soc. N.S.W. 45(Pt. 2): 185-202.

VOLUME 84, NUMBER 3 627

Hobby, B. M. 1931. The British species of Asilidae (Diptera) and their prey. Trans. Entomol.
Soc. South England 6: 1-42.

—. 1933. Supplementary list of the prey of Asilidae (Dipt.). J. Entomol. Soc. South

England |: 69-74.

. 1934. Predacious Diptera and their prey. J. Soc. Brit. Entomol. (2): 35-39.

Ionescu, M. A. and M. Weinberg. 1960. Specii de Asilide (Diptera, Asilidae) din R. P. R.
Colectia Petru Suster. Acad. Repub. Pop. Rom. Stud. Cercet. Biol., Ser. Biol. Anim.
11: 301-313.

Iwata, K. and A. Nagatomi. 1962. Prey of some Japanese asilids (Diptera). Kontyu 30: 87—
100.

Lavigne, R. J. 1979. Notes on the ethology of Efferia argyrogaster (Diptera: Asilidae) in
Mexico. Proc. Entomol. Soc. Wash. 81: 544-551.

Lavigne, R. J. and D. S. Dennis. 1980. Ethology of Proctacanthella leuacopogon in Mexico
(Diptera: Asilidae). Proc. Entomol. Soc. Wash. 82: 260-268.

Lavigne, R. J. and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming
robber flies (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr. No. 18,
61 pp.

Lavigne, R. J., L. Rogers, and F. Lavigne. 1976. Ethology of Efferia benedicti (Diptera:
Asilidae) in Wyoming. Proc. Entomol. Soc. Wash. 78: 145-153.

Melin, D. 1923. Contributions to the knowledge of the biology, metamorphosis and distribution
of the Swedish Asilids. Zoolog. Bidr. Upps. 8: 1-317.

Myers, J. G. 1928. The habits and prey of three Asilidae in New Zealand. Proc. R. Entomol.
Soc. Lond. 2(3): 54-56.

Parmenter, L. 1952. Diptera. Notes on the Asilidae (robber flies). Entomol. Rec. J. Var. 64:
295-299.

Poulton, E. B. 1907. Predacious insects and their prey. Trans. Entomol. Soc. Lond. 1906, pt.
III, pp. 323-409.

Scarbrough, A. G. 1978. Ethology of Cerotainia albipilosa Curran (Diptera: Asilidae) in Mary-
land: predatory behavior. Proc. Entomol. Soc. Wash. 80: 113-127.

Scarbrough, A. G. and A. Norden. 1977. Ethology of Cerotainia albiplosa Curran (Asilidae:
Diptera) in Maryland: diurnal activity rhythm and seasonal distribution. Proc. Entomol.
Soc. Wash. 79: 538-554.

Valentine, E. W. 1967. A list of the hosts of entomophagous insects of New Zealand. N.Z. J.
Sci. 10: 1100-1209.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 628-631

A NEW SPECIES OF SYMMETRISCHEMA POVOLNY
(LEPIDOPTERA: GELECHIIDAE) FROM TEXAS

ANDRE BLANCHARD AND EDWARD C. KNUDSON

(AB) 3023 Underwood, Houston, Texas 77025; (ECK) 804 Woodstock,
Bellaire, Texas 77401.

Abstract.—Symmetrischema kendallorum, n. sp., is described. Imagines,
male and female genitalia, and wing venation are figured. The host is Phys-
alis virginiana Mill. var. spathulaefolia (Torr.) (Solanaceae).

The following new species was discovered in Neuces and Kleberg coun-
ties, Texas. Adults were reared from larvae and pupae found in galls on the
upper stems of Physalis virginiana var. spathulaefolia of the family Solan-
aceae.

Symmetrischema kendallorum Blanchard and Knudson, NEw Species
Figs. 1-7

Description.—Head: Front and vertex light ochreous variably irrorated
with fulvous. Labial palpi: Exceeding front by 3 eye diameters, basal seg-
ments covered by ventro-anteriorly directed divided brush of ochreous
scales, terminal segment light ochreous with 2 black bands, | submedial, |
supramedial, scales closely appressed. Ocelli: Present. Antenna: Simple,
flagellum with 2 scale rows per segment. Proximal scale row black with
narrow whitish base. Distal 19 scale rows are as follows: 1 white 3 black,
| white 3 black, | white 3 black, | white 3 black, | white 2 black.

Thorax: Light ochreous, speckled with fulvous.

Forewing: Ground color light ochreous, largely obscured by extensive
fulvous overscaling and grayish-black patches. Grayish-black patches ap-
pear, under magnification, to consist of black scales having a grayish-white
base and a narrow yellowish-white tip. An extensive grayish black patch
occupies middle % of forewing from costal margin to fold and is interrupted,
in the cell, by 2 fulvous spots, narrowly edged with ground color. A single
row of grayish-black scales extends along dorsal margin, broadening to form
a grayish-black patch near base. Extensive fulvous present along termen
and above dorsal margin along fold. Fringe consisting of 3 rows of scales,

VOLUME 84, NUMBER 3 629

Figs. 1-7. Symmetrischema kendallorum. 1, Male holotype, North Padre Island, Nueces
Co., Texas, 17-I[X-81. 2, Female paratype, same locality, 18-IX-81. 3, Male genitalia, side view
and separate aedeagus, slide ECK 239, paratype from same locality, 18-IX-81. 4, Male genitalia,
| ventral view, slide ECK 232, paratype from same locality, 22-I[X-81. 5, Female genitalia, slide
| ECK 240, paratype from same locality, 18-IX-81. 6, Female genitalia, same slide, greater

magnification, to show ostium bursae, 8th segment and its apophyses. 7, Wing venation of

male paratype, same locality, 25-IX-81, slide A.B. 5050. Segments in Figs. 3, 4, 6, represent
) 0.5 mm; in Fig. 5, 1.0 mm.

630 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

inner 2 rows whitish with broad black apices, outer row ochreous with
deeply dissected tips.

Hindwing: Light fuscous. Fringe concolorous, longer than width of wing.

Abdomen: Light fuscous, usually with some ochreous scaling at caudal
end.

Length of forewing: Male: (N = 20) 4.9-6.0 mm, average 5.5 mm. Fe-
male: (N = 20) 5.0-6.6 mm, average 5.9 mm.

Venation: As in Fig. 7. which represents a male paratype.

Male genitalia (Figs. 3, 4): Sacculus with 2 sets of paired processes; small,
thornlike, outer pair more sclerotized. Median unpaired saccular process
prominent, lightly sclerotized, not reaching tips of paired processes, but at
base covers 2 of inner pair of paired processes. Valvae exhibiting typical
chaetotaxy of genus (row of hairs passing obliquely from tip towards base),
but in addition, have 6 to 8 rows of thornlike projections on middle 4%.
Circumanal membrane prominent, densely scobinate, extending beyond un-
cus laterally and supported by thin rodlike processes. Subscaphial mem-
brane prominent, subtriangular. Along lateral margins of tegumen, near base
of uncus, are complicated curved processes which seem to support the
cephalad margin of uncus. Aedeagus with short lateral process tipped with
numerous fine spines.

Female genitalia (Figs. 5, 6): Bursa lacking signum. Fig. 6 shows simpli-
fied 8th segment sternum and ostium bursae.

Holotype (Fig. 1)—d, Nueces Co., Texas, North Padre Island, ex-larva
Physalis virginiana Mill. var. spathulaefolia (Torr.) (Solanaceae), emerged |
17-[X-81, collected by Roy O. and C. A. Kendall and deposited in the
National Museum of Natural History, Smithsonian Institution, Washington,
Dic

Paratypes.—Nueces Co., Texas, North Padre Island, 24-VIII-79, 1 3, 1
2; same location, 25-VIII-79, 3 ¢ (all collected by A. and M. E. Blanchard
in blacklight traps). Nueces Co., Texas, North Padre Island, ex-larva or
pupa Physalis virginiana var. spathulaefolia, dates of emergence as follows:
25-IX-79, | 3; 2-IX-81, 1 ¢; 3-[X-81, 2 6; 4-IX-81, 1 3; 13-IX-81, 2 d, 1
2; 14-I[X-81, 2 ¢, 1 9; 16-IX-81, 4 9; 17-IX-81, 4 5, 4 2; 18-IX-81, 11
3, 10 2 (Fig. 2); 19-IX-81; 4 6, 5 9; 20-IX-81,3 6,4 2; 21-IX-817 42 ae
9; 22-IX-81, 4 d, 2 9; 23-IX-81, 1 9. Kleberg Co., Texas, Padre Island
National Seashore, 5-IX-81, 1 2 (all collected and reared by Roy O. and C.
A. Kendall).

Distribution.—So far known only from Nueces and Kleberg counties,
Texas:

Life history.—Larvae and pupae found in galls on upper stems of Physa-
lis virginiana Mill. var. spathulaefolia (Torr.) (Solanaceae). Examples were
collected on sand dunes at various locations in Padre Island National Sea-
shore, in Nueces and Kleberg counties.

VOLUME 84, NUMBER 3 631

Remarks.—The genus Symmetrischema was erected by Povolny (1967)
in his notes on Nearctic and Neotropical Gnorimoschemini. Symmetrische-
ma is separated from its nearest relative, Gnorimoschema Busck (1903),
mainly by characteristics of the male genitalia which include tendency to-
ward the formation of a median unpaired process of the sacculus, enlarge-
ment of the terminal portion of the valvae to form irregularly trapezoidal
planes with typical chaetotaxy, strongly developed circumanal membrane,
and a lateral process of the aedeagus. Additionally, the female genitalia
show a tendency towards reduction of the signum. Symmetrischema ken-
dallorum clearly exhibits all of the above characteristics, but it is obviously
distinct from all other known species. The authors take pleasure in naming
this new species for Roy O. and C. A. Kendall, who discovered the larval
food plant and reared nearly all of the type-series.

ACKNOWLEDGMENTS

The authors are extremely grateful to J. F. Gates Clarke, research asso-
ciate of the National Museum of Natural History, for examining the type-
series and critically reviewing the manuscript. Appreciation is also due the
U.S. National Park Service for their cooperation in allowing collecting of
this new species within the Padre Island National Seashore.

LITERATURE CITED

Busck, A. 1903. A revision of the American moths of the family Gelechiidae, with descriptions
of new species. Proc. U.S. Natl. Mus. 25(1304): 767-938.

Povolny, D. 1967. Genitalia of some nearctic and neotropical members of the tribe Gnori-
moschemini (Lepidoptera, Gelechiidae). Acta Entomol. Mus. Natl. Prague 37: 51-127.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 632-635

THREE NEW LEAFHOPPERS (HOMOPTERA: CICADELLIDAE)
FROM COCOS ISLAND'

PAUL S. CWIKLA AND PAUL H. FREYTAG

Department of Entomology, University of Kentucky, Lexington, Ken-
tucky 40546.

Abstract.—Two new species, Scaphytopius (Cloanthanus) biflavus and
Chlorotettix canolaterus, belonging to the subfamily Deltocephalinae, and
one new species, /dona murrayae, belonging to the subfamily Typhlocybi-
nae, are described. All are from Cocos Island, Costa Rica.

Through the kindness of Charles Hogue, Natural History Museum of Los
Angeles County, we were able to examine leafhoppers collected on Cocos
Island. This isolated tropical island lies 500 km southwest of Costa Rica and
445 km northeast of the Galapagos Islands.

Although most of the examined leafhopper fauna of Cocos Island is shared
with the mainland, the three new species may be endemic. All types are
deposited in the Natural History Museum of Los Angeles County, Califor-
nia.

Scaphytopius (Cloanthanus) biflavus Cwikla and Freytag, NEw SPECIES
Figs. 14

Description.—Length: Male 3.2 mm, female unknown.

Structural characteristics: Body relatively slender. Head as wide as pro-
notum. Crown produced anteriorly, median coronal length greater than least
interocular width.

Coloration: Anterior margin of crown with dark brown band. Anterior
Yy of crown yellow, posterior 4% whitish yellow. Face yellowish. Pronotum
whitish yellow with lateral margins brown and irrorate with yellow. Fore-
wing transparent with brownish markings on area of discal cell and around
apical and subapical cells.

Male genitalia: Pygofer broadly rounded in apical 2. Genital plates short,

' The investigation reported in this paper (81-7103) is in connection with a project of the
Kentucky Agricultural Experiment Station and is published with approval of the Director.

VOLUME 84, NUMBER 3 633

———
0.05 mm

| Figs. 1-4. Scaphytopius (Cloanthanus) biflavus. 1, Head and pronotum, dorsal view. 2,

_ Aedeagus and pseudostyle, lateral view. 3, Pseudostyle, ventral view. 4, Style, ventral view.

ee

634 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Ree

Figs. 5-8. 5, 6, Chlorotettix canolaterus. 5, Aedeagus, lateral view. 5a, Aedeagal apex,
dorsal view. 6, Style, ventral view. 7, 8, dona murrayae. 7, Aedeagus, lateral view. 7a, Ventral
aedeagal process apex, ventral view. 8, Style, ventral view.

triangular, with row of setae on lateral margins. Style with apex blunt.
Aedeagal shaft slender, truncate at apex. Pseudostyle (=paraphysis) fused
at base, long, apex tapering to a point in ventral view.

Type.—Holotype 6, Cocos Island (Costa Rica), Wafer Bay, 17-22 April
1975, Cc. i, Hogue.

Discussion.—This species will key to couplet 2 in Linnavuori’s (1959: 73)
key to the Neotropical Scaphytopius (Cloanthanus) Ball; however, a defi-
nite association with any of the keyed species cannot be made. It can be
separated from other Scaphytopius (Cloanthanus) by the two shades of
yellow on the crown and by the slender aedeagus that is apically truncate.

Chlorotettix canolaterus Cwikla and Freytag, NEW SPECIES
Figs. 5-6

Description.—Length: Male 2.9 mm, female unknown.

Structural characteristics: Body slender. Crown short, slightly produced
anteriorly, granulose microsculpturing only on anterior margin.

Coloration: Crown, pronotum, and scutellum whitish yellow. Face yel-

VOLUME 8&4, NUMBER 3 635

low. Forewing whitish yellow with yellow patch on costal area and brown
patch on 3rd apical cell. Legs and thorax brown.

Male genitalia: Style with apex hooked, blunted pointed. Aedeagus sim-
ilar to C. berryi DeLong (DeLong, 1945: Plate 3) but with aedeagal shaft
curved anteriorly, with pair of short dorsally curved processes.

Type.—Holotype ¢, Cocos Island (Costa Rica), Wafer Bay, 24 March
1978, Malaise trap, Station 3, C. Hogue and S. Miller, Steele Exped. 1978.

Discussion.—The generic placement of this species is based on genitalic
similarities. This species is similar to C. berryi DeLong, but can be sepa-
rated from it by the small size and the apical processes which are curved
dorsally but not laterally.

Idona murrayae, Cwikla and Freytag, NEw SPECIES
Figs. 7-8

Description.—Length: Male 2.5 mm, female 2.6 mm.

Coloration: Body pale yellow. Forewing light yellow with white patch on
costal area.

Male genitalia: Style with apex broadly hooked, bluntly pointed. Aedea-
gus similar to J. aptera (Beamer) (Beamer, 1943: Fig. 6D), but 2 spines on
ventral margin of shaft near apex and ventral aedeagal process only bifurcate
at apex.

Female genitalia: Posterior margin of 7th sternum truncate with median
slightly produced.

Types.—Holotype ¢, Cocos Island (Costa Rica), Wafer Bay, 26 March
1978, Malaise trap, Station 3, C. Hogue and S. Miller, Steele Exped. 1978.
Paratype 2, Cocos Is. (500 km SW of Costa Rica), Chatham Bay, Malaise
trap over stream, 9-11 April 1979, R. Silberglied.

Discussion.—The generic characters are like those described by Young
(1952: 62) for Idona DeLong. This species is similar to 7. aptera (Beamer)
but can be separated from it by the two spines on the venter of the aedeagal
shaft and by the apically bifurcate aedeagal process. This species is named
in honor of Evelyn Murray.

LITERATURE CITED

Beamer, R. H. 1943. Notes and descriptions of some new species of Dikraneura (Homoptera:
Cicadellidae) J. Kans. Entomol. Soc. 16: 54-63.

DeLong, D. M. 1945. The genus Chlorotettix (Homoptera: Cicadellidae) in Mexico. Lloydia
(Cinci.) 8: 1-30.

Linnavuori, R. 1959. Revision of the Neotropical Deltocephalinae and some related subfamilies
(Homoptera). Ann. Zool. Soc. “‘Vanamo’ 20: 1-370.

Young, D. A. 1952. A reclassification of Western Hemisphere Typhlocybinae (Homoptera:
Cicadellidae). Univ. Kans. Sci. Bull. 35: 3-217.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 636-650

FIVE NEW SPECIES OF DIACHLORUS (DIPTERA: TABANIDAE)
FROM SOUTH AMERICA WITH A REVISED KEY TO
SPECIES AND NEW LOCALITY RECORDS'”

RICHARD C. WILKERSON AND G. B. FAIRCHILD

Entomology and Nematology Department, University of Florida, Institute
of Food and Agricultural Sciences, Gainesville, Florida 32611; Research
Associates, Florida State Collection of Arthropods.

Abstract.—Five new species of the genus Diachlorus are described and
figured and a revised key to the species is presented. New species are: D.
leticia, D. trevori, D. leucotibialis, D. habecki, and D. heppneri. Males of two
species, D. xynus Fairchild and D. pechumani Fairchild, are described for
the first time. D. aitkeni Fairchild is reduced to a subspecies of D. pechu-
mani. New distribution records are given for D. anduzei Stone, D. bicinctus
Fabricius, D. curvipes Fabricius, D. fuscistigma Lutz, D. jobbinsi Fairchild
and D. podagricus Fabricius.

The genus Diachlorus was reviewed and a species key was provided by
Fairchild (1972). The recent discovery of five undescribed species and
change in status of | species prompts us to report these taxa, describe males
of two species, revise the key, and add new records which extend the geo-
graphic ranges of several species. Species of Diachlorus are primarily char-
acterized by a patch of silvery gray (‘‘pearly’’) pollinosity on their pleura
and a bare, shiny frontoclypeus. Most are small to medium sized yellow
and black species, but a few are primarily black and at least one has a
banded abdomen suggesting that it is a wasp mimic. In addition, many have
apical wing patches and all have multicolored black and yellow and/or white
legs. Eye patterns are quite varied and many resemble Chrysops spp. The
genus ranges from eastern and southern United States (New Jersey to Tex-
as) and the Bahamas to Argentina but is absent in Chile and the Antilles.

' Florida Agricultural Experiment Station Journal Series No. 3207.
* This paper was prepared during the tenure of National Science Foundation grant DEB-
8020081.

VOLUME 84, NUMBER 3 637

4(1).

KEY TO SPECIES OF DIACHLORUS BASED ON FEMALES

Subcallus bare and shiny. Largely shiny black species. Apical
dark wing patch a vertical band which leaves apex hyaline .... 2
Subcallus wholly pollinose. Apical dark wing patch present,
faint, or absent; if outwardly dilute, then abdomen not largely
DICK rest rs cee Sees ee en ee 4
Subcallus with a median silvery pollinose streak. Foretibia whol-
ly black and black haired. Subapical dark wing band concave
outwardly. Abdomen wholly shiny black, occasionally with faint
sparsely pale-haired median triangles on terga 2 to 5 (northern
SreAMmehica, Wrimadagd) 2 eee eee scutellatus Macquart
Subcallus entirely bare and shiny. Foretibia basally white and
white-haired. Subapical dark wing band straight or convex out-
WARY AM Sete Oey a cote c TaN cre ee ee Oe 3
Frons about 2.5 as high as basal width. Abdomen with a mid-
dorsal yellow integumental stripe, or vestige thereof. Terga 2 and
often 3 yellowish or brown laterally. Frontal callus, subcallus
and palp yellowish brown. All femora yellowish to brown, the
hind pair often with subapical brown band (Brazil, (Bahia to
NATORGTOSSONE cee ee Co ee ee ae ee neivai Lutz
Frons about 3.5 as high as basal width, abdomen wholly shiny
black. Frontal callus black, subcallus black or brown, palp yel-
low to black, largely pale-haired. All femora black (Brazil (Para)
TORE ICRU cere Se Te ters ec noe oo oe eee ee xynus Fairchild
Antennal scape longer than basal plate. Abdomen black, the first
2 terga with conspicuous white transverse bands. Mesonotum
black, shiny, with small pale triangles connected to the yellow-
haired notopleural lobes. Scutellum yellow pollinose and yellow
haired. Wing with broad dark costal band to apex, a narrow dark
band covering crossveins at ends of basal cells and small clouds
on fork of 3rd vein (N.E. South America) ..... bicinctus Fabricius
Antennal scape shorter than basal plate. Abdomen rarely black;
if so, then otherwise marked. If wing with costal band, then
lacking Gand al-ends OF basal CElls.~ <<. J. scat aoe cere ee ee 5
Frons about 3.6 as high as basal width, distinctly narrower at
vertex, the callus drop-shaped. Antennae unusually long and
slender, markedly longer than frons, the style as long or longer
than basal plate, the 3rd segment about 3x length of scape. Ab-
domen dull yellowish with a pair of sublateral blackish stripes
covering terga 2 to 6, the median yellowish stripe wider than the
black. Fore- and hindlegs mainly brown to black, midleg wholly

638

OG):

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

pale. Apical wing spot dilute, brownish, often fading out in cell

Pa(S: Brazil), ob csc ese eee bivittatus Wiedemann
Without the above combination of characters ................ 6
Frons narrow, over 3.5 as high as basal width, generally par-
allel sided or widened in the middle (rarely narrowed above) .. 7
Frons broad, less than 2.5 as high as basal width, nearly always
Narrowed. Above... ev rom Horses = pasceadhko ete es Shia ee 19
Mesonotum and scutellum both pollinose, the former unstriped.
Stigma: yellow... brown. or blacks «=o oo nce eee 8

Either mesonotum or scutellum or both with bare shiny areas 9
Median yellow-haired abdominal stripe broad, at least 4% width
of abdomen, often appearing as a series of broadly overlapping
triangles. Distal ends of basal cells distinctly brown bordered
and often with brown streaks in Ist basal and base of Ist pos-
terior cells:(USA: (New Jersey) to Costa Rica), 2522 32-4
ethene tos cot cents AoE EY SRNR oR ea coe ferrugatus Fabricius
Median yellow-haired abdominal stripe narrow, often indistinct,
not over 4 width of abdomen on terga 2 to 4, occasionally wider
on posterior terga. Wings without dark clouds except apical

patch (NorthernsySAinenica): ee eee ae ee fuscistigma Lutz
Mesonotum without pollinose stripes on bare shiny areas. Stigma
intensely black and wing patch strongly black ................ 10

Mesonotum with at least a median yellow pollinose stripe on
bare shiny areas, rarely absent. Stigma yellow to brown, rarely
black... Wing. patchyvamabley .. cis. 6 7 acter coerce eee eee 13
Abdomen translucent yellow on first 2 segments, 3 with diffuse
dark patches on posterolateral corners, 4 largely brown, 5 to 7
shiny black, all black pilose dorsally except yellow hairs laterally
on terga | to 3. All tibiae largely black (E. Colombia) .........
Biss lene: Becorckins, Mognge an Wes cca Mae eae ee ec reas leticia, new species
Abdomen yellow in the middle of first 4 terga and with broad
blackish dorsolateral stripes. Terga 5 to 7 shiny black. Pilosity
yellow in middle and at extreme sides of first 4 terga, black
CISEWIRETE 013 )5 oi 2)..b a)h 5.3 oom apatereity lange enc eas ae 1]
Frontal callus square, narrower than frons. Scutellum shiny
black. Forecoxa, distal % of forefemur, all of mid- and hindfe-
mora and fore- and hindtibiae black, only midtibia and mid- and
hindtarsi white. Pale pollinosity and pilosity whitish (E. Ecua-
don? Brazil (Amazonas)) see cae leucotibialis, new species
Frontal callus higher than wide, filling width of frons. Scutellum
yellow, as is pale pollinosity and pilosity. Forecoxa and mid- and
hindfemoralargely pale’ . ....5... 5.02 esse « Gene eee 12

VOLUME 84, NUMBER 3 639

13(9).

All femora and midtibia pale, at most fore- and midfemora dis-
tally dusky. Fore- and hindtibiae black (E. Ecuador) .........

BONES ERP) BE CR ENE? HBS SUP Bitte cy.n) 5, Sy ogy anasto habecki, new species
Distal 2 to 4% of fore- and hindfemora black. All tibiae black (E.
Benad one yc46 eset Jets nhs ace aed Ko eek trevori, new species
Mesonotum without a pair of anterolateral dark shiny oval spots
above pronotal lobes. Median pale pollinose stripe on meso-
notum widened at level of wing insertions. Frons about 4.5 as
high as basal width. Scutellum yellow margined (Panama to Am-

AZONEDASIONN, Jie Wen eae dus .d i Poel A Teepe curvipes Fabricius
Mesonotum with a pair of anterolateral dark shiny oval spots or
Streaks abo eipronotallobes lech enkises bal eee = 14

Abdomen with a pair of prominent black integumental dorsolat-
eral stripes from Ist through 3rd terga; terga 4 to 7 black with
broad middorsal yellow-haired triangles. Apical wing patch rath-
er dilute, not obvious much posterior to vein M,. Frons about
5x as high as basal width. Scutellum black at base (E. Peru)
forte behind, Stem Fees rae 4. thd sé cise pechumani pechumani Fairchild
Abdomen without prominent black integumental dorsolateral
stripes, at most with faint black spots on terga 2 to 4, and strong-
SipSPotswOMmengaOotoun mele ssl se eticew, Lapse, eee er 15
Abdomen with faint blackish spots dorsolaterally on terga 2 to
4, and small shiny black spots on terga 5 to 7. A pair of diffuse
broad black-haired stripes extends from terga 2 to 4, becoming
narrower posteriorly. Apical wing patch as in pechumani, but
frons slightly narrower and scutellum wholly yellow (Brazil

(Rata rae it. SAR Bie He pechumani aitkeni Fairchild
Abdomen otherwise. Scutellum at least dark at base .......... 16
Median yellow stripe of mesonotum conspicuous, slender. Scu-
tellum wholly black (Brazil (Rio de Janeiro)) .... varipes Rondani
Median yellow stripe of mesonotum faint or, more usually, ab-
sentaSeutellumnarelyawhollyvdark ¢222 .nGa2 0G) ae eae: 17

Dark wing patch rarely evident beyond Ist posterior cell, and
then only in dilute form (Costa Rica to Ecuador, and Brazil

CAmmazonas) ard tos 2as2 Sk Resour s OS. cae esp Jobbinsi Fairchild
Dark wing patch extended along hindborder of wing in full in-
fensityatosanalseclhe 3.6 Se cde tls iar ve Das ee He eRe 18

Hindtibia black and densely black pilose. Last 2 abdominal seg-
ments all black and black pilose. Frontal callus higher than wide,
frontaliinders.0:105:4.(EPer). 60 deen8s heppneri, new species
Hindtibia yellow to brown, yellow to brown pilose. Last 2 ab-
dominal segments at most with shiny black dorsolateral patches,

640

19(6).

9/9)

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

largely yellow pilose. Frontal callus as wide as high. Frontal
index 4.0 or less (Venezuela to Brazil (Mato Grosso)) ........
ot SUSE. RRR sy ad Ss A ss ol nuneztovari Fairchild and Ortiz
Abdomen with a large prominent black median integumental spot
on 2nd tergum, often extending onto 3rd segment. Inner margin

of dark apical wing patch proximal to fork of 3rd vein ......... 20
Abdomen otherwise. Apical wing patch distal to fork of 3rd vein,
Often faint hones MIS USS es Pee Se 1 ae 21

Apical wing patch complete, extensive and intense, its proximal
border curved inward, nearly reaching end of discal cell (North-
Erne S America) .at AT Pa ee ee podagricus Fabricius
Apical spot a broad vertical band, leaving apex of wing hyaline,
its proximal border straight (S. Central Brazil) ... fascipennis Lutz
Mesonotum dark with at most a slender median pale line and
margins and scutellum pale haired. Abdomen with pale-haired
hind marginal bands widened into low median triangles ........ Ip
Mesonotum with a pattern of dark shiny areas separated by 3
pale pollinose stripes. Abdomen with a median pale-haired stripe
or series of (connected! thangles: 2428s ee ee 23
Mesonotum with a slender pale-haired median stripe; sides, pos-
terior margin, and scutellum also pale haired. Abdomen largely
black, the hindmargins of all terga pale, yellow haired, and with
small median yellow-haired triangles. Sides of Ist 2 terga with
yellowish patches (Brazil (Minas Gerais)) ......... altivagus Lutz
Mesonotum without median stripe. Abdomen shining light yel-
low-brown, with anterior parts of terga darkened dorsolaterally.
Hindmargins of all terga pale margined, with median triangles
and pale hairs) (Brazil): Gea n6 She eee eee glaber Wiedemann
Shiny black areas of undenuded mesonotum consisting of a pair
of inverted comma-shaped marks nearly meeting in median line
just anterior to scutellum. Abdomen light yellow on first 3 terga;
succeeding terga each with a dorsolateral pair of black patches
on anterior border. Each tergum with a yellow-haired median
triangle reaching anterior border and broad yellow-haired hind-
margins. Wings with but a faint trace of apical spot in marginal
and ‘submarginal cells (S. ‘Central Brazile s-2 - 22. 3a6e eee

Black pattern of mesonotum more extensive. Abdomen other-
WISE A eek. PUA ROLL, SRT Ca Ee es ae 24
Abdomen light yellow brown, tergum 2 with golden yellow mid-
stripe, terga 3 to S with a gradually darkening and more distinct
dark midstripe, and 3 to 6 with lateral brown streaks. Black

VOLUME 84, NUMBER 3 641

mesonotal pattern consisting of 4 dorsal stripes, the outer pair
curved dorsad and joining before scutellum. Femora yellow, ex-
cept tips of fore- and hindfemora. Wing as in distinctus (Brazil

(alia) eer oe 8 ose ta Pa, Se afflictus Wiedemann
- Abdomen.withontmedian' dank stripe | 7.2) oeiia. 22. ees eee 25
25. Abdomen with a broad diffuse middorsal yellow-haired stripe,

widening on posterior terga. Apical wing spot dilute but well
defined, its proximal border sharp, straight, to fork of 3rd vein.
Mesonotum on disk shiny black with 3 grayish pollinose stripes,
the margins and scutellum yellow haired. Palp dark brown to
blackish,shimyi(SaE: Brazil)ict 308 Jeet. Sees distinctus Lutz
- Abdomen with a narrow, even yellow-haired middorsal stripe.
Apical wing spot dilute and diffuse, seldom reaching fork of 3rd
vein. Median pale stripe of mesonotum broader than laterals,
nsmallysyellowahained> 95248 Be 2 SASS: J See tke Tee 26
26. All femora brown or blackish, at least darker than tibiae of mid-
pair. Abdomen largely brown to blackish (S. Brazil, Paraguay,
AMOS MUNA Meeps ATH Te be a At. o bs eA flavitaenia Lutz
- All femora yellow. Abdomen generally extensively yellow ..... bs |
27. Abdomen yellow to yellowish brown, with a pair of diffuse dark
integumental spots on tergum 2, and terga 4 to 6 generally dark-
ened (S. Brazil, Bolivia, E. Ecuador) .... bimaculatus Wiedemann
~ Abdomen yellow, with more or less distinct dark patches or tri-
angles on extreme sides of terga 4 to 6 (Venezuela, Ecuador,
WOlOMIDI Aa) ee Eee SOs ae Ce RM SE Asta, Os anduzei Stone

Diachlorus leticia Wilkerson and Fairchild, NEw SPECIES
Figs. 1A—C

A slender species resembling fuscistigma Lutz, the mesonotum blackish,
unstriped, scutellum yellowish, all femora and coxae largely pale, all tibiae
blackish. Frons very narrow, abdomen without conspicuous pattern, and
wing with black stigma and strong apical spot.

Female.—Length 9 mm; of wing 8 mm. Head structures as figured. Fron-
tal index 8.2, index of divergence 1.2. Frons pale silvery gray pollinose with
shiny brown callus and diffuse black patch at vertex. Vestiture of short,
sparse, pale yellow hairs, becoming darker around dark patch at vertex.
Integument of subcallus and face dark brown; the subcallus shiny silvery
pollinose, gena pale yellow pollinose. Frontoclypeus mostly bare, shiny
dark brown, lateral area below tentorial pit thinly pale pollinose. Beard
sprase, of pale yellow hairs except for small patch of black hairs and dark
pollinosity above base of palp next to eye. Eye in life green with dark blue
border and horizontal blue mark as figured by Lutz (1913) for D. fuscistig-

642 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 1-5. 1, Diachlorus leticia. 2, D. trevori. 3, D. leucotibialis. 4, D. habecki. 5, D.
heppneri. All holotypes. A, Antenna. B, Palp. C, Frons.

VOLUME 84, NUMBER 3 643

ma. Antennal scape, pedicel, and base of flagellum yellow to orange, re-
mainder of basal plate pale brown becoming dark brown on annuli. Antennal
hairs pale yellow above and below, dark brown laterally. First palpal seg-
ment yellow pollinose, pale yellow haired; 2nd segment with brown integ-
ument showing through sparse shiny yellow pollinosity, hairs pale yellow.
Proboscis blackish, stylets about as long as palp, labella large and wholly
fleshy.

Mesonotum black in ground color. Central area in a broad diffuse trans-
verse band between transverse sutures, and prescutellar area shiny blackish.
Area anterior to transverse suture thinly pale pollinose. Notopleural lobes,
postalar lobes, prescutellum and scutellum thickly yellowish pollinose and
yellow pilose. Pilosity on dark areas sparse, black. Pronotal lobes, propleu-
ral lobes and forecoxa yellow and pale pollinose and pilose. Mesopleuron
including midcoxa infuscated, mesoanepisternum and mesokatepisternum
shiny black in ground color, sparsely covered with pearly pollinosity; re-
mainder of pleura including portions of the mesoanepisternum and meso-
katepisternum near the wing base sparsely yellow pollinose. Hindcoxa pale
yellow and pale yellow pollinose. Basal % of forefemur, all of midfemur,
and all but apex of hindfemur pale yellow. Apical 2 of forefemur and apex
of hindfemur heavily infuscated. Tibiae black; midtibia whitish apically.
Foretarsus black; Ist tarsomere of mid- and hindlegs contrastingly white,
remainder brown. Leg hairs yellow, black or whitish, corresponding to un-
derlying color. Wings hyaline with smoky brown apical patch reaching from
end of vein R,, through fork of 3rd vein and continuing posteriorly more
faintly through distal area of posterior cells. Costal cell brown. Stigma dark
brown, vein R, from the stigma to its termination yellow. Wing veins brown
margined, more intensely so at apices of basal cells. Halter with yellow stem
and dark brown knob. Integument of abdominal terga |, 2, and most of 3
translucent yellow. Tergum 3 with dorsolateral faint, diffuse black patches,
remaining terga shiny black. Dorsal pilosity of numerous short black hairs
except for pale yellow hairs laterally and mesially on tergum | and a quite
small inconspicuous median patch on tergum 2. Abdominal sterna I yel-
low pollinose and yellow haired; remaining sterna black and black haired.

Types.—Holotype, 2, Colombia, Comisaria of Amazonas, 17 km N. Le-
ticia, 25 July 1973, elev. 100 m, Malaise trap, Wilkerson and Young coll.
To be deposited in Florida State Collection of Arthropods (F.S.C.A.).

Paratypes, 2 2, same locality but 26 July 1973. To be retained in collec-
tions of the authors.

Paratypes have wing lengths of 8.5 and 9.0 mm and frontal indices of 8.2
and 9.3. They agree in detail with the holotype, one differs slightly in that
there is faint indication of a pale middorsal abdominal stripe.

Discussion.—Diachlorus fuscistigma is similar to D. leticia but the two
are readily distinguished by leg coloration and abdominal vestiture. The

644 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

midtibia of D. leticia is black and the middle and back first tarsomeres
white. The midtibia of D. fuscistigma is pale yellowish brown and the mid-
tarsus pale yellowish brown, the hindtarsus pale basally darkening to brown
apically. Diachlorus fuscistigma has a broad middorsal yellow-haired ab-
dominal stripe not evident in D. leticia. These two species share the same
eye pattern which, to our knowledge, is unique in the genus.

It is unlikely that D. /eticia is but a color form or subspecies of D. fus-
cistigma since it occurs in the middle of the range of fuscistigma: Surinam;
Brazil (Amazonas, Para, Bahia); Bolivia; Colombia; Ecuador; Peru. Both
have been taken in the vicinity of Leticia, Colombia, and we have seen no
intermediates.

Diachlorus trevori Wilkerson and Fairchild, NEw SPECIES
Figs. 2A—C

A slender species with narrow frons, unstriped mesonotum, largely yellow
coxae and femora, black tibiae and white mid- and hindtarsi. Abdomen
yellow with broad sublateral dark stripes and apex. Wings with dark stigma
and a conspicuous black apical patch.

Female.—Length 10 mm; of wing 9 mm. Head structures as figured. Fron-
tal index 8.4; index of divergence 1.3. Head structures, pilosity and polli-
nosity as in D. leticia. Eye pattern not noted.

Thorax, wings and legs as in D. leticia, except that the scutellum is mostly
yellow in ground color, darkened slightly at base and lateral bare shiny areas
are anterior to transverse sutures.

First abdominal tergum yellow; 2-4 yellow with broad dorsolateral black
stripes formed of large contiguous spots; remaining terga black. Dorsal hairs
largely black but yellow laterally and in central portion of mid-dorsal yellow
stripe. Abdominal sterna 14 yellow and yellow haired; remaining sterna
black and black haired.

Male.—Length 9.5 mm; of wing 9.0 mm. Like female with following dif-
ferences. Eyes bare, holoptic, a central area of poorly demarcated larger
facets occupies about % eye area. Vertex with small black tubercle which
does not reach eye level and bears no visible vestiges of ocelli. Palp greatly
swollen, drop-shaped, shiny black and sparsely yellow haired. Sublateral
abdominal spots less extensive than in female, not contiguous.

Types.—Holotype, 2, Ecuador, Napo Province, Limoncocha, Playaco
River, Malaise trap, 23—28-VIII-1980, Knopf and Dunkle coll. Allotype,
3d, same information as holotype but netted. Paratypes 12 2, 7 d6, same
data as holotype. Holotype and Allotype to be deposited in F.S.C.A.

Female paratypes, length 8.5—11.0 mm; wings 8.5—9.5 mm. Frontal indices
7.0-9.3; indices of divergence 1.2-1.6. Male paratypes, length 9-11 mm;
wings 8-9 mm, paratypes to be retained in collections of the authors.

Discussion.—The sublateral abdominal stripes of the males vary from

VOLUME 84, NUMBER 3 645

very faint, with some lacking the spot on the fourth segment, to nearly as
prominent as in the female.

Diachlorus trevori is similar to two species described here, D. habecki
and D. leucotibialis. Diachlorus leucotibialis has black mid- and hindfemora
and a white midtibia; habecki has wholly yellow midlegs; D. trevori has
largely yellow mid- and hindfemora and a black midtibia. Diachlorus leu-
cotibialis has a black scutellum and grayish yellow thoracic pollinosity in-
stead of the yellow scutellum and yellow thoracic pollinosity of trevori and
habecki.

Named for Trevor Alan Wilkerson, son of the senior author.

Diachlorus leucotibialis Wilkerson and Fairchild, NEw SPEcIES
Figs. 3A—C

A slender species with a narrow frons, extensively black legs but midtibia
and basitarsus and hindbasitarsus contrasting ivory white. Scutellum black
and shiny. Abdomen yellow with broad dorsolateral black stripes and a
black tip, the wing with a black stigma and dilute black apical patch not

_ extending along hind border beyond 3rd posterior cell.

Female.—Length 8 mm; of wing 7.5 mm. Head structures as figured.
Frontal index 4.6, frons slightly widened below. Color of vestiture of head
and appendages as described for D. leticia. Mesonotum as in D. leticia but
yellow integumental and pollinose borders paler, whitish yellow instead of

_ yellow. Scutellum shiny black in ground color with sparse yellow hairs and

pollinosity. Propleuron pale yellow pollinose and sparsely yellow haired.
Remainder of pleural integument and midcoxa black, showing through
sparse silvery gray pollinosity typical of genus. Foreleg with coxa black in
ground color, sparsely yellow pollinose and yellow haired; femur bicolored,
basally yellow, apically black; tibia and tarsus entirely black. Midleg with
trochanter yellow, femur black, tibia and basitarsus white, remainder of
tarsus brown. Hindleg with coxa, trochanter, and extreme base of femur
yellow; rest of femur and all but extreme base of tibia black; basitarsus
white, remainder of tarsus brown. Wing and halter as in D. leticia. Abdom-
inal tergum 1, sides of 2 and a middorsal broad stripe through 5, yellow and
yellow haired; the remainder black and black haired. Sterna |, 2, and large
median areas of 3—S yellow and yellow haired; lateral areas of 3-5 and
remaining segments black and black haired.

Types.—Holotype, 2, Ecuador, Napo Province, Primavera, netted, 26-
VIII-1980, Dunkle and Knopf coll. To be deposited in F.S.C.A.

Paratypes: | 2 same data as holotype; 8.5 mm long, wing 8.0 mm, frontal
index 4.9. 2 2, Brazil, Amazonas, Manaus, Reserva Ducke, VI-1976, L.
Albuquerque; length 6.5 and 7.0 mm, of wing 7.0 and 7.5 mm, frontal indices
4.3 and 4.9. All paratypes agree well with the holotype though the 2 from
Brazil are paler yellow than the Ecuadorian specimens.

646 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Discussion.—Diachlorus leucotibialis is quite similar to D. trevori and D.
habecki. The differences between these three are discussed under D. tre-
VOri.

Diachlorus habecki Wilkerson and Fairchild, New Species
Figs. 4A—C

A slender species with a narrow frons, unstriped mesonotum, and legs
mostly yellow, but with the fore- and hindtibiae black, the foretarsus black
and the mid- and hindtarsi white. Abdomen with large sublateral black spots
on terga 2 to 3 or 4, the last 3 segments of the abdomen black.

Female.—Length 8 mm; of wing 8 mm. Head structures as figured. Fron-
tal index 6.8. Frons slightly widened in middle. Color of integument and
vestiture of head structures as in D. leticia except that palp is darker, Ist
segment concolorous with 2nd black, not orange brown as in leticia. As
noted from figures, basal plate of antenna of habecki not markedly widened.
Mesonotum, scutellum, pleura, wing, and halter all as in D. leticia. Coxae,
femora, and midtibia yellow and yellow haired. Foretibia and tarsus and
hindtibia black and black haired. Mid- and hindbasitarsi white and white
haired, remainder of tarsus brown.

Abdominal terga 1-4 yellow and yellow haired with broad dorsolateral
black and black pilose stripes, broadest on tergum 2, narrower on 3, and
obsolete or absent on 4. Abdominal segments 5—7 shiny black and black
haired. Sterna I-4 yellow and yellow haired.

Types.—Holotype, 2, Ecuador, Napo Province, Limoncocha, Playaco
River, 23—28-VIII-1980, Malaise trap, Knopf and Dunkle coll. To be depos-
ited in F.S.C.A. Paratypes, 3 2, same data as holotype. The paratypes are
7.5-9 mm long; wings 7-8.5 mm with frontal indices of 6.5—6.7. One has a
parallel-sided frons, another is slightly widened in the middle, another slight-
ly wider below than above. To be retained in the collections of the authors.

Discussion.—Diachlorus habecki is very similar to D. leucotibialis and
D. trevori. The differences between these three are discussed under D.
trevori.

Named in honor of Professor Dale Habeck of the Department of Ento-

mology and Nematology, University of Florida, in grateful acknowledgment

of numerous kindnesses to both of us.

Diachlorus heppneri Wilkerson and Fairchild, NEw SPECIES
Figs. 5A—C

A slender species similar to D. nuneztovari, with yellow palpi, striped
mesonotum, mostly yellow scutellum and legs mostly yellow except for
black fore- and hindtibiae and foretarsus. Wings with a distinct dark apical
patch extending in full intensity along hind border to anal cell. Abdomen
orange with a sharply black and shiny tip.

VOLUME 84, NUMBER 3 647

Female.—Length 9.5 mm; of wing 10.0 mm. Head structures as figured.
Frontal index 5.4. Frons nearly parallel sided, widened slightly in middle
and above. Frons grayish yellow pollinose with shiny dark brown callus and
diffuse black patch at vertex. Vestiture of short sparse pale yellow hairs
becoming black and numerous at dark patch at vertex. Subcallus, gena, and
lateral area of frontoclypeus silvery gray pollinose. Remainder of fronto-
clypeus shiny, dark brown. Scape and pedicel yellow, outer aspect brown
| haired, remainder yellow haired. Flagellum brown, annuli slightly darker
_ than basal plate. Palp yellow and yellow haired. Stylets about as long as

palp, proboscis brown and brown pollinose, labella large and wholly fleshy.
' Beard of sparse pale yellow hairs.

Mesonotum yellow and yellow pollinose with broad median shiny black
stripe reaching to prescutellar area and projecting laterally to transverse
suture and back to wing base. Anterior to transverse sutures are a pair of
subshiny dorsolateral spots formed of dark integument showing through
sparse pollinosity which are separated from median dark stripe by dense
pale pollinosity. Scutellum shiny dark orange, sparsely yellow haired and
with dark spot at base. Propleuron yellow pollinose and yellow haired. Re-
mainder of pleura with black integument showing through silvery gray pol-
linosity. Area below wing base sparsely yellowish pollinose. Foreleg with
. coxa and femur yellow; tibia and tarsus black. Midleg with coxa black;
femur and tibia yellow; basitarsus white, remainder of tarsus pale brown.
| Hindleg with coxa and femur yellow; tibia black; basitarsus white, remain-
\ der black. Leg hairs concolorous with integument. Halter yellow. Wing with
| distinct smoky brown apical patch beginning at end of vein R,, continuing
| through fork of 3rd vein and posteriorly in broad band through Sth posterior
\ cell. Costal cell tinted yellow, Ist basal cell with broad anterior brown
streak, and stigma pale brown.

Abdominal terga I—5 dark yellow, 6 and 7 black. Tergum | and 4 and 5
yellow haired; 2 and 3 black haired with broad median yellow haired stripe;
6 and 7 black haired. Sterna 1-4 yellow and yellow haired; 5 dusky and
_ yellow haired; 6 and 7 black and black haired. Five yellow anterior segments
\ thinly yellow pollinose, 2 terminal black segments shiny.
| Types.—Holotype, 2, Peru, Madre de Dios, Rio Tampopata Reserve, 30

air km SW Pto. Maldonado, 290 m, 6—10-XI-1979, Subtropical Moist Forest,
\J. B. Heppner coll. To be returned to National Museum of Natural History,
Washington, D.C. (USNM). Paratypes 2 2, same locality as holotype, one,
21-25-XI-1979, the other 16—20-XI-1979. To be returned to USNM.

Paratypes are 9.5 and 9.0 mm long and have wing lengths of 9.5 mm.
Frontal indices are 5.0 and 4.8.

Discussion.—Diachlorus heppneri is quite similar to a sympatric species,
D. nuneztovari. Diachlorus nuneztovari has a slightly broader frons, pale
hindtibia, and a distinct broad abdominal yellow-haired stripe reaching

648 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

through terga 6 or 7. Diachlorus jobbinsi is also similar but has a pale
hindtibia and less extensive wing coloring, the apical patch not extended
posteriorly, and the first basal cell not infuscated brown.

Named in honor of Dr. John B. Heppner, lepidopterist of the National
Museum of Natural History and the collector of this and many other inter-
esting tabanids.

Diachlorus pechumani Fairchild

This species was based on 4 females from Quince Mil, Prov. Cuzco, Peru.
An additional female collected by Wilkerson and Young is from Colombia, |
Amazonas, 17 km W. of Leticia, 22-VII-73. It is paler than the available
paratype, the dark abdominal stripes being only a little darker than the
darkest available paratype of D. aitkeni, while the dark pilose areas on the
hindlegs are less dark and less extensive than on the paratypes of D. pechu- |
mani. Three additional females from Surinam, though not in perfect con- |
dition, are nearly as dark as the Colombian example cited above, although |
geographically closer to the type-locality of aitkeni. Further specimens from |
the type-locality of aitkeni are a male and a female taken by D. G. Young
in July-Aug. 1974, and 4 females taken in the forest canopy by I. S. Go- |
rayeb, and J. A. Rafael, 1-6-VIH-1981. Surprisingly, the species proves to |
be sexually dimorphic, the male being even darker than pechumani, the
female with almost entirely yellow legs and abdomen. A description of the |
male follows.

Male.—Length 9 mm, of wing 8 mm. Eyes bare, holoptic, bearing a poorly |
demarcated area of greatly enlarged facets in middle, covering about % or |
less of whole eye area. Vertex with a black subshiny tubercle beset with ;
long black hairs and reaching to eye level. Subcallus and frontal triangle
brown in ground color, the former silvery pollinose, the latter bare and
shiny. Frontoclypeus inflated, black, shiny, with sunken orange pollinose
lower median subtriangular area. Cheeks thinly gray pollinose, sparsely pale |
haired. Antenna as in female but basal plate and style more slender. Palp
inflated, banana-like, black and shiny, the Ist segment orange, contrasting |
with the 2nd, both sparsely haired. Proboscis blackish, theca sclerotized, |
labella soft, exceeding length of palp.

Thorax marked as in female, though dark areas more extensive and less |
pollinose, so that thorax appears more dark and shiny. Scutellum blackish, |

cell. Stigma yellow. Forecoxa yellow, mid- and hindcoxae lightly infuscated §
on outer surface. Forefemur apically black, basally pale, bicolored. Mid-

pale basal '/5 and extreme apex. Foretibia and tarsus black to dark brown. |
Midtibia white and white pilose, as is tarsus except for dusky tip. Hindtibia

VOLUME 84, NUMBER 3 649

about 4% pale basally, otherwise black, tarsus white except for a dusky tip.
Vestiture follows integumental color. Halter with yellow stem and brown
knob.

Abdomen translucent yellowish, Ist segment brownish on posterior mar-
gins, 2nd to 6th terga with broad blackish patches which form dorsolateral
stripes on terga 2 and 3 but reach lateral margins on terga 4 to 6. Terga 2
and 3 also have extreme sides somewhat infuscated. Median yellow strip

_ widened to form complete or nearly complete pale hindmargin on segments

4 to 6. Beneath abdomen is translucent on first few segments, except for
chalky white patch at sides of Ist segment. Posterior segments yellow, in-

creasingly brown infuscated to nearly black terminally. Vestiture sparse, of
long hairs, and integument largely shiny both above and below.

Discussion.—Plesiotype 6, Brazil, Para, Belem, APEG forest, 29-VII to

| 6-VIII-1974, D. G. Young coll., flight trap.

The additional material here reported and the appearance of the male

_make it necessary to reduce D. aitkeni to a subspecies of D. pechumani
since the latter has page precedence and was the species illustrated.

Diachlorus xynus Fairchild

Two additional females from Colombia, Meta, El Porvenir and Carima-
-gua, II and III 1979, Wilkerson coll., agree closely with paratypes from
Surinam and eastern Colombia in having the wings more lightly marked and
in being smaller than Peruvian examples. Two male examples from Brazil,
Mato Grosso, Rio Aripuana, Humboldt, 59°27’W., 10°10’S., 12—16-VIII-74,
_D. G. Young coll., flight trap, are unfortunately not accompanied by fe-
/ males. We believe, however, that they belong with xynus rather than scu-
| tellatus, as they lack the median pollinose stripe on the subcallus and have
) very dark costal cells, irregular proximal margin of apical spot, and strong
clouds around apices of basal cells and discal cell. Both specimens are
almost entirely shiny black and sparsely black-haired, showing pale vesti-
| ture or integument only on the four posterior basitarsi, extreme bases of
| four posterior tibiae, and small tufts of dark golden hairs on notopleural
lobes and disk of scutellum. The halteres have blackish stems and orange
heads. The eyes are bare with large facets confined to a small area in the
front of the eye, the facets not much enlarged nor demarcated from the
small facets. Although holoptic, the eyes are in contact for only a short
a and there is a prominent brownish subshiny frontal triangle and
a large black and hirsute tubercle at vertex rising well above eye level. The
antennae are like those of the female, though more slender, the palpi greatly
inflated, both segments shiny black and the 2nd with long outstanding hairs.
Wings are like those of females from eastern Peru, heavily marked, and
with the basal cells strongly brownish tinged.

|
|
|

650 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

NEw DISTRIBUTION RECORDS

Diachlorus anduzei Stone. 1 2, Ecuador, Napo Prov., Limoncocha, 18—25-
V-76, T. E. Rogers coll., flight trap.

Diachlorus bicinctus Fabricius. 1 2, Bolivia, Dept. Beni, Rio Itenez at
mouth of Rio Baures, 30-IX-1964, J. K. Bouseman coll. 2 9 Brazil, Mato
Grosso, Rio Aripuana, Humboldt, 12—16-VIII-1974, D. G. Young coll.,
flight trap. These specimens are darker than those from Surinam, with
more extensive and darker wing spots at ends of basal and discal cells
and at fork, while the costal dark band is extended in a faint apical patch
along the hind border to the 3rd posterior cell.

Diachlorus curvipes Fabricius. 3 2, Bolivia, Dept. Beni, Rio Mamore and
Rio Itenez, Aug., Sept. 1964, Bouseman coll.

Diachlorus fuscistigma Lutz. 1 ° , Bolivia, Rio Itenez, Pampa de Meio, 11-
13-[X-1964, J. K. Bouseman coll.

Diachlorus podagricus Fabricius. 2 2 , Brazil, Para, Mocambo, (APEG for-
est), I-VI and 4-6-VIH-1981, flight trap in forest canopy | 2, 29-VII to
6-VIII-1974, flight trap, D. G. Young coll.; 4 2, Brazil, Amazonas, Re-
serva Ducke near Manaus, 24, 25, 29-VIH-1981 and 14-VIII-1981, arboreal
flight trap; 2 2, Brazil, Amazonas, Manaus, Parque Laranjeiras, 29-VII-
1981, arboreal flight trap and | 2, Brazil, Para, Belem-Brasilia highway,
km 94, 10-I-1962, in tree. The species seems almost wholly arboreal. The |
recent specimens were taken in an arboreal flight trap designed by J. A. |
Rafael and I. S. Gorayeb.

Diachlorus jobbinsi Fairchild. 2 2 , Brazil, Amazonas, Lago Amana, 10-[X-
1979, at light, Robin Best coll. In coll. INPA, Manaus.

ACKNOWLEDGMENTS

We express our appreciation to D. G. Young and D. H. Habeck for their |
kind help in reviewing this manuscript. |
LITERATURE CITED

Fairchild, G. B. 1972. Notes on neotropical Tabanidae (Diptera) XIII. The genus Diachlorus |
O. S. Fla. Ent. 55(4): 219-229.
Lutz, A. 1913. Tabanidas do Brazil e alguns Estados vizinhos. Mem. Inst. Oswaldo Cruz (Rio |
de J.) 5(2): 142-191.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 651-652

BooK REVIEW

The World of the Tent-Makers. A Natural History of the Eastern Tent
Caterpillar, by Vincent G. Dethier. The University of Massachusetts
Press, Amherst. 1980. 148 pp.

This delightful book is a popular account of the biology of the Eastern
Tent Caterpillar. The book is very much in the tradition of the author’s
previous To Know a Fly and Howard Evans’ Wasp Farm and Life on a
Little Known Planet. The story line follows the fate of a clutch of tent
caterpillar eggs from the dormant eggs in midwinter to the emergence of the
adults and the laying of the next generation of eggs the following summer.

The book begins with a discussion of how the overwintering first-instar
larvae in the egg shells survive the sub-freezing temperatures and how they
measure the gradually lengthening days of spring and hatch after the proper
period of light and warmth. The newly hatched young are followed through
the first stages of their colonial life, and in the process, the topics of pher-
omone trails, social behavior, silk production, movement, and orientation
are presented. One of the more interesting aspects of their loose social life,
I think, is the heterogeneity of the larvae in terms of their activity. Dethier
discusses admirably the paradox of a social animal that depends on active
and slightly asocial individuals to keep the colony well fed and healthy. As
early summer progresses, Dethier follows the caterpillars through their
molts, briefly touching on the hormonal control of metamorphosis. The se-
lection of foodplants and the relationship between chemicals produced by
the plant and the food-seeking behavior of the larvae are reviewed. The
nemesis of death is also portrayed: the predators, parasitoids, and patho-
gens. As last-instar larvae, they leave the nest and wander. As random as
it looks to those of us who have watched it, their search for a pupation site
is not haphazard, and the discussion of the search gives Dethier a chance
to elaborate on how the larvae orient. Once the larva has located a pupation
site, the author talks about the construction of the cocoon, the molt into the
pupal stage, and hints, too briefly I thought, at the reorganization of the
body tissues of the larva into the adult stage. The book concludes with the
emergence of the adult and the laying of the next generation of eggs.

The book is filled with interesting details and tidbits of information, all of
it presented in a light and easy manner. The writing style, at times, tends
to become a bit romantic, but never enough to irritate. The author’s back-
ground as a physiologist is apparent as physiology and neurobiology tend
to dominate the ecology and evolution. Perhaps a bit more information on
the coevolutionary relationship between the host plants and the caterpillars

652 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

might have been welcome. Dethier’s list of references has one important
omission: John G. Franclemont’s treatment of the genus Malacosoma and
the beautiful color plates in The Moths of North America North of Mexico,
fascicle 20.1, 1973.

These ritual carpings do not detract, however, from the enjoyability and
quality of the book. It’s the sort of book I would buy to give to a relative
who wondered what I’m doing and why I’m doing it.

Robert W. Poole, Systematic Entomology Laboratory, IBII, Agricul-
tural Research Service, USDA, % National Museum of Natural History,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
84(3), 1982, pp. 652-653

Book REVIEW

Taxonomic Studies of the Encyrtidae with the Description of New Species
and a New Genus, by Gordon Gordh and V. A. Trjapitzin. University of
California Publications in Entomology. 1981. Vol. 93, 55 pp., 6 pls. of 23
figs. Cost: $7.00 (paperback).

This paper discusses 13 North American genera of Encyrtidae. One genus
and five species are described as new, three lectotypes are selected, one
tribe and three genera are placed in synonymy, a key to four genera of one
subtribe is given, and keys to the species of five genera are provided.

Basically the paper is divided into a brief introduction, a taxonomic sec-
tion, a summary, a bibliography, and six plates of figures. Of these sections,
possibly the introduction and summary are the most important to the ma-
jority of readers, and it is herein that the reader might be confused. The
authors state that ‘‘there are no keys available to the genera’’ of North
American Encyrtidae (first paragraph), but then they cite two recent generic

keys for this region (third paragraph). The summary states that the paper |

treats ‘“‘the chalcidoid family Encyrtidae as it is represented in North Amer-
ica,’ but actually it treats only 13 of 131 genera. These statements, I feel,
are possibly just misworded, but the abstractor or reader who relies solely
on these sections for information will be misinformed or misled as to what
is accomplished or presented in the main body of the text. Additionally, the
summary incorrectly lists the taxa Tetralophiellus brevicollis Ashmead and
Cerapterocerus floridanus Ashmead as ‘‘Generic Redescriptions,’’ when
the authors really meant new combinations in the genus Tetracnemus (brev-
icollis is not cited as such in the text or the summary).

|
q
'
|
|

VOLUME 84, NUMBER 3 653

This paper will have limited appeal to most readers, as it mainly consists
of descriptive and taxonomic notes on a few selected genera. Keys to
species of five genera may prove useful to some readers. There are 23
figures, unfortunately without a figure legend, so that the identity of each
must be found in the text. (On page 46, the reference to Fig. 24 should read
Fig. 23.)

On a personal note, I do not care much for the format adopted for this
volume by the University of California Press. It has apparently used a cam-
era-ready, typed manuscript (unjustified right margin) with wide-spaced
lines. By my estimate, the older style issues contain at least twice the num-
bers of words per page as the new, and thus require half the number of
pages as the presently used format. At the same time, I believe the text of
the older issue is more easily readable, with the various sections and sub-
sections being quite distinct. I hope the press will consider returning to its
previous format. Additionally, it has been my impression that the U. C.
Publications in Entomology have been used for comprehensive taxonomic
and biological treatments of subjects rather than fragmentary works such as
the one discussed here. I believe this series should be reserved for compre-
hensive works, and that other papers should be referred to the appropriate
journals.

E. E. Grissell, Systematic Entomology Laboratory, IBII, Agricultural
Research Service, USDA, % National Museum of Natural History, Wash-
ington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.

84(3), 1982, pp. 654-660

MEMBERSHIP LIST OF THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

Honorary Members

Muesebeck, C. F. W., 1920
(1955), Honorary President
(1970)

Poos, F. W., 1923 (1966)

St. George, R. A., 1918 (1975)

Gurney, A. B., 1936 (1979)

Regular and (*) Life Members

Abercrombie, J., 1975
Adams, J. R., 1963
Addington, R. J., 1964
Adler, V. E., 1961
Agafitei, N. J., 1981
Agarwal, A. K., 1977
Aitken, T. H. G., 1957
Albuquerque, L. P. de, 1978
Alexander, J. L., 1971
Al-Gboory, I. J., 1979
Altman, R. M., 1964
Anderson, D. M., 1954
Anderson, L. D., 1944
Anderson, W. H., 1937
App, B. A., 1952
Armstrong, E., 1977
Arnaud He 1955
Arnett, R. H., Jr., 1980
Arnold, S. L., 1975
Ashlock, P. D., 1958
Austin, D. F., 1977
Autry, H. V., 1961
Baker, E. W., 1944
Baksh, E., 1980

Ball, G. E., 1948
Barber, K., 1981
Barker, Z. A., 1964
Barnes, J. K., 1979
Barnett, D. E., 1976
Barnum, A., 1956
Barr, A. R-. 195i]

Barr, W. F., 1958
Barrows, E. M., 1976

April 1, 1982

Barry, C., 1963
Baumann, R. W., 1973
Beal, R. S., Jr., 1958
Beaver, O., 1974
Bechtel, R. C., 1960
Beck, T. R., 1979
Becker, E. C., 1951
Bell, R. T., 1955
Bellinger, R. G., 1972
Berg, €; OF 1975
Bergman, P. W., 1966
Berner, L., 1949
Benny. Rae sal o72
Beyer, W. N., 1977
Bezark, L., 1974
Bicha, W., 1981
Bickel, D. J., 1981
Bickley, W. E., 1949*
Billings, S. C., 1952
Bissell, T. L., 1941
Blanchard, A., 1968
Bodri, M. S., 1979
Boese, J. L., 1977
Boesel, M. W., 1973
Boettcher, R. A., 1955
Bohart, R. M., 1944
Bohnsack, K. K., 1958
Borchelt, R., 1979
Bouseman, J. K., 1975
Bowen, T. W., 1980
Brassard, D. W., 1978
Braum, B. H., 1969
Brigham, W. U., 1975
Brown, F. M., 1975
ebony, lal. 14 IMT)
Brown, R. L., 1979
Bryce, G. K., 1977
Bueno-Soria, J., 1977
Burditt, A. K., 1965
Burger, J. F., 1973
Burke, H. R., 1981
Burks, B. D., 1938
Burks, K. O., 1950

VOLUME 84, NUMBER 3

Burnett, J. A., 1975
Burns, J. M., 1975
Butler, L., 1966
Calabrese, D. M., 1980
Campbell, J. M., 1966
Carbonell, C. S., 1946
Carestia, R. R., 1981
Carlson, R. W., 1970*
Caron, D. M., 1971
Carrington, J. H., 1943
Carroll, J. F., 1977
Carver, M., 1980
Casdorph, D. G., 1977
Gave, R., 1977
Chang, L. W. H., 1976
Chapin, J. B., 1973
Charpentier, P. O., 1962
Chilson, L. M., 1977
Chittick, H. A., 1946
Clark, W. E., 1975
Clark, S. T., 1981
Clarke, J. F. G., 1936
Cochran, D. G., 1981
Coffman, C. C., 1980
Cole A. Be 1977,
Cole, F. R., 1963
Collins, M. S., 1977
Connell, J. G., 1974
Cook, D. R., 1952
Cooper, C. M., 1976
Cooper, K. W., 1955
Coovert, G. A., 1977
Copeland, T. P., 1977
Corkum, L. D., 1979
Coulson, J. R., 1961
Covell, C. V., 1971
Crabill, R. E., Jr., 1968
Craig, G. B., 1954
Crooks, E. E., 1964
Cross, H. F., 1954
Curéié, B., 1982
Currado, I., 1978
Curtin, T. H., 1956
Darling, D. C., 1981
Darsie, R. F., 1949
Daum, R. J., 1969
Davidson, J. A., 1957
Davidson, R. L., 1975
Davis, D. R., 1961
Davis, I. 'Ge, 1952
Davis, M. M., 1979

655

DeBold, K. J., 1981
Deeming, J. C., 1974
Delgado, H., 1979
DeLong, D. M., 1936
Denning, D. G., 1966
Dennis, S., 1976
Deyrup, M. A., 1979
Dicke, F. F., 1933
Dodds, P. J., 1980
Donley, D. E., 1955
Donnelly, T. W., 1962
Doria, J. J., 1977

Dos Passos, C. F., 1947
Downing, W. D., 1975
Downs, W. G., 1981
Dozier He lee. Loses
Drummond, R. O., 1954
Duckworth, W. D., 1961
Duffield, R., 1978
Duret, J. P., 1969
Dutky, S. R., 1953
Eads, R. B., 1963
Edmiston, J. F., 1975
Edmunds, G. F., Jr., 1951
Eikenbary, R. D., 1979
Elder, A. S., 1973
Elias, M. K., 1972
Emerson, K. C., 1952
Emsley, M. G., 1970
Enns, W. R., 1960
Erwin, T. L., 1972
Evans, H. E., 1948
Evans, W. G., 1957
Evenhuis, N. L., 1980
Fairchild, G. B., 1939
Fales, J. H., 1944
Faran, M. E., 1977
Fedde, G., 1962
Fedde, V. H., 1977
Fennah, R. G., 1941
Ferguson, D. C., 1969
Field, G., 1966

Field, W. D., 1965
Fisher, E. M., 1977
Fisk, F. W., 1968
Flechtmann, C., 1968
Flint, O. S., Jr., 1961
Floore, T. G., 1967
Fluno, J. A., 1957
Foerster, W. P., 1977
Foote, B. A., 1958

656

Foote, R. H., 1950
Forattini, O. P., 1956
Foster, J. R-, 1953

Foxe 131936
Franclemont, J. G., 1947
Freeman, J. V., 1981
Freytag, P. H., 1979
Friedberg, A., 1979
Froeschner, R. C., 1961
Gagné, R. J., 1966*
Garcia, C. M., 1981
Gentry, J. W., 1958
Gerberg; E- Je, 1953
Gerberich, A. G., 1981
Gerdes, C. F., 1976
Gibson, L. P., 1981
Giles, F. E., 1981

Gill, G. D., 1958
Gimpel, W. F., Jr., 1970
Glick, J. I., 1979
Godfrey, G. L., 1971
Gonzalez, R. H., 1974
Gordh, G., 1975
Gordon, R. D., 1968
Gorham, J. R., 1974
Gotwald, W. H., Jr., 1977
Grabowski, W. B., 1970
Grant, C. D., 1948
Greenbaum, H. N., 1968
Greenfield, M. D., 1979
Gregg, R. E., 1945
Gressittq Jae 19455
Grimes, L. R., 1981
Gnissell, E. E., 1979
Grogan, W. L., Jr., 1974
Grothaus, R. H., 1981
Gunther, R. G., 1981
Habeck, D. H., 1957
Hacker, J. D., 1971
Hagen, K. S., 1949
Haines, K. A., 1952
Hamid, A., 1976
Hamman, R. E., 1968
Hannemann, H.-J., 1979
Harbach, R. E., 1972
Harding, W. C., Jr., 1955
Hardy, A. R., 1974
Harman, D. M., 1966
Harmston, F. C., 1940
Harper, PS P1977
Harris, S. C., 1979

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Harrison, B. A., 1976
Harrison, F. P., 1954
Haskins, C. P., 1945
Hawkins, L. S., Jr., 1970
Hayes, D. K., 1970
Heitzman, R. L., 1980
Hendrickson, R. M., Jr., 1978
Henry, Cas slo7s
Henty, Redeenl975
Heppner, J. B., 1974
Hermans > Hearel96s
Herring, J., 1963
Hespenheide, H. A., III, 1981
Hevel, G. F., 1970
Higgins, H. G., 1948
Hodges, R. W., 1960
Hoebeke, E. R., 1980
Hoelscher, S. M., 1980
Hoffmann, C. H., 1945
Hoffmann, W. E., 1944
Hoogstraal, H., 1946
Hopla, C. E., 1961
Hopper, H. P., 1978
Fone; Jee elo
Howden, H. F., 1948
Huang, Y.-M., 1968
Hubbard, M. D., 1976
Hudson, B. N. A., 1980
Hull, W. B., 1949
Hung, A. C. F., 1981
Hunter, P. E., 1961
Husband, R. W., 1973
Inai, E., 1980

Ingles, R., 1975

Irwin, M. E., 1976
Ivie, M. A., 1981
Jackson, D. L., 1966
Jacobs, T. T., 1981
Jakob, W. L., 1977
Johnson, M. D., 1973
Johnson, N. F., 1980
Johnson, P. T., 1951
Jones, J. C., 1980
Jones, R. H., 1955
Joseph, S. R., 1957
Kasten (GH al979
Kennedy, J. H., 1977
Kethley, J. B., 1974
Kimball@aPl952
Kimsey, L. S., 1976
Kingsolver, J. M., 1963

VOLUME 84, NUMBER 3

Kirchner, R. F., 1981
Kissinger, D. G., 1955
Kitayama, C., 1974
Kiteley, E. J., 1971
Kittle, P., 1975

Knight, K. L., 1944
Knipling, E. F., 1946
Kniser, S. G., 1981
Knisley, C. B., 1981
Knutson, L. V., 1963*
Kormilev, N. A., 1941
Kosztarab, M., 1978
Krafsur, E. S., 1968
Kramer, J. P., 1957
Krombein, K. V., 1941*
Kuenzel, N. T., 1975
Kurcezewski, F. E., 1970
Lager, T. M., 1976
Lamb, N. J., 1978
Lambdin, P. L., 1974
ampert, 2.1, 1976
Lanchester, H. P., 1957
Langford, G. S., 1924
Lassmann, G. W., 1946
Lavigne, R., 1973
Lewis, P. A., 1974
Lewis, R. E., 1958
Lewis, S. E., 1974
Wiens Ue Ca) 1967

Lin, N., 1978

Linam, J., 1982
Linkfield, R. L., 1959
Pipes, J. E., 1974
Lipovsky, L. J., 1949
Ludvik, G. F., 1978
Lund, H. O., 1952
Lyon, R. J., 1961
Mabry, J. E., 1954
MacLachlan, W. B., 1980
Magner, J. M., 1953
Maier, C. T., 1976
Main, A. J., Jr., 1965

Maldonado-Capriles, J., 1947

Mallack, J., 1957
Mallis, A., 1977
Mangan, R. L., 1977
Manglitz, G. R., 1956
Mankins, J. V., 1976
Mari Mutt, J. A., 1976
Marsh, P. M., 1960
Marshall, S., 1982

Masner, L., 1969
Mason, T. L., Jr., 1977
Mason, W. R. M., 1970
Mathis, W. N., 1976
Matta, J., 1978

Mayor, A. J., 1981
McCabe, T. L., 1977
McCafferty, W. P., 1968
McComb, C. W., 1976
McDaniel, B., 1964
McFadden, M. W., 1956
McGann, D. R., 1979
McIntyre, T., 1957
McMurty J. A., 1981
Mead, F. W., 1976
Mendez, E., 1976
Menke, A. S., 1969
Messersmith, D. H., 1965
Miller, C. E., 1981
Miller, D. D., 1975
Miller, D. R., 1969

Miller, G. L., 1981
Miller, R. M., 1974
Miller, R. S., 1981
Miller, S. E., 1980

Mockford, E. L., 1955
Moore, T. E., 1950
Moraes, A. P. A. de, 1978
Morgan, N. O., 1969
Morse, J. C., 1976
Moser, J. C., 1973
Moulding, J. D., 1979
Mullens, B. A., 1979
Muraleedharan, D., 1981
Murdoch, W. P., 1966
Nakahara, S., 1968
Neal, J. W., Jr., 1982
Neff, S. E., 1969
Negrobov, O. P., 1979
Nelson, C. H., 1969
Nelson, G. H., 1949
Nelson, R. H., 1933
Neunzig, H. H., 1956
Newkirk, R. A., 1968
Nickle, D. A., 1980
Nielson, L. T., 1951
Norton, R. A., 1978
Novak, R. J., 1974
Nuhn, T. P., 1981
Nutting, W. H., 1973
Oatman, E. R., 1980

657

658 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

O’Brien, M. F., 1979
Oman, P. W., 1930
Orth, R. E., 1981
Osgood, E. A., 1977
Pakaluk, J., 1981
Palacios-Vargas, J. G., 1980
Palmer, M. K., 1976
Parker, €. R., 1977
Parrish, D. W., 1963
Parsons, M., 1963
Pena-G., L. E., 1980
Perez-H., A. M., 1974
Perkins, E. M., Jr., 1977
Perkins, P) D:; 1973
Peniyn De Rel 979
Petenss levies 979
EeterssaW sleet S71
Peterson, B:. L., 1979
Peterson, B: V., 1952
Retersona eles lO8i
Peyton, E. L., 1968
Phillips, W. G., 1955
Rifers D> Hee98il
Pitkin, B. R. 1977
Pogue, M. G., 1980
Polhemus, J. T., 1964
Poyner, M. M., 1969
Pratt, G. K., 1974
RraticpeDss943
Preval, S., 1978
Prices Ds Weil
Price Rebs, 1963
Pulawski, W. J., 1975
RackaiGe 1975
Radovic; I., 1981
Rainwater, H. I., 1964
Ramalingam, S., 1970
Ramos, J. A., 1947
Ramsay, M. J., 1968
Rawlins, J. E., 1974
Reed, W. D., 1931
Reichart, C. V., 1946
Reinert, J. F., 1969
Ribble, D. W., 1981
Richards, L. L., 1978
Riegel, G. T., 1952
Robbins, R. G., 1979
Roberts, D. R., 1968
Robinson, H., 1963
Robinson, W. H., 1975

Rohwer, G. G., 1964
Rolston, L. H., 1973
Rose, S., 1981

Ross, M. H., 1981
Roth, L. M., 1944
Roth, M., 1968
Rozen. JniG:, Jr..01956
Ruiter, D., 1976
Russell, L., 1979
Russell, L. M., 1930
Sabrosky, C. W., 1946
Sailer, R. I., 1943
Santana, F. J., 1966
Saugstad, E. S., 1979
Scanlon, J. E., 1952
Scarbrough, A. G., 1971
Schaber, B. D., 1980
Scharf, W. C., 1981
Schauff, M. E., 1980
Schmidt, C. H., 1969
Schroder, R. F. W., 1976
Schroeder, P. M., 1969
Schwan, R. G., 1980
Sedmane YouSs.95i
Selander, R. B., 1954
Shaffer, J. G 1974
Shands, W. A., 1940
Shanks, S. S., 1977
Shannon, M. H., 1980
Shaw, S. R., 1979
Shenefelt, R. D., 1946
Shepard, H. H., 1927
Sherman, R. W., 1947
Shewell, G. E., 1949
Shinohara, A., 1981
Shockley, C. W., 1955
Sholess OF DAVE 1979
Shubeck, P. P., 1982
Simpson, K. W., 1975
Sirivanakarn, S., 1969
Skaptason, J. L., 1979
Skiles, D. D., 1978
Slater, J. A., 1949
Sleeper, Ewe 1976
Slusssal E70
Smiley, R. L., 1964
Smith, C. F., 1967
Smith, Gs .. 1977
Smith, D. R., 1965*
Smith, F. F., 1921

VOLUME 84, NUMBER 3

Snelling, R. R., 1968
Sollers-Riedel, H., 1938*
Sommerman, K. M., 1947
Southem=bs.S-. 1977,
Spaeth, V. A., 1976
Spanelem vel 958=
Shrsmeeir, (C, 13),, Yrs, WS)
Spicer, G., 1978
Spilman, T. J., 1950
Sprance. Hebel oi
SUNS, (Cs Ihsan dirs, NEWS
Stannard, L. J., 1948
StarckesHe. 955,
Steffan, W. A., 1970
Sieomalen Gabe eboOS
Steiner, W. E., Jr., 1979
Steinhauer, A. L., 1958
Stewart, K. W., 1981
Steyskal, G. C., 1947
Stibick, J. N. L., 1966
Stimmel, J. F., 1979
Stoetzel, M. B., 1971
Stoltzfus, W. B., 1967
Stones Aes) 1931
Surdick, R. F., 1979
Sutherland, C. M., 1974
Sutherland, D. W. S., 1973
Matt Sees 979

Tanner, V. M., 1930
Thompson, F. C., 1968
Thompson, J. V., 1953
Thompson, P. H., 1965
Thornburg, M. C., 1961
Thorpe, K. W., 1980
Threlfall, W., 1977
Tibbetts, T., 1955
Tidwell, M. A., 1981
Todd, E. L., 1953
Townes, G. F., 1956
Townes, H., 1941

Townsend, L. H., Jr., 1977

inacere. We.Jn.. 1962
Trapido, H., 1948
Traub, R., 1947
Triplehorn, C. A., 1972
Trumble, J. T., 1979
Tyson, W. H., 1970
Ulrich, H., 1978*
Unzicker, J. D., 1981
Wittnars Js Aw. 1974"

Valley, K., 1976

Van Wie, L., 1979
Vazquez, A., 1957
Villegas, B., 1977
Vincent, D. L., 1980
Viraktamath, C. A., 198]
Voegtlin, D., 1981
Vogt, G. B., 1947
VoshellieselcesnL oui,
Walker, H. G., 1941
Wallenmaier, T. E., 1979
Wallis, R. C., 1948
Walton, M., 1937
Ward, R. A., 1975
Watrous, L. E., 1977
Weaver, J. S. III, 1980
Webb, D. W., 1982
Webb, R. E., 1967
Weems ih. Ve. Jn 1953
Weisman, D. M., 1956
Wendleton, D. S., 1965
Werner, F., 1948
Wharton, B., 1981
Wheeler, A. G., 1974
Wheeler, G. C., 1949
Whitcomb, R. F., 1966
White, G. B., 1977
White, R. E., 1966
White, T. R., 1979
Whitehead, D. R., 1974
Whitsel, R. H., 1967
Whittemore, F. W., 1974
Wilder, D. D., 1980
Wilkerson, R. C., 1980
Wilkinson, R. S., 1971
Williams, H. B., 1977
Williams, M. L., 1971
Williams, R. W., 1946
Wills, W., 1977
Wilson, A. M., 1978
Wilson, N., 1957
Wirth, W. W., 1945
Wojtowicz, J. A., 1981
Wolfe, G. W., 1977
Wood, F. E., 1968
Wood, T. K., 1974
Yonke, T. R., 1971
Young, D. A., Jr., 1950
Young, D. K., 1981
Zenner-Polania, I., 1977

659

660

Zimmerman, E. C., 1965
Zungoli, P. A., 1978
Zuska, J., 1974

Emeritus Members

Cartwright, O. L. (1948; 1980)
Gibson, E. H. (1916; 1965)
Hatch, M. H. (1921; 1975)
James, M. T. (1942; 1972)
Knowlton, G. F. (1935; 1973)
McGovran, E. R. (1937; 1973)
McGuire, J. U., Jr. (1954;
1980)

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Mason, H. C. (1949; 1973)

Mitchell, R. T. (1949; 1978)

Munson, S. C. (1938; 1976)

Parker, H. L. (before 1918;
1963)

Rainwater, C. F. (1954; 1975)

Richardson, H. H. (1939;
1976)

Spilman, R. E. (1950; 1977)

Swartzwelder, E. B. (1948;
1974)

Weber, N. A. (1941; 1981)

Woke, P. A. (1936; 1976)

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CONTENTS
(Continued from front cover)

GORDH, G. and B. A. HAWKINS—Tetrastichus cecidobroter (Hymenoptera: Eulo-
phidae), a new phytophagous species developing within the galls of Asphondylia
(Diptera: Cecidomyiidae) on Atriplex (Chenopodiaceae) in southern California ...

HARRIS, S. C., P. K. LAGO, and R. W. HOLZENTHAL—An annotated checklist
of the caddisflies (Trichoptera) of Mississippi and southeastern Louisiana. Part
II: Rhyacophiloidea

HOLZENTHAL, R. W., S. C. HARRIS, and P. K. LAGO—An annotated checklist
of the caddisflies (Trichoptera) of Mississippi and southeastern Louisiana. Part III:
Limnephiloidea and conclusions

KORMILEV, N. A.—Six new species of Neotropical Aradidae (Hemiptera)
LAGO, P. K., R. W. HOLZENTHAL, and S. C. HARRIS—An annotated checklist

of the caddisflies (Trichoptera) of Mississippi and southeastern Louisiana. Part I:
Introduction and Hydropsychoidea

LAVIGNE, R. J.—Ethology of Neoitamus vittipes (Diptera: Asilidae) in South Au-
stralia

MAIER, C. T.—Abundance and distribution of the seventeen-year periodical cicada,
Magicicada septendecim (Linnaeus) (Hemiptera: Cicadidae—brood II), in Connec-

MAIER, C. T.—Larval habitats and mate-seeking sites of flower flies (Diptera: Syrphi-
dae, Eristalinae)

MATHIS, W. N.—Description of a new species of Nocticanace Malloch (Diptera:
Canacidae) from Sri Lanka with notes on two related species

MATTA, J. F.—The bionomics of two species of Hydrochara (Coleoptera: Hydrophili-
dae) with descriptions of their larvae

MAYOR, A. J.—A review of the genus Chaetocoelus LeConte (Coleoptera: Malachi-

PETERS, T. M. and D. ADAMSKI—A description of the larva of Dixella nova
(Walker) (Diptera: Dixidae)

SCHAUFF, M. E. and E. E. GRISSELL—Nomenclatural notes on Polynema (Hymen-
optera: Mymaridae), with description of a new species

WEBB, D. W.—Smittia lasiops (Malloch): A redescription of the adults with a descrip-
tion of the immature stages (Diptera: Chironomidae)

WILKERSON, R. C. and G.-B. FAIRCHILD—Two new species of Esenbeckia
(Diptera: Tabanidae) from Mexico

WILKERSON, R. C. and G. B. FAIRCHILD—Five new species of Diachlorus (Dip-
tera: Tabanidae) from South America with a revised key to species and new
locality records

WIRTH, W. W.—The cacao-pollinating midges of the Forcipomyia argenteola group
(Diptera: Ceratopogonidae)

VOL. 84 OCTOBER 1982 NO. 4

SIS. 7TEB ee
| ae PROCEEDINGS,

of the
{

ENTOMOLOGICAL SOCIETY
| of WASHINGTON

DEPARTMENT OF ENTOMOLOGY
SMITHSONIAN INSTITUTION
WASHINGTON, D.C. 20560

PUBLISHED QUARTERLY

CONTENTS
FISHER, E. M. and H. A. HESPENHEIDE—Taxonomy and ethology of a new

Central American species of robber fly in the genus Glaphyropyga (Diptera:
EASTTITTEY EN 1 eis oc nomeceete ort Rae IR tn un ene SA OR ro mS CURE ery coe fr Ph 716

FOOTE, B. A.—Biology and immature stages of Setacera atrovirens, a grazer of
Wate dieal mats) (Diptera: Ephydridae)) .. 5.0 sonic 122.0 <u e wets be ein d's es nine 828

GILES, F. E. and W. W. Wirth—New species and new collection records of Cerato-

perdae (oiters) from or Lanka... a.s56ecios nak ote Ne awed be ooh Re 822
HEPPNER, J. B.—Synopsis of the Hilarographini (Lepidoptera: Tortricidae) of the

Mcreari lcd MMe Te Pe eee sens rea 2 Veh Rise Pot ier a ver ctes\ ial sal oifess adv alarael mus ey TRUE UMNO ele ele eae » core Peete pee 704
HUDSON, A.—Evidence for reproductive isolation between Xestia adela Franclemont

and Xestia dolosa Franclemont (Lepidoptera: Noctuidae) ...............+...0055 Hake,

| HUNG, A. C. F.—Chromosome and isozyme studies in Trichogramma (Hymenoptera:

MPVANMAATOAE) |) civ ket lec enc lannd nated ene dennieewaw nok «tea tonne e mene 791
KINGSOLVER, J. M.—Taxonomic studies in the genus Rhipibruchus Bridwell (Cole-
optera: Bruchidae), with descriptions of four new species ........ 66.660. e eens 661

' KINGSOLVER, J. M.—Conicobruchus albopubens (Pic) (Coleoptera: Bruchidae) and
its host Cyamopsis tetragonoloba (L.) (Leguminosae), with the designation of

I Scape as en N g eid tdi Ptinie SS & 2 aiy a Caw & San NS KO Rte Ss ome ae 845
_ KIRCHNER, R. F.—A new Allocapnia from West Virginia (Plecoptera: Capniidae) ... 786
) KONDRATIEFF, B. C. and J. R. VOSHELL, JR.—The Perlodinae of Virginia,
lee )6CUUSA (Plecoptera: Perlodidac) .............0scccccceerense cree seenceseseacess 761
LAGER, T. M., M. D. JOHNSON, and W. P. McCAFFERTY—The mayflies of
northeastern Minnesota (Ephemeroptera) .........---- +e eee cece ee ee eeees 729

(Continued on back cover)
.
he

rt

THE

ENTOMOLOGICAL SOCIETY
OF WASHINGTON

ORGANIZED MARCH 12, 1884

OFFICERS FOR 1982

MARGARET S. COLLINS, President HELEN SoOLLeRS-RIEDEL, Hospitality Chair
MAnyA B. STOETZEL, President-Elect Jay C. SHAFFER, Program Chairme
THOMAS E. WALLENMAIER, Recording Secretary Joyce A. Urmar, Membership Chairman
RALPH E. HARBACH, Corresponding Secretary JOHN F. CARROLL, Custodian |
F. CHRISTIAN THOMPSON, Treasurer Marcartt S. Coins, Delegate, Wash. Acad. Si. |

Davip R. Situ, Editor

Publications Committee
E. Eric GRISSELL GEORGE C. STEYSKAL
JOHN M. KINGSOLVER WAYNE N. MATHIS THOMAS E. WALLENMAIER

Honorary President
C. F. W. MUESEBECK

Honorary Members
FREDERICK W. Poos ASHLEY B. GURNEY RAYMOND A. ST. GEORGE

All correspondence concerning Society business should be mailed to the appropriate officer at the following address: Entomological
Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution, Washington, D.C. 20560.

MEETINGS. — Regular meetings of the Society are held in the Natural History Building, Smithsonian Institution, on the first Th
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Please see p. 183 of the January 1981 issue regarding preparation of manuscripts.

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Title of Publication: Proceedings of the Entomological Society of Washington.
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PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 661-684

TAXONOMIC STUDIES IN THE GENUS RHIPIBRUCHUS BRIDWELL
(COLEOPTERA: BRUCHIDAE), WITH DESCRIPTIONS OF
FOUR NEW SPECIES

JOHN M. KINGSOLVER

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—The South American genus Rhipibruchus is apparently re-
stricted to seeds of Prosopis spp. (Leguminosae), common name “‘mes-
quite.’’ Six species are now assigned to it including Rhipibruchus picturatus
(Fahraeus), R. prosopis Kingsolver, and four new species, R. rugicollis, R.
oedipygus, R. atratus, and R. psephenopygus. Host records, geographical dis-
tribution, a key to species, and illustrations of salient characters are pro-
vided.

Since my summary of Rhipibruchus was published in 1967, four additional
new species and numerous host associations have been discovered. In this
paper the generic description is revised, new species are described, host
records are tabulated, and a key to species is given. Life histories of some
of these species are being investigated by Arturo Teran and Susana M. de
L’ Argentier in Argentina. Collections have also been made by Hugo Cordo
and associates in Argentina for possible biological control of mesquite in
that country and in the United States.

Genus Rhipibruchus Bridwell

Rhipibruchus Bridwell, 1932: 105. New name for Megalorhipis Philippi,
1859: 668 (not Lacordaire, 1857). Type-species: Megalorhipis leiboldi Phi-
lippi, 1859: 668 = Rhipibruchus picturatus Fahraeus, 1839: 2. Automatic
type designation.

Genus in subfamily Bruchinae, tribe Acanthoscelidini. Body ovate, rather
broad. Head short, broad; width across eyes approximately equal to height
of head, strongly constricted behind eyes; eyes prominent, more narrowly
separated in d than in 2 (Fig. 23); frons with prominent median carina
expanded dorsally into triangular boss; d antenna strongly pectinate from
4th segment (Fig. 16), 2 antenna strongly serrate from 4th segment (Fig.

662 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

17). Pronotum campaniform, disk convex with narrow median sulcus sep-
arating paired basal and apical tumescences, or pronotum uniformly convex
without prominent tumescences; median basal lobe rounded, lateral margins
slightly convex or sinuate; lateral carina obsolete or faintly indicated by
threadlike line. Scutellum small, narrow. Elytra subquadrate, each elytron
evenly rounded at apex; disk flat, or subdepressed medially; striae usually
well defined, 3rd and 4th striae commonly arising at base from prominent,
rounded, bidentate gibbosity. Pro- and mesolegs slender, procoxae contig-
uous apically, mesocoxae narrowly separated; metafemur moderately in-
crassate, finely serrate in basal %4 on ventroposterior margin, pecten with 3
or 4 denticles (Fig. 24). Male pygidium arcuate in lateral profile, apex fitting
into emargination of Sth sternum; sterna 2-5 telescoped; ° pygidium nearly
flat in lateral profile, vertical or oblique. Male genitalia with median lobe
long and slender (Fig. 2); lateral lobes elongated, slightly expanded apically
(Fig. 3).

The male genitalia in the six known species of Rhipibruchus (and one
species of Pectinibruchus Kingsolver, a closely related genus) are basically
similar with only subtle specific differences; however, the basic structure is
so unlike that of any other species or group of species that for future ref-
erence in relating Rhipibruchus and Pectinibruchus to other groups, the
following description is given (Fig. 10): The median lobe is elongated, par-
allel-sided in apical two-thirds but cucullate in basal one-third; the ventral
valve is ovate-acuminate and varies but little throughout the genus. Details
of the armature of the internal sac are:

1. A pair of acute, obliquely positioned transparent plates with their apices
extending into apical orifice: the surfaces of the plates appear to be minutely
pitted perhaps with glandular openings but the shape of the plate does not
vary to supply specific differences except in R. picturatus.

2. A transparent plate bearing 3—6 spines on its dorsal face: the shape of
this plate and the number of spines is diagnostic.

3. A cluster of setae resembling a brush: length and shape of the brush is
diagnostic.

4. A cluster of thornlike spines or denticles: the shape, number, and
relative number of spines is diagnostic.

5. A saclike structure near apex of internal sac carrying two rodlike scler-
ites; not diagnostic.

The lateral lobes are not strikingly different among the species, the prin-
cipal diagnostic features being the length of the lobes, shape of the apex of
each lobe, and the depth of the cleft between them.

Color and pattern of the pronotum and pygidium are fairly consistent for
each species but the pattern on the elytra is somewhat variable and must
be used with caution in concert with other characters.

VOLUME 84, NUMBER 4 663

Table 1. Rhipibruchus species and their Prosopis host plants.

Rhipibruchus Species Prosopis Species

R. atratus P. abbreviata Benth.; P. alba Griseb.; P. caldenia Burkart; P. flexu-
osa DC.; P. nigra (Griseb.) Hieron.; P. ruscifolia Griseb.; P. torquata
(Lag.) DC.

R. oedipygus P. kuntzei Harms.

R. picturatus P. alba; P. affinis Spreng.; P. caldenia; P. chilensis (Mol.) Stuntz; P.

elata (Burk.) Burk.; P. ferox Griseb.; P. flexuosa; P. humilis Hook.
& Arn.; P. nigra; P. strombulifera (Lam.) Benth.; P. torquata.

R. prosopis P. affinis Spreng.; P. alpataco Phil., P. calingastana Burk.; P. chi-
lensis; P. juliflora (Sw.) DC.; P. kuntzei; P. nigra; P. sericantha Hook.
& Arn.; P. strombulifera.

R. psephenopygus P. alba; P. alpataco; P. caldenia; P. chilensis; P. flexuosa; P. juliflora;
P. nigra; P. torquata.

R. rugicollis P. kuntzei; P. sericantha.

Species in this genus are so far known to breed only in seeds of various
species of Prosopis L. (Leguminosae) (mesquite, algarrobo, algarobillo).
Rhipibruchus is restricted to South America principally in north and central
Argentina and Chile but records from Uruguay and a questionable record
from Colombia are known. Host species and their associated bruchids are
listed in Tables | and 2.

Rhipibruchus is closely related to Pectinibruchus Kingsolver but the af-
finities of these genera remain obscure. The South American genus Pseu-
dopachymerina superficially resembles Rhipibruchus but | believe that they
are only remotely related. The male genitalia of Pseudopachymerina are
quite different from those of Rhipibruchus in basic structure; the metafemur
is broader than in Rhipibruchus; the antenna is not sexually dimorphic as
it is in Rhipibruchus; and the basal denticles of the third and fourth elytral
striae are not elevated on a tumescence.

Rhipibruchus is tentatively separated into three groups primarily on the
basis of body form and color pattern, (1) the rugicollis group including only
rugicollis; (2) the oedipygus group including only oedipygus; and (3) the
picturatus group including picturatus, prosopis, atratus, and psephenopy-
gus.

KEY TO SPECIES OF RAHIPIBRUCHUS

1. Body large, 3.6-5.4 mm in length; with no distinct pattern on prono-
tum, elytra, or pygidium; vestiture mostly gray with pronotum and
elytra with mixed gray and bronze; pronotum (Fig. 7) without prom-

664 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Prosopis species and their Rhipibruchus predators.

Prosopis Species Rhipibruchus Species
P. abbreviata R. atratus
P. affinis R. picturatus
P. alba R. atratus; R. picturatus; R. psephenopygus
P. algarobilla R. picturatus (literature record, Bosq, 1942) = P. affinis
P. alpataco R. prosopis; R. psephenopygus
P. caldenia R. atratus; R. picturatus; R. psephenopygus
P. calingastana R. prosopis
P. chilensis R. picturatus; R. prosopis; R. psephenopygus
P. elata R. picturatus
P. ferox R. picturatus
P. flexuosa R. atratus; R. picturatus; R. prosopis; R. psephenopygus
P. humilis R. picturatus (literature record, Bosq, 1942)
P. juliflora R. prosopis; R. psephenopygus
P. kuntzei R. oedipygus; R. prosopis; R. rugicollis
P. nigra R.atratus; R. picturatus; R. prosopis; R. psephenopygus
P. ruscifolia R. atratus
P. sericantha R. prosopis; R. rugicollis
P. silaquastrum R. picturatus (literature record, Zacher, 1952) = P. chilensis
P. strombulifera R. picturatus; R. prosopis
P. torquata R. atratus; R. picturatus; R. psephenopygus

inent median tumescences but with subbasal tumescences ............

SE ee OS a Mk Gra Ae ht a ery ae oedipygus, new species
— Body large or small; usually with pattern on dorsal surfaces, color
mixed; pronotum with longitudinal tumescences adjacent to midline;
sub-basal tumescences prominent? 22s. 221 «272 e022. eee 2
Pronotum in lateral aspect with dorsal profile strongly sinuate (Fig.
6); elytra without strongly contrasting pattern (Fig. 1); pygidium with
short, narrow, white basal triangle and small tuft of white either side

th

at basal\4.(Figei'4y SP UO a es, See rugicollis, new species
— Pronotum in lateral aspect with dorsal profile only slightly sinuate
(Fig. 18); elytra and pygidium usually with contrasting pattern ...... 3

3. Antenna strongly pectinate (Fig. 16); eyes narrowly separated by
frons (Fig. 23, 6); pygidium with faint sublateral depressions (Fig.
28); Male PY oA SOY SEO TA ee 4
— Antenna serrate (Fig. 17); eyes more widely separated by frons (Fig.
23, 2); pygidium with prominent sublateral depressions often con-
trastingly marked in integument and vestiture (Fig. 29); female ..... v
4. Pygidium with integument uniformly dark brown to black with at
most a narrow reddish-yellow stripe from middle of disk to apex,
vestiture sparse, evenly ‘distributed (Fis: 33)) 2202. ates. Ci ae
SGlncig attanshite 216 wiaiep ores. Gre Croat ee eee psephenopygus, new species

VOLUME 84, NUMBER 4 665

— Pygidial integument with extensive yellowish to yellowish-red areas,

vestiure usuallyan dense patches (Figg 28) 6 ss... coca s crud ves Bee 5
5. Pronotum mostly piceous with contrasting narrow, white basal patch

of setae and narrow median line, rarely with cruciate mark in middle

of disk (Fig. 25); elytra with lateral and apical margins broadly black,

median area of disk with 24 isolated dark spots but without large

median, common discal spot, or broad, transverse band (Fig. 25)

a tS 2 IR oO ee eee atratus, new species
— Pronotal disk piceous with flanks and cruciate median mark gray;

elytra with margins usually mottled or streaked with longitudinal

spots, median area usually with large discal spot or continuous or

brokem bandi(Bigs-12 19, 30) ccc.as. 040 See vcs dem on age See 6
6. Vestiture and integument of basal % of elytra yellowish, middle of

elytra usually with large, common discal spot (Fig. 19), occasionally

with transverse band; male genitalia with transparent plates at apical

Onbleercloneabed (HIS. 21) 51. oc. 52 ein inyesbye0 Sin yetoves eh prosopis Kingsolver
— Vestiture of basal % of elytra gray; middle of elytra occasionally

with discontinuous transverse dark band but never with discal spot

(Fig. 12); male genitalia with transparent plates short and broad (Fig.

IAIN, pct st eat o,1S e ie MRTP picturatus (Fahraeus)
7. Apical 1/5 of elytra largely black, occasionally with paler elongate

spot on 7th interval (Fig. 25); pygidium with strongly contrasting

black or piceous sublateral depressions, these separated by median,

SUMS IS LIS 2D) pec... 20s. «,paciniaim ah ep arches m= Sei ane atratus, new species
— Apices of elytra variegated black and yellowish brown (Figs. 12, 19,

30); pygidium with contrasting spots or these obscure (Figs. 13, 20,

BN ge ER ead ha chats bt tal ac blak » is tyeie're Sepa arate tesa Ae 8
8. Pygidial integument uniformly piceous with at most a reddish stripe

from middle of pygidium to apex (Fig. 34); sublateral spots obscure

See Mig or hte ok fo ho Saat. oe Sackalayn yis psephenopygus, new species

— Pygidial integument with contrasting dark sublateral spots (Figs. 13,

ZO) VA, oe ee ea ene re ees meee mpe es reer en 9
9. Pygidium with basal triangle and short median stripe white, remain-

ing vestiture yellowish, lateral spots piceous or black (Fig. 20)

SON Sag Sto Se Perret eee errr C prosopis Kingsolver
— Pygidial vestiture silvery gray, basal triangle and ‘“‘eyebrow”’ spots

above sublateral black or piceous spots appearing more intensely

white than intervening areas (Fig. 13) .......... picturatus (Fahraeus)

RuGICOLLIS GROUP

Pronotum with prominent medial and subbasal tumescences; elytra and
pygidium with mottled pattern; pygidium not sexually dimorphic and with-
out darkened depressions in female.

666 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Rhipibruchus rugicollis Kingsolver, NEw SPECIES
Figs. 1-6, 38

Body length.—3.9-4.4 mm; width, 2.1—2.3 mm. Pronotal length, 1.3-1.5
mm; width, 1.4-1.6 mm.

Color.—Integument dark red to piceous, without distinct pattern; head
piceous except eyes black, spot above each eye reddish; antenna uniformly
piceous; pronotum, elytra, pygidium, and venter of body uniformly piceous
except metacoxa reddish, metafemur darker in ventral 2. Vestiture of gray
and yellowish hairs in mottled pattern on both dorsal and ventral surfaces
(Fig. 1); pygidium mottled with hairs obliquely positioned toward midline,
with short basal triangle and 2 small lateral clumps of white hairs (Figs.
4, 5).

Structure.—Body elongated; pronotum campaniform; elytra rectangular,
together slightly longer than wide. Head obovate, eyes strongly protuberant
laterally, vertex finely, densely punctate, frons strongly carinate in both
sexes, narrower ind than in@, slopes of carina punctulate, clypeus micro-
rugose, postocular fringe narrow. Pronotum (Figs. |, 6) with lateral margins
sinuate; longitudinal, median, undulate elevation broadly sulcate, sublateral
gibbosities prominent; surface of pronotum densely foveolate, each foveola
setiferous; lateral carina traceable from posterolateral corner to a point im-
mediately above procoxal cavity; cervical sulcus fine, short; cervical boss
prominent, bisetiferous; prosternum short, intercoxal piece extending 2
length of coxae; mesosternum narrow, triangular; postmesocoxal sulci meet-
ing on midline, fine, parallel to coxal cavity. Scutellum elongate, 2 as long
as wide, bidentate apically, densely pilose. Elytra elongate (Fig. 1), gently
convex but subdepressed along suture and basally between 3rd intervals;
striae fairly regular in course except 3rd and 4th bent laterally at base,
shallowly sulcate, punctures elongate, setiferous; Ist stria arising near scu-
tellar apex, 2nd arising behind vertical basal ridge, 3rd and 4th arising from
fine denticles on summit of prominent subbasal gibbosity, 5th arising on
lateral limits of gibbosity; all striae free apically, 4th and Sth abbreviated;
intervals flat, strigulate, densely setose. Abdomen of d strongly telescoped,
length of segments 2-5 less than that of Ist segment, Sth segment with
posterior border emarginate for reception of apex of pygidium; pygidium
elongate and narrowly triangular, basal margin arcuate, surface convex,
densely, evenly microfoveolate; abdomen of 2 with segments 2—5 together
slightly longer than Ist segment, Sth segment with posterior margin evenly
rounded; pygidium (Figs. 4, 5) elongate-obovate, oblique, surface convex,
shallowly impressed sublaterally toward apex, surface densely microfo-
veolate. Pro- and mesolegs not modified; metacoxa densely, finely punctate
except for elongate, polished proximal area; metafemur moderately incras-
sate, dorsal margin evenly arcuate, ventral margin arcuate in basal 4/5,

VOLUME 84, NUMBER 4

Figs. 1-6. Rhipibruchus rugicollis. 1,

667

6

Habitus, dorsal. 2, 6 genitalia, median lobe. 3, d

genitalia, lateral lobes. 4, Pygidium 3. 5, Pygidium 2. 6, Prothorax, lateral profile.

668 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sinuate apically, pecten with | long and 3 shorter denticles on its distal
slope; metatibia slender, more narrowed proximally and gradually widened
distally, mucro short, acute, lateral denticle short, coronal denticles 3 or 4;
lateral, ventral and dorsomedial carinae distinct and complete to apex, la-
teroventral carina obsolete in apical 4/5. Male genitalia (Figs. 2, 3) with
internal sac of median lobe with brush of setae inconspicuous, poorly de-
fined; spinous plate elongate, with 4—S spines; median cluster of 12—15 thorn-
like or sickle shaped denticles. Lateral lobes with apices oblique, cleft to
about % their length.

Holotype d.—ARGENTINA: Salta Prov., Tartagal, December 1950, Da-
guerre (USNMNH type #76390).

Allotype 2 and 6d, 32 paratypes.—ARGENTINA; Santiago del Estero
Prov., Termes de Rio Hondo, 20 April 1972, ex Prosopis sericantha.

Other paratypes.—ARGENTINA: Tucuman Prov., Tucuman, 1941, J.
Hayward, 46, 2 2. Formosa Prov., Rt. 81, 14 mi SE Cdte. Fontana, 12
March 1977, ex Prosopis kuntzei, 1¢, 19; Rt. 81, 58 mi SE Ing. Juarez,
11 March 1977, ex Prosopis kuntzei, 1°. Chaco Prov., in “‘itin’’ (Prosopis
kuntzei), 1d, 19; Chaco, ex Prosopis kuntzei, 14, 1 2. Paratypes de-
posited in the National Museum of Natural History, Washington, D.C.
(USNMNH), the Fundacion Miguel Lillo, Tucuman, and Bernadino Riva-
davia Museum of Natural Sciences, Buenos Aires, Argentina.

Discussion.—This species is easily recognized by the prominent medial
tumescences, the mottled pattern on the elytra and pygidium, the lack of
darkened depressions on the female pygidium, and the larger size.

Remarks.—The name rugicollis is derived from the Latin ruga (wrinkled)
and collum (neck) and refers to the rugose pronotum. In Kingsolver et al.
(1977: 115), R. rugicollis was listed as R. sp. D.

OEDIPYGUS GROUP

Pronotum convex, without medial but with subbasal tumescences; elytra
and pygidium with vestiture evenly distributed, not sexually dimorphic,
without darkened depressions in female.

Rhipibruchus oedipygus Kingsolver, NEw SPECIES
Figs. 7-11, 38

Body length.—3.6—-5.4 mm; width, 2.0-2.6 mm. Pronotal length, 1.8—2.0
mm; width, 2.0-2.5 mm.

Color.—Integument mostly piceous with following areas reddish: trian-
gular spot above eye, lateral margin of pronotum, posterior “3 of metaster-
num, abdominal sterna, pro- and mesofemora, dorsal 2 of metafemur, meta-
tibia; with following areas yellowish red: antenna with rami darker ind,
pro- and mesotibiae. Vestiture of intermixed gray and bronze slender hairs

VOLUME 84, NUMBER 4 669

7 as
7
Figs. 7-11. Rhipibruchus oedipygus. 7, Prothorax, lateral profile. 8, Pygidium ¢, lateral

profile. 9, Pygidium 9, lateral profile. 10, ¢ genitalia, median lobe. 11, ¢ genitalia, lateral lobes.

evenly distributed over body, elytra with predominantly bronze vestiture,
remainder of body predominantly gray with bronze setae set in foveolae.
Structure.—Body elongate, head subtriangular, eyes strongly protuber-
ant, nearly divided by ocular sinus with about 7 rows of facets between
sinus and posterior margin of eye, supraocular sulcus with setiferous, um-
bilicate punctures; vertex densely foveolate posteriorly, with subtriangular,
minutely granulose boss anteriorly leading to frontal carina, frons foveolate
on lateral slopes, narrower ind (width across eyes: narrowest frontal width
7:1) than in 2 (4:1); clypeus broadly concave, shallowly foveolate, densely
setose; labrum finely granulose; d antenna with scape cucumiform, pedicel

670 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

short, Ist flagellar segment angular, 2nd with short ramus, remaining seg-
ments each with long, setose ramus; 2 antenna serrate from 4th segment.
Pronotum campaniform, apex evenly arcuate, lateral margins moderately
sinuate, posterolateral angles slightly flared, basal margin sinuate, basal lobe
prominent, prominent condyle near basal margin either side of basal lobe;
in lateral aspect, dorsal profile nearly flat in basal +/s, strongly convex at
apex; surface densely, evenly foveolate, each foveola bearing short, bronze
seta; lateral carina short (Fig. 7); cervical sulcus short, obscure, cervical
boss with 3-5 setae; prosternum short, triangular, not separating procoxae
apically; mesosternum narrow, rounded and slightly sulcate at apex; post-
mesocoxal sulcus absent. Scutellum 142 as long as wide, apex shallowly
emarginate, bidentate. Elytra slightly longer than wide, evenly convex; striae
uniform, evenly spaced, shallowly sulcate, foveolate, well marked in medial
4, partly to completely obliterated laterally, especially in basal 2, sutural
stria deep, narrow, 2nd stria arising basally from deep pit beneath marginal
bead, 3rd and 4th arising from prominent bidentate basal gibbosity, Sth on
lateral margin of gibbosity, bases of remaining discal striae obliterated in
imbricate caudal extension of humeral umbo, striae free apically with 4th
and Sth abbreviated and approximate; intervals finely imbricate and fo-
veolate, especially toward base. Abdomen with basal sternum 2x as long
as Sth sternum ind, subequal in length in 2, caudal margin of Sth deeply
emarginate for reception of apex of pygidium in d, evenly rounded in 2;
pygidium obovate in caudal aspect in both sexes, strongly convex ing (Fig.
8), only slightly convex in 2 (Fig. 9) in lateral aspect, apex of pygidium
fitting into emarginate Sth sternum, 2 pygidium nearly vertical, disk in both
sexes densely, evenly, shallowly foveolate, foveolae nearly concealed by
dense vestiture. Pro- and mesolegs not modified; metacoxa finely, densely
punctate except for anterior polished area; metafemur moderately incras-
sate, pecten with | large and 2 small denticles; metatibia slender, mucro
short, acute; lateral, ventral, and dorsomedial carinae distinct and complete,
lateroventral carina obsolete in apical 4%.

Male genitalia (Figs. 10, 11)—Internal sac of median lobe with brush of
setae short; spinous plate small with 5 spines; median cluster of about 40
small denticles; lateral lobes rounded apically, cleft to about /% their length.

Holotype d.—ARGENTINA: Chaco, ex seed of Prosopis kuntzei, Harms,
#1365. In collection of Fundacion Miguel Lillo, Tucuman, Argentina.

Allotype 2.—ARGENTINA: Tucuman, 1941, J. Hayward; on temporary
loan to the National Museum of Natural History, Washington, D.C.

Other paratypes—ARGENTINA: Formosa Prov., Rio Bermejo (Pto.
Lavalle), 23 August 1972, ex seeds Prosopis kuntzei, 23, 3°; Ibarreta, 8
April 1976, ex seeds ‘‘itin’’ (Prosopis kuntzei) 2 6. Santiago del Estero
Prov., north of the city, July 1965, ex seeds Prosopis kuntzei, 23.

VOLUME 84, NUMBER 4 671

Diagnosis.—This distinctive species differs from R. prosopis, R. pictur-
atus, R. atratus, and R. psephenopygus by the absence of a distinct color
pattern, its larger size, strongly convex 6 pygidium, and details of the d
genitalia. It differs from R. rugicollis by its color, lack of median pronotal
tumescences, color patterns of the d and 2 pygidia, and details of the 3
genitalia.

Remarks.—The name oedipygus is derived from the Greek oidema (swell-
ing) and pyge (rump) and refers to the bulbous pygidium of the male. In
Kingsolver et al. (1977: 115) R. oedipygus was listed as R. sp. E.

PictTuRATUS GROUP

Pronotum with slight medial tumescences and prominent subbasal tu-
mescences; elytra and pygidium with prominent pattern; pygidium sexually
dimorphic with darkened depressions in female.

Discussion.—The four known species in this group are quite closely re-
lated. Color patterns on the pronotum and elytra are comparatively uniform
for each sex of the four species. Sexual dimorphism in pygidial patterns is
quite striking and offers excellent distinguishing features. Characters given
in the key should be sufficient for identification but they are reiterated here
for convenience.

Rhipibruchus psephenopygus is easily separated by the dark median,
transverse band extending nearly or completely across the middle of the
elytra (Fig. 30), and by the uniformly distributed pygidial vestiture in both
sexes (cf. picturatus below) (Figs. 33, 34) except for the dark, slightly de-
pressed sublateral patches in the female.

Rhipibruchus atratus is distinctive in its contrasting black-and-white pro-
notal and elytral patterns (Fig. 25). The male pygidial pattern (Fig. 28) is
quite similar to those of picturatus and prosopis but elytral color and pattern
will serve to separate these three species. The pygidial pattern (Fig. 29) of
females of atratus with strikingly contrasted large, black subapical patches
margined medially by reddish brown hairs, and with dorsal patches of the
same reddish brown is diagnostic.

Rhipibruchus prosopis is easily separated from others in the genus by the
yellow vestiture of the elytra in both sexes. The pygidium in the female
(Fig. 20) is distinctively marked with a basal triangular and narrow median
line white, subapical marginal, depressed patches black, and the remainder
of the vestiture yellow. The middle of the elytra is usually marked with a
large, dark common spot (Fig. 19).

Rhipibruchus picturatus is most easily confused with psephenopygus but
the pale areas on the elytra are more extensive with the dark spots discon-
nected and not forming a transverse band. The white basal triangle and white
median line in both sexes of picturatus as well as the more prominent white

672 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

subbasal patches in the male (as in Fig. 28) and the more densely white
‘‘eyebrow’’ patches above the dark subapical depressed patches in 2 pic-
turatus (Fig. 13) are diagnostic.

Rhipibruchus picturatus (Fahraeus)
Figs. 12-18, 35

Bruchus picturatus Fahraeus, 1839: 2.

Megalorhipis leiboldi Philippi, 1859: 669.

Megalorhipis leyboldi: Pic, 1913: 12 (error).

Rhipibruchus picturatus: Bridwell, 1932: 105; Blackwelder, 1946: 758; King-
solver, 1967: 320; Kingsolver et al., 1977: 115.

The redescription (Kingsolver, 1967) and the group diagnosis given above
are sufficient to distinguish this species from others in the group.

Male genitalia (Figs. 14, 15)—Internal sac of median lobe with brush of
setae short; spinous plate short, broad, with 4-5 spines; median cluster of
5 large, thornlike, curved spines and 3-4 smaller denticles; lateral lobes
blunt apically, somewhat oblique on median margin, cleft to about % their
length.

New host records.—ARGENTINA: Catamarca Prov., Rt. 307 6 km N
Sta. Maria, 11 January 1980, in Prosopis flexuosa; 30 mi N Catamarca, ex
seeds Prosopis nigra; Arauco, near Aimogasta, ex seeds Prosopis chilensis.
Santiago del Estero Prov., Rt. 34, 31 mi SE La Banda, 12 June 1976, in
Prosopis alba. La Rioja Prov., Aimogasta, 25 February 1978, in Prosopis
chilensis; Rt. 9, 46 km N from Rt. 38, in Prosopis chilensis. Cordoba Prov.,
Rt. 38, Capilla del Monte, 22 February 1978, in Prosopis alba; Rt. 38,
Camino a los Mogotes, 25 February 1977, in Prosopis chilensis; Rt. 7, 1 mi
E. Cordoba-San Luis border, 19 November 1976, in Prosopis caldenia.
Entre Rios Prov., Rt. 126, 9 mi NE La Paz, 15 March 1977, in Prosopis
nigra. San Juan Prov., 3 mi E Caucete, 19 February 1978, in Prosopis
torquata; Calingasta, 17 May 1977, in Prosopis strombulifera. Mendoza
Prov., Rt. 40, Mojon km 19, 24 November 1976, in Prosopis strombulifera.
San Luis Prov., Alto Pencosa, 6 February 1951; Lavaisse, Don Roberto
Estancia. January 1943, 17 October 1942, February—March 1943; La Pampa
Prov., Rt. 35, 85 mi N Sta. Rosa, 15 February 1978. CHILE: Santiago Prov.,
Antumapu, I1 June 1974.

Rhipibruchus prosopis Kingsolver
Figs. 19-24, 36

Rhipibruchus prosopis Kingsolver, 1967: 322; Kingsolver et al., 1977: 115.

The original description (Kingsolver, 1967) and the group diagnosis given
above are sufficient to distinguish this species.

Ne

VOLUME 84, NUMBER 4 673

18

16 17

Figs. 12-18. Rhipibruchus picturatus. 12, Left elytron, integumental pattern. 13, Pygidium
2. 14, genitalia, median lobe. 15, d genitalia, lateral lobe. 16, Antenna ¢. 17, Antenna °.

18, Prothorax, lateral profile.

674 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

CUEING a

MNO A MG fee
SM EE Yen
Shy UY a,

Wi hetull icf
.\ mG

21

Figs. 19-24. Rhipibruchus prosopis. 19, Left elytron, integumental pattern. 20, Pygidium
2. 21,6 genitalia, median lobe. 22, 3 genitalia, lateral lobe. 23, Heads, d, 2. 24, Metafemur
and metatibia.

VOLUME 84, NUMBER 4 675

Male genitalia (Figs. 21, 22).—Internal sac with brush of setae moderately
long; spinous plate short, broad, with 2-3 spines; median cluster of 5 large,
curved, thornlike spines and 2-3 denticles; lateral lobes rounded apically,
cleft to about +/s their length.

New host records.—ARGENTINA; Formosa Prov., Rt. 81, 15 mi NW
Formosa, 10 December 1976, in Prosopis nigra; Rt. 81, 14 mi SE Cdte.
Fontana, 12 March 1977, in Prosopis kuntzei. Catamarca Prov., Catamarca,
31 January 1970. La Rioja Prov., Aimogasta, 25 February 1978, Prosopis
chilensis. San Juan Prov., Rt. 20, 1 mi SE Caucete, 19 February 1978, in
Prosopis alpataco and strombulifera; Rt. 40, 18 mi NE Mendoza-San Juan
border, 22 February 1977, in Prosopis flexuosa; Rt. 141, 12 mi S Calingasta,
16 May 1977, in Prosopis calingastana; Airport, Rt. 20, 10 km from San
Juan, 26 November 1976, in Prosopis strombulifera. Cordoba Prov., Rt. 38,
8 mi NW Cruz del Eje, 24 February 1977, in Prosopis chilensis. Entre Rios
Prov., Rt. 126, 9 mi NE La Paz, 15 March 1977, in Prosopis nigra. Mendoza
Prov., Rt. 40, 8 mi NE Mendoza, 22 February 1977, in Prosopis strombu-
lifera and alpataco; Rt. 40, Mojon, km 19, 24 November 1976, in Prosopis
strombulifera; Rt. 40, 4 mi N. Mendoza, 19 February 1978, in Prosopis
strombulifera.

Rhipibruchus atratus Kingsolver, NEw SPEciES
Figs. 25-29, 39

Body length.—2.9-3.1 mm; width, 1.5-1.7 mm. Pronotal length, 0.8—0.9
mm; width, |.1—-1.3 mm.

Color.—Integument reddish brown to black, head black except apex of
clypeus and postocular spots reddish; antenna piceous; pronotum black ex-
cept basal triangle and lateral and apical margins reddish; elytra (Fig. 25)
with reddish-brown median vitta extending laterally to 5th or 6th interval
and with subapical band extending to lateral margin, 3rd interval with sub-
apical black spot grading to apical 4 of elytra black, lateral margins black;
venter of body piceous to black, abdomen often reddish; pro- and mesolegs
reddish brown, darker proximally, metafemur piceous with dorsal reddish
brown area; metatibia reddish brown, darker proximally; pygidium of ¢
reddish brown, with subapical lateral margins dusky; pygidium of 2 dark
red. Vestiture of black, dark brown, yellowish-brown, yellow, and white
slender setae, those on head yellowish brown, on pronotum yellowish with
narrow, median white stripe expanded basally, white often intermixed with
yellow, darker areas of disk with dark brown hairs, flanks with yellowish
and white; elytra with white or yellowish setae on reddish integument, and
black with scattered intermixed white on black integumental areas; venter
of body and legs evenly covered with white hairs; pygidium of ¢ (Fig. 28)
densely covered with white hairs condensed into basal triangle and vague

676 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 25-29. Rhipibruchus atratus. 25, Habitus, dorsal. 26, 6 genitalia, median lobe. 27,

3 genitalia, lateral lobes. 28, Pygidium 3. 29, Pygidium ° .

VOLUME 84, NUMBER 4 677

lateral patches, of 2 (Fig. 29) with condensed basal triangle, median stripe,
and basolateral patches of white hairs intermixed with yellow; apicolateral
depressed, ovate spots with dark brown hairs.

Structure.—Body ovate, elytra quadrilateral, pronotum campaniform.
Head subtriangular, eyes strongly protruding laterally, sexually dimorphic,
relatively larger and more narrowly separated medially in d, ocular sinus
narrow, extending about +/s length of eye, vertex finely reticulate, frons
narrow, strongly carinate, finely punctate, clypeus scabriculous, labrum
semicircular, sparsely punctate, supraantennal ridges prominent, frons be-
tween ridges concave; ¢ antenna strongly pectinate, reaching metacoxa, in
2 serrate from 4th segment, reaching humerus. Pronotum (Fig. 25) with
basal margin strongly lobed, apical margin semicircular, lateral margins sin-
uate; disk strongly convex, shallowly sulcate either side of broad median
gibbosity, this with shallow sulcus in basal 2; obsolete subbasal gibbosity
near each posterolateral angle, surface of disk densely microfoveolate, each
foveola with a hair arising from its center or from its anterior rim; lateral
carina present as an obsolete ridge extending to procoxal cavity; cervical
sulcus short, fine; cervical boss small but distinct, bisetiferous; prosternum
short, triangular, acute posteriorly, procoxae contiguous in apical 4%. Scu-
tellum short, narrow, 2X as long as wide, deeply bifid apically. Elytra (Fig.
25) slightly longer than wide, widest at basal 4%, disk moderately convex,
subdepressed at basal, median 4; striae slightly sinuate, | and 2 each arising
from basal fovea, 3 and 4 arising from a prominent, minutely dentate, sub-
basal gibbosity, 5 arising laterad of gibbosity, striae 4 and 5 and 6 and 7
conjoined apically, remaining striae ending free; intervals flat, finely stri-
gose, 3 and 5 wider than 2, 4, and 6. Mesosternum acute apically; post-
mesocoxal sulci conjoined on midline, ending laterad of coxal cavity. Ab-
domen of ¢ telescoped, Ist sternum longer than remaining 4 together, Sth
sternum with posterior margin shallowly emarginate and narrowly reflexed
ventrad to receive apex of pygidium, of 2, Ist sternum subequal in length
to remaining 4 together, Sth sternum slightly tumescent in profile, percep-
tibly emarginate on posterior border; 6 pygidium strongly reflexed, disk
moderately convex transversely, surface finely, sparsely punctate, of 2 not
reflexed, disk convex with lateral darker areas subdepressed and finely,
densely punctulate, white setae nearly concealing punctuation. Pro- and
mesolegs not unusually modified; metacoxa densely, evenly punctate except
for elongate, polished area near trochanteral insertion; metafemur moder-
ately incrassate, more strongly arcuate on dorsal margin than on ventral,
pecten with | long and 2 shorter distal denticles, metatibia slender, wider
distally, mucro short, acute, lateral denticle prominent, 4 coronal denticles
fine, lateral, ventral, and dorsomedial carinae distinct and complete, later-
oventral carina obsolete in apical 4.

678 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Male genitalia (Figs. 26, 27).—Internal sac of median lobe with brush of
setae short; spinous plate elongate, parallel-sided, with 4 spines; median
cluster of 5 large, thornlike and 2—3 smaller spines; lateral lobes elongate,
apices bluntly oblique, cleft to about % their length.

Holotype ¢6.—ARGENTINA: Formosa Prov., Rt. 11, 12 mi N. Chaco-
Formosa border, 8 December 1976, in pods Prosopis nigra.

Allotype 2 .—Same data as holotype.

Other paratypes—ARGENTINA: Catamarca Prov., Rt. 60, 47 mi SE
Tinogasta, 25 February 1978, in seed pods, Prosopis torquata, 2°; Rt. 307,
6 km N Santa Maria, 11 January 1980, in seed pods Prosopis flexuosa, 1°;
Rt. 62, 26 mi S Andalgala, 1 March 1977, in pods, Prosopis abbreviata, |
2; 50 km W Andalgala, 31 October 1972, on Cassia, 136, 12; same data
except in pods, Prosopis flexuosa, 12. Formosa Prov., Rt. 81, 15 mi NW
Formosa, 10 December 1976, in pods on ground, Prosopis nigra, 23,1°.
La Pampa Prov., Rt. 188, 38 mi W Realico, 15 February 1978, in pods,
Prosopis caldenia, 1°. La Rioja Prov., Rt. 20, 23 mi E Chepas, 20 February
1978, in pods on ground, Prosopis torquata, 12; La Rioja, January 1970,
Pena, in Prosopis abbreviata 5°. San Juan Prov., Rt. 20, 3 mi E Caucete,
19 February 1978, in pods on ground, Prosopis torquata 4°; Matagusanos,
29 February 1970, 446, 62. Santiago del Estero Prov., Rt. 34, 31 mi SE
La Banda, 6 December 1976, in seed pods, Prosopis alba, 1°. San Luis
Prov., Lavaisse, Don Roberto Estancia, 12 and 18 October 1942, 2d. PAR-
AGUAY: Pres. Hayes Prov., Hwy. Trans Chaco 365 km NW Asuncion, 18
January 1976, DeLoach #72179, in pods, Prosopis ruscifolia, 13,2°.

Type and paratypes deposited in the Fundacion Miguel Lillo, Tucuman;
paratypes deposited in the National Museum of Natural History, Washing-
ton, D.C., Rivadavia Museum of Natural Sciences, Buenos Aires, Utah
State University, Logan, British Museum (Natural History), London, and
the Museo Nacional de Historia Natural, Santiago, Chile.

Diagnosis.—This species is easily recognized by the intensely black bor-
der of the elytra, the predominantly dark pronotum, and the strikingly marked
pygidium. It is probably most closely related to R. picturatus. See the pic-
turatus group diagnosis above.

Remarks.—The name atratus is derived from the Latin atratus (dressed
in black) and refers to the prominent black border of the elytra.

Rhipibruchus psephenopygus Kingsolver, NEW SPECIES
Figs. 30-34, 37
Body length.—2.2-4.3 mm; width, 1.4-1.9 mm. Pronotal length, 0.7-0.9
mm; width, 0.9-1.3 mm.
Color.—Integument yellowish brown to reddish brown or piceous; head
piceous, clypeus and labrum usually reddish, head behind eye reddish, an-

VOLUME 84, NUMBER 4 679

Figs. 30-34. Rhipibruchus psephenopygus. 30, Habitus, dorsal. 31, ¢ genitalia, median
lobe. 32,4 genitalia, lateral lobes. 33, Pygidium d. 34, Pygidium °.

680 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tenna dark reddish brown with segments | and 2 paler beneath, pronotum
mostly piceous fading to reddish brown along anterior border and occasion-
ally along posterior border; elytra with contrasting piceous and reddish-
yellow to reddish-brown variable pattern with humeri and basal umbones
darker, circumscutellar area reddish yellow to brown sometimes extended
to surround umbones, median transverse band piceous divided laterally by
paler blotch, this sometimes small or absent, apical /2 of elytra usually with
oblique pale band extending to lateral margins, apical “3 variable from nearly
all piceous to all mottled; venter of body piceous with some reddish areas
especially on abdomen; legs pale reddish yellow except ventral 2 of metafe-
mur and base of metatibia dusky; pygidium uniformly piceous to black with
depressed lateral spots of 2 slightly darker. Vestiture of black, gray, and
brownish hairs, paler areas of integument generally with gray hairs, piceous
or reddish areas with brown or black hairs; head with brownish hairs, an-
tenna with gray, pronotum with a broad, black median stripe divided by
gray stripe and transversely bisected by grey band, sublateral stripe and
flanks intermixed gray and brown, subbasal gibbosities piceous; scutellum
gray; elytra with pale area gray, darker areas black with gray and brown
intermixed; venter with evenly distributed gray hairs; d pygidium (Fig. 33)
with gray hairs nearly evenly distributed, some specimens with faint median
gray line of hairs; 2 pygidium (Fig. 34) with basal white triangle leading to
intermittent median line of hairs, lateral depressed areas sometimes with
black hairs intermixed with gray; legs evenly covered with gray hairs.
Structure.—Body ovate, base of pronotum narrower than elytral base
(Fig. 30). Head subtriangular; d eyes larger than 2 eyes, ¢ frons narrow;
ocular sinus narrow, about %4 length of eye, carina above base of antenna
prominent; vertex densely, finely punctate, frons strongly carinate arising
dorsally as impunctate, triangular boss, clypeus broader in 6 than in 2,
postocular fringe narrow; d antenna strongly pectinate from 4th segment,
? antenna serrate from 4th segment. Pronotum (Fig. 30) campaniform, lat-
eral margins slightly sinuate, disk strongly convex, with narrow median
sulcus, and shallow longitudinal sulcus extending anteriorly from sinuation
of basal margin, subbasal gibbosities prominent; surface of disk microfo-
veolate, each foveola bearing a hair at its anterior rim, interspaces micro-
punctate; lateral carina fine in basal 42, obsolete apically; cervical sulcus
fine, short; cervical boss prominent, bisetiferous; prosternum short, acutely
triangular medially, separating procoxae for only 2 their length. Scutellum
2x as long as wide, bidentate apically. Elytra quadrilateral, slightly longer
than wide, gently convex except subdepressed along suture, and more
strongly depressed around scutellum; striae slightly sinuate, Ist stria arising
near scutellum, 2nd arising in basal pit beneath oblique marginal ridge, 3rd
and 4th arising from prominent, dentate subbasal gibbosity, Sth arising along

681

VOLUME 84, NUMBER 4

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Figs. 35-36. Distribution of Rhipibruchus spp. 35, R. picturatus. 36, R. prosopis.

682 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lateral border of gibbosity; strial punctures fine, elongated, closely spaced;
3rd, Sth, 7th, and 9th intervals slightly wider than 2nd, 4th, 6th, and 8th;
intervals flat, finely strigose; mesosternum strongly narrowed apically, post-
mesocoxal sulci fine, meeting on midline, inconspicuous behind coxae. Ab-
domen and pygidium as in R. atratus. Pro- and mesolegs not modified,
metacoxa finely punctate except for narrow, polished sulcus; metafemur
incrassate but narrow, dorsal margin more strongly arched in distal /% than
proximal 2, ventral margin nearly straight, pecten consisting of 2 or 3 short
denticles, proximal denticle longer than distal denticles; metatibia slender,
more narrowed at base, gradually broadening distally, mucro short, acute,
lateral denticle small, coronal denticles fine, inconspicuous; lateral, ventral,
and dorsomedial carinae fine but complete to apex, lateroventral carina
obsolete in apical '/s.

Male genitalia (Figs. 31, 32).—Ventral valve of median lobe with elon-
gated apex; internal sac with brush of setae nearly 2 as long as sac; spinous
plate short, broad, with 5—6 spines; median cluster with 5 large, thornlike
spines and 3 smaller denticles; lateral lobes blunt apically, cleft to about 3/s
their length.

Holotype d.—ARGENTINA: La Pampa Prov., Rt. 35, 1 mi N Santa
Rosa, 18 November 1976, from pods Prosopis caldenia.

Allotype 2, 3d, 12 paratypes.—Same data as holotype.

Other paratypes —ARGENTINA: San Juan Prov., Rt. 20, 32 mi SE Cau-
cete, 19 February 1978, in Prosopis alpataco and chilensis, 23; Rt. 20, 3
mi E Caucete, 19 February 1978, in Prosopis torquata, 1°; Rt. 20, 1 mi
SE Caucete, 19 February 1978, in Prosopis alpataco, 1d6.Catamarca Prov.,
Rt. 62, 26 mi S Andalgala, 1 March 1977, in Prosopis flexuosa, 2 3; Rt.
307, 6 km N Santa Maria, 11 January 1980, in Prosopis flexuosa, 13; Rt.
40, 14 mi NE Tinogasta, February 1978, sweeping Larrea cuneifolia, | 2;
Andalgala, 23 October 1973, from Prosopis nigra, 1 3; Andalgala, 25 Oc-
tober 1972, 1d; Colpes, 19 October 1973, from Prosopis nigra, 16. San-
tiago del Estero Prov., Rt. 34, 31 mi SE La Banda, 6 December 1976, in
Prosopis alba, 12. Las Termas, 11 October 1972, Prosopis alba, 13; La
Rioja Prov., Rt. 74, 1 mi SE Chilecito, 30 November 1976, on Acacia aroma,
1d; Aimogasta, 11 February 1978, Prosopis chilensis, 43,52, 25 Feb-
ruary 1978, in Prosopis chilensis, 3 6. Cordoba Prov., Rt. 7, | mi E Cor-
doba-San Luis border, 19 November 1976, in Prosopis caldenia, 4d; Rt.
38, 9 mi SW Cruz del Eje, 24 February 1977, in Prosopis chilensis, 16.
Mendoza Prov., Rt. 40, 8 mi NE Mendoza, 22 February 1977, in Prosopis
alpataco, 1°. San Luis Prov., Rt. 188, 20 mi W La Pampa-S. Luis border,
15 February 1978, in Prosopis caldenia, 12; Rt. 188, 20 mi W La Pampa-
S. Luis border, 17 February 1977, in Prosopis flexuosa, 13, 1 2; Alto
Pencosa, 6 February 1951, 1 2. Lavaisse, Don Roberto Estancia, 17 Oc-

VOLUME 84, NUMBER 4 683

Figs. 37-39. Distribution of Rhipibruchus spp. 37, R. psephenopygus. 38, R. oedipygus
(solid circles); R. rugicollis (open circles). 39, R. atratus.

684 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tober 1942, 1 6; La Pampa Prov., Rt. 188, 38 mi W Realico, 15 February
1978, in Prosopis caidenia, 3 3, 4 2; Rt. 35, 85 mi N Santa Rosa, 15
February 1978, in Prosopis caldenia, 13, 1°. Salta Prov., Oran Salta, 14—
17 August 1917, Prosopis juliflora, 23. BOLIVIA: Tarija Prov., Tarija, 7
July 1980, in seed Prosopis sp., 1°.

Holotype and paratypes deposited in Fundacion Miguel Lillo, Tucuman.
Allotype and paratypes deposited in National Museum of Natural History,
Washington, D.C.; paratypes in Bernardino Rivadavia Museum of Natural
Sciences, Buenos Aires, British Museum (Natural History), London, and
Utah State University, Logan.

Diagnosis.—This species is recognized by the evenly distributed pygidial
vestiture in both sexes and by the broad, transverse dark band across the
middle of the elytra. It is probably most closely related to R. picturatus.

Remarks.—The name psephenopygus is derived from the Greek pse-
phenos (dark, obscure) and pyge (rump) and refers to the dark pygidium.

ACKNOWLEDGMENTS

For supplying specimens, I thank Arturo Teran, Fundacion Miguel Lillo,
Tucuman; Luciano Campos, Universidad de Chile, Santiago; Hugo Cordo,
SEA, USDA, Biological Control of Weeds Laboratory, Buenos Aires; R.
T. Thompson, British Museum (Natural History), London; and W. J. Han-
son, Utah State University, Logan.

I thank Mary Lou Cooley for preparing the illustrations for Figures 1, 4,
5, 25, 28, 29, 30, 33, 34. The remaining illustrations were prepared by the
author. My thanks also go to E. Grissell and C. D. Johnson for manuscript
review and to Phyllis Iglehart for typing the manuscript.

LITERATURE CITED

Blackwelder, R. E. 1946. Checklist of the coleopterous insects of Mexico, Central America,
the West Indies, and South America. U.S. Natl. Mus. Bull. 185(4): 551-763.

Bosq, J. M. 1942. Segundo lista de Coleopteros de la Republica Argentina, daninos a la
agricultura. Ing. Agron. 4(18—22): 1-80.

Bridwell, J. C. 1932. The subfamilies of the Bruchidae (Coleoptera). Proc. Entomol. Soc.
Wash. 34(6): 100-106.

von Fahraeus, O. I. 1839. (New species). Jn Schoenherr, Genera et species Curculionidum,
cum synonymia hujus familiae 5(1): 1-456.

Kingsolver, J. M. 1967. On the genus Rhipibruchus Bridwell, with descriptions of a new
species and a closely related new genus (Coleoptera: Bruchidae: Bruchinae). Proc. Ento-
mol. Soc. Wash. 69(4): 318-327.

Kingsolver, J. M., C. D. Johnson, S. R. Swier, and A. Teran. 1977. Prosopis fruits as a
resource for invertebrates. Jn Simpson, B. B., Mesquite. Its biology in two desert scrub
ecosystems. Dowden, Hutchinson, and Ross, Stroudsburg, Pa. 250 pp.

Philippi, F. H. E. 1859. Algunas especias nuevas de Coleopteros de la Provincia de Valdivia.
Anal. Univ. Chile 16: 656-678.

Pic, M. 1913. Coleopterorum catalogus, pars 55, Bruchidae (Vol. 26): 1-74.

Zacher, F. 1952. Die Nahrpflanzen der Samenkafer. Z. Angew. Entomol. 33(3): 461-480.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 685-689

A NEW SPECIES OF TETANOCERA DUMERIL FROM COLORADO
(DIPTERA: SCIOMYZIDAE)

R. E. ORTH AND T. W. FISHER

Department of Entomology, Division of Biological Control, University of
California, Riverside, California 92521.

Abstract.—A new species, Tetanocera arnaudi, is described from Teller
County, Colorado. It is one of the smaller Tetanocera species found in
North America, and shows some similarities with 7. spreta van der Wulp
(externally) and 7. brevihirta Steyskal (male terminalia). Illustrations of the
new species are included.

The genus Jetanocera is widespread in both the Nearctic and Palaearctic
regions. In North America there are 31 species (including the species de-
scribed herein), 11 of which are Holarctic. The genus is well represented in
collections as these flies are relatively large and easily identified. The body
coloration varies from yellowish to reddish-brown. Many species within the
genus exhibit little diversity in external differences and in many cases dis-
section of the adult terminalia is the only effective means of separation.

The most complete work on North American Tetanocera was by Steyskal
(1959), in which he listed 27 species, of which 8 were described as new.
Illustrations and keys were provided for all species and a phylogenetic
grouping was suggested where relationships could be shown. Since that
work two new species occurring in North America have been described,
Tetanocera andromastos Steyskal (1963), a very small species from Church-
ill, Manitoba, and Tetanocera freyi Stackelberg (1963), a Holarctic species
from the NW European part of the USSR and North America (Alaska).
Stackelberg (1963) also provided extensive illustrations for other members
of the genus in the European part of USSR.

Tetanocera arnaudi Orth and Fisher, NEw Species
Figs. 1-6

Holotype male.—Height of head 7/10 width. Medifacies, parafacies, and
cheeks whitish pruinose with a tinge of yellow. Parafacies with hairs ex-
tending less than halfway from lower margin of eyes to antennal sockets.
Frons yellow, slightly narrowed anteriorly. Midfrontal stripe orangish brown,
extending more or less parallel sided to anterior margin of frons. Ocellar

686 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Tetanocera arnaudi, holotype male. Before terminal segments were excised. Photo
by M. E. Badgley, University of California, Riverside.

triangle and orbital plates with brownish pruinosity. Orbital plates tapered
anteriorly, extending well beyond ocellar triangle. Orbitoantennal spot lack-
ing, narrow strip of white pruinosity along upper orbital margin. Two pairs
of fronto-orbital bristles of approximately equal length; ocellars, postocel-
lars, and inner and outer verticals well developed. Occiput whitish pruinose.
Lateral occipital margins with strong setae and bristles. Antenna testaceous,
segment 2 approximately % length of 3; segment 3 elongate, dorsal margin
slightly concave. Arista black, plumosity of moderate length and density.
Palpi yellowish, labium and labella brownish.

Thorax tannish yellow pruinose; dorsum with brownish longitudinal stripes.
Pleura bare except sternopleuron with fine short hairs over most of the
surface and well-developed bristles ventrally; propleuron with | or more
fine hairs above base of forecoxa. Prosternum bare.

VOLUME 84, NUMBER 4

687

Figs. 2-6. Tetanocera arnaudi, Florissant, Petrified Forest Area, Teller County, Colorado,
2530 m, 11 August 1973, P. H. Arnaud, Jr. 2-5, Paratype, male. 2, Postabdomen, sinistral
view. 3, Hypopygium, rear view. 4, Protandrium, ventral view. 5, Antenna. 6, Allotype, female,
ventral view of terminalia.

688 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Coxae whitish pruinose with some testaceous areas. Legs mostly yellow-
ish, forefemur more testaceous. Foretarsus and apical % of foretibia black;
mid- and hindtarsal segments 4 and 5 black, preceding segments with some
brown. Posterior surface of midfemur with a single strong bristle near tip.

Wing length 4.4 mm. Membrane greyish yellow hyaline with costal brown-
ing; costal margin and veins yellow, slightly testaceous; crossveins arcuate,
infuscated. Halter and squama pale yellow.

Abdominal segments tannish yellow pruinose; postabdomen as in Figs.
2-5.

Allotype female.—Similar to holotype except in abdominal morphology.
Fig. 6. Wing length 5.2 mm.

Holotype.—d , USA, Colorado, Teller County, Florissant, Petrified For-
est Area, 2530 m, 11 August 1973, Paul H. Arnaud, Jr., CAS. California
Academy of Sciences Type no. 13967.

Allotype.— , same data as holotype. Deposited with holotype.

Paratypes.—Same data as holotype (7d, 22); Colorado, Teller County,
21 August 1959, N. Marston (2 3d, 2 2); Colorado, Saguache County, |
August 1937, 9000 feet [2743 m], R. H. Painter (1 3d). At the California
Academy of Sciences, University of California at Riverside, and the Na-
tional Museum of Natural History, Washington, D.C.

Variation.—The specimens at hand show very little variation in color and
size. Wing length of 104, 4.4 to 4.8 mm. Wing length of 4°, 4.8 to 5.4
mm.

Diagnosis.—In Steyskal’s (1959) *‘Key to the American Species of Te-
tanocera,’’ T. arnaudi would be placed close to T. spreta van der Wulp, a
species from the highlands of Mexico. However, examination of the ter-
minalia reveals them to be entirely different. The male terminalia of T.
arnaudi show some similarity to T. brevihirta Steyskal, a species known
only from Fairbanks, Alaska, but the external characters are quite different.
At rubric 10 in Steyskal’s key, 7. spreta and T. arnaudi would be separated
as follows:

10(11). Part of foretibia and all of foretarsus black.

a. Apical '/7 of foretibia black, abdomen with middorsal brown
stripe, terminalia as illustrated (by Steyskal, Fig. 9) .........
wise Spiny Hse Sas tes ergaken tooo VSR ea aE Pr a toa T. spreta van der Wulp

b. Apical % of foretibia black, abdomen dorsally unicolorous, ter-
minalia as illustrated (in our Figs. 2-6) .. 7. arnaudi, new species

Discussion.—Tetanocera arnaudi is one of the smallest of the North
American Tetanocera; perhaps only T. andromastos is smaller. The type-
locality, Florissant, Teller County, is in the center of the state of Colorado
at approximately 39°N latitude. This species is named after Dr. Paul H.
Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San Fran-

VOLUME 84, NUMBER 4 689

cisco, California, who for many years has provided material to further our
studies on the Sciomyzidae.

ACKNOWLEDGMENTS

We extend our thanks to the following people who have given us assis-
tance: P. H. Arnaud, Jr., California Academy of Sciences, San Francisco;
L. Knutson, IIBIII, USDA, Beltsville, Maryland; and G. C. Steyskal, Sys-
tematic Entomology Laboratory, USDA, % National Museum of Natural
History, Washington, D.C.

LITERATURE CITED

Stackelberg, A. A. 1963. Species of the genus Tetanocera Dum. (Diptera: Sciomyzidae) in
the European part of the USSR. Entomol. Rev. 42(4): 492-497.

Steyskal, G. C. 1959. The American species of the genus Tetanocera Dumeril (Diptera).
Pap. Mich. Acad. Sci. Arts Lett. (1958)44: 55-91.

——. 1963. Taxonomic notes on Sciomyzidae (Diptera: Acalyptratae). Pap. Mich. Acad.
Sci. Arts Lett. (1962)48: 113-125.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 690-703

PSALLUS VARIABILIS (FALLEN) AND P. ALBIPENNIS
(FALLEN), TWO EUROPEAN PLANT BUGS ESTABLISHED
IN NORTH AMERICA, WITH NOTES ON TAXONOMIC
CHANGES (HEMIPTERA: HETEROPTERA: MIRIDAE)

A. G. WHEELER, JR. AND E. RICHARD HOEBEKE

(AGW) Bureau of Plant Industry, Pennsylvania Department of Agricul-
ture, Harrisburg, Pennsylvania 17110; (ERH) Department of Entomology,
Cornell University, Ithaca, New York 14853.

Abstract.—The Palearctic phyline mirids Psallus variabilis (Fallén) and
P. albipennis (Fallén) were collected and found established on Long Island,
New York, in 1981-1982. These are the first substantiated records for vari-
abilis in North America and the first published record of a breeding popu-
lation of albipennis in North America. On Long Island, adults and nymphs
of P. variabilis were collected from staminate inflorescences of scrub oak,
Quercus ilicifolia Wang.; adults also were taken from Q. coccinea Muenchh.
and several other plants. Adults and nymphs of P. albipennis were collected
from leaves and stems of common sagewort, Artemisia campestris L.

The adult and fifth-instar nymph of both species are described and illus-
trated; the male genitalia of each species also are illustrated. A review of
the European literature discussing host plants and seasonal history is pre-
sented for both species. In addition, an unpublished record of P. albipennis
from Minnesota is given. Psallus waldeni Knight is proposed as a junior
synonym of P. albipennis, new synonymy; and albipennis is formally trans-
ferred from Plagiognathus to Psallus, new combination.

As part of the USDA-APHIS ‘‘High Hazard Pest Survey’’ program, Hoe-
beke (1980) identified the mirid Psallus variabilis (Fallen), based on a spec-
imen collected at Oyster Bay, Long Island, N.Y. Although Van Duzee (1889)
had reported this European plant bug from Ontario and later (1894) from
New York, Knight (1927), after examining Van Duzee’s specimens, consid-
ered them to represent a native species, Lepidopsallus rubidus var. atricolor
Knight. The Long Island collection thus is the first New World record for
P. variabilis.

We visited the Oyster Bay area in May 1981 and 1982 to confirm the
establishment of P. variabilis and to determine its host range. On the Long

VOLUME 84, NUMBER 4 691

Island trips we also collected the Palearctic Psallus albipennis (Fallén), a
mirid recorded in the New World only from Bradore Bay, Quebec (Moore,
1950). We now confirm that this Artemisia feeder is established in North
America.

In this paper we provide descriptions and illustrations for the adult and
fifth-instar nymph of both species; the male genitalia of each species also
are illustrated. We also propose Psallus waldeni Knight, described from
Connecticut in 1923, as a junior synonym of the Palearctic Psallus albipen-
nis, and transfer albipennis from Plagiognathus to Psallus.

Psallus variabilis (Fallen)

Butler (1923) considered P. variabilis one of the most common British
heteropterans. Although in England it apparently is confined to species of
Populus and Salix (Southwood and Leston, 1959), on the continent it feeds
mainly on oak (Quercus spp.). Kullenberg (1944) recorded this mirid from
Quercus macranthera Fisch. and Mey., Q. petraea L. ex Liebl. (cited as
Q. sessiliflora Salisb.), and Q. robur L. Scudder (1956) also said it is com-
mon on Q. petraea, and Ehanno (1965) recorded populations on Q. robur
(cited as Q. pedunculata Ehrh.) and on Q. ilex L. On oak, Kullenberg
observed that feeding takes place on young shoots and leaf veins, as well
as on catkin stalks and pollen sacs. He noted that when staminate inflores-
cences of oak begin to dry the late instars and adults move to the pistillate
flowers. We believe that the large number of “‘hosts”’ listed in the European
literature (e.g., Stichel, 1956) reflect a movement to other plant species,
especially their inflorescences, when reproductive structures of oak have
shriveled. Kullenberg (1944) and Strawinski (1964) stated that P. variabilis
often supplements its diet with soft-bodied prey occurring on its host plants.

Psallus variabilis is a univoltine species that overwinters in the egg stage:
adults are present from mid-June to early August (Kullenberg, 1944) or may
appear as early as mid-May (Aukema, 1981). The October record of Butler
(1923) must be based on a misidentification. Indeed, the subgenus to which
variabilis belongs, Hylopsallus Wagner, includes several closely related oak
feeders (e.g., perrisi Mulsant) that can be separated reliably only by char-
acters of the male genitalia (Aukema, 1981). Woodroffe (1957) believes that
many of the British records of variabilis should be referred to perrisi.

The first North American record of P. variabilis resulted from the collec-
tion of an adult female in the Planting Fields Arboretum, Oyster Bay, Long
Island, on June 6, 1979; the specimen was taken in a vegetable garden by
K. Uchida as part of the USDA-APHIS ‘High Hazard” survey. Following
the identification of this specimen (by ERH) in June 1980, we made plans
to make additional collections in 1981.

On May 29, 1981, we found a small population of P. variabilis at the
Planting Fields Arboretum; five adults were collected on scarlet oak, Quer-

692 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

cus coccinea Muenchh., growing in a hedgerow. Single specimens were
taken on a nearby red maple, Acer rubrum L., and a quaking aspen, Populus
tremuloides Michx. The next day we collected two adults from the inflo-
rescences of Viburnum recognitum Fernald growing in the same hedgerow,
and two adults at Port Jefferson on a fir, Abies sp., whose needles were
coated with honeydew from an aphid infestation.

We found a larger population of P. variabilis on May 30 at Center Mor-
iches on southeastern Long Island; about 40 adults and six fifth-instar nymphs
were beaten from staminate inflorescences of scrub oak, Q. ilicifolia Wang.
The trees, part of a dense thicket dominated by bayberry, Myrica pennsyl-
vanica Loisel., were growing along a railroad.

In May 1982 we found large numbers of adults on flowers of black cherry
(Prunus serotina Ehrh.), hawthorn (Crataegus sp.), and downy serviceberry
(Amelanchier arborea (Michx.) Fern.) after catkins had largely dropped
from nearby oaks. We collected P. variabilis at these additional localities
in Suffolk Co., Long Island, during May 28-30, 1982: Bayard Cutting Ar-
boretum, Oakdale; Blue Point; and near Tiana Bay.

Our limited observations suggest that P. variabilis is a univoltine, early-
season mirid associated mainly with staminate inflorescences of oaks. Based
on the European literature and our collections, adults on Long Island prob-
ably appear by mid-May and are present only until mid- to late June. In the
Planting Fields Arboretum the staminate flowers had withered and fallen
from most of the scarlet oaks, and the mirid population already may have
begun to decline. Our collection of adults on inflorescences of various plant
species probably reflected a dispersal to an alternate food source. On south-
eastern Long Island at Center Moriches, the scrub oaks still retained their
male catkins although some had withered; the development of mirid popu-
lations on scrub oak in 1981 lagged slightly behind that on scarlet oak at
Oyster Bay.

Psallus variabilis appears well established on Long Island, and the large
numbers found on native scrub oak suggest that it has been present for
several years, if not longer. This mirid is widely distributed on Long Island;
we have seen a specimen collected by Roy Latham from the northern pen-
insula. His unlabeled specimen was among material donated to Cornell Uni-
versity and known to have been collected in the Orient area between 1968
and 1978 when this well-known naturalist was in his 80’s, or nearly 90 (L.
L. Pechuman, personal communication).

Psallus variabilis most likely was introduced with nursery stock imported
from Europe. Long Island, site of the first nursery in the New World (Bailey,
1935; Hedrick, 1950), was an early center of plant importation. Imported
plants were purchased by nurseries, as well as by private estates. In fact,
the Planting Fields Arboretum, originally a private estate, is known to have

VOLUME 84, NUMBER 4 693

been landscaped partly with plants that originated in Europe (G. Clarke,
personal communication).

The following adult description of P. variabilis is taken from Aukema
(1981), who discussed color variation found among populations in the Neth-
erlands and Great Britain. Similar color variation is found among Long
Island populations we examined. Females tend to be reddish brown, males
dark brown or fuscous and more slender.

Adult.—Small (3.3-4.4 mm), oval, dorsal habitus as in Fig. 1, yellow
brown, red brown, bright orange red, sometimes dark brown. Pubescence
consisting of both shiny golden scales and semi-erect black setae.

Head dirty yellow brown to dark brown, sometimes with a reddish tinge.
Antenna yellow brown, with extreme base of segment I more or less dark
or black.

Pronotum light yellow brown to dark brown, tinged with black and some-
times anteriorly and/or posteriorly with dark markings; sometimes (espe-
cially in female) tinged with red or orange red. Mesoscutum and scutellum
colored as pronotum; lateral margins of mesoscutum and scutellum and apex
of scutellum usually somewhat lighter.

Hemelytra light yellow brown to dark brown, especially near apex more
or less tinged with red or orange red. Cuneus yellow brown, red brown to
deep red, basally more or less white along fracture. Wing membrane dark
gray to black, with a clear spot just behind the outer cell. Veins colorless
to grayish, usually tinged with red.

Femora dark brown, broadly red to orange red apically, sometimes with
a few indistinct dark spots beneath. Tibiae light yellow with black spines
arising from distinct dark spots, especially on hindtibia. Tarsi yellow, seg-
ment 3 dark apically.

Ventral surface light red brown to dark brown.

Structural differences of the aedeagus and parameres offer the most de-
pendable characters for separating species of Psallus. The male genitalia of
P. variabilis are briefly described here to facilitate recognition. The right
paramere is large and oblong (Fig. 3a), while the left paramere is also large
with the sensorial process broadly rounded apically (Fig. 3b, c). The ae-
deagus is oblong and robust, with two lateral processes (Fig. 3d); the basal
process is short and hooked at apex.

Kelton (1980) listed approximately 40 species of Psallus occurring in North
America; Knight (1941) treated 13 of these species in a key to the mirid
fauna of Illinois and eastern North America. Psallus variabilis will key near
P. amorphae Knight in Knight’s key, but differs from amorphae mainly by
its lighter dorsal coloration, by its pubescence consisting of shiny golden
scales and semi-erect black setae, and by its distinctive male genitalia (Fig.
3a-d).

694 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. |, 2. Dorsal habitus of adults. 1, Psallus variabilis. 2, P. albipennis.

Fifth-instar nymph.—(In alcohol), Fig. 4. Length 3.0-3.1 mm; background
color pale brown to reddish brown; head, pro-, meso- and metanotum and
wing pads moderately, densely clothed with erect silvery setae; abdomen
moderately clothed with erect pale to dark setae. Head: Length 0.40 mm,
width across eyes 0.67 mm. Rostrum: Length 1|.30-1.34 mm, reaching me-
tacoxae. Antenna: Pale yellowish, extreme base of segment I dark or black,
clothed with fine recumbent setae; segment I, length 0.25—0.26 mm; II, 0.67—
0.70 mm; III, 0.50 mm; IV, 0.30 mm. Pronotum: Length 0.40—-0.50 mm,
median width 0.80 mm, anterior and posterior angles broadly rounded,
somewhat uniformly dark brown to fuscous, especially borders; median line
and other adjacent median areas pale, rather moderately and uniformly
punctured. Meso- and metanotum: Length 0.60-0.67 mm, median width
across wing pads 1.40—-1.47 mm, moderately and uniformly punctured; wing
pads relatively long, reaching slightly beyond apex of abdominal segment
Ill, moderately coarsely punctured. Abdomen tapered to apex, mostly red-
dish; basal segments on mid-dorsal line marked with dark brown; small,
irregular brownish spot on each side of midline on segments V—VIII. Apical
2 abdominal segments brownish dorsally. Dorsal abdominal gland opening
conspicuous along suture between terga 3 and 4; opening (=Type 3 of Ak-
ingbohungbe, 1974) with a sinuate sclerotized bar.

Forefemur dark reddish brown, apex pale; foretibia pale yellow, slightly
darkened apically, clothed with fine, recumbent setae; foretarsus pale
brownish. Mid- and hindfemora reddish brown; mid- and hindtibiae pale

VOLUME 8&4, NUMBER 4 695

Fig. 3. Male genitalia of Psallus variabilis. a, Right paramere. b, c, Opposing views of left
paramere. d, Aedeagus. (All structures reproduced from Aukema, 1981.)

yellow, with black, erect setae arising from black spots. Mid- and hindtarsi
pale brownish. Venter pale reddish to fuscous, extreme lateral areas of
abdominal segments brownish.

Material examined.—2 fifth-instar nymphs, collected from male catkins
of Quercus ilicifolia, near Center Moriches (Suffolk Co.), Long Island, on
May 30, 1981. Determined by association with adults.

696 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

i Pm
zs ez
.
é ‘eS
™. ~\
\
AA >
7? = |
f :
?/ \ \
f Is
a ‘i ! X
j } Veg ee
a;| Me ; me Ye
} x |)
j a
“i Hon
j | Rd
SY
Va j *
/ J
RI (
BS p-R~
4

Fig. 4. Dorsal habitus of Psallus variabilis nymph, fifth instar.

Psallus albipennis (Fallen), NEw CoMBINATION

This common and widely distributed European species has been termed
a ‘‘Holopalearctic’’ mirid (Wagner, 1952). Kullenberg (1944) noted that P.
albipennis is restricted to feeding on plants of the genus Artemisia, with A.
absinthium L. as the principal host, although Kullenberg also observed feed-
ing on A. vulgaris L. and on an ornamental species. He found that the bugs
prefer unopened flower buds but also will feed on leaves and stems. Adults
of this bivoltine mirid are present from mid-July to early September (Kul-
lenberg, 1944), or late June to October (Southwood and Leston, 1959). Kul-
lenberg determined that the egg represents the overwintering stage of this
species.

Psallus albipennis so varies in coloration that at least seven varieties have

VOLUME 84, NUMBER 4 697

been described (Reuter, 1878). Wagner (1941) described two new Artemisia-
feeding species of Plagiognathus that had been confused with albipennis;
since then, several other species, all restricted to Artemisia, have been
described in the subgenus Poliopterus Wagner of Plagiognathus (see Wag-
ner, 1975). In disagreeing with Wagner’s (1941) interpretation, Southwood
and Leston (1959) regarded the extreme morphological forms occurring on
different species of Artemisia as a single, quite variable species. The dif-
ferent forms show ‘‘no constant differences between them,’’ and second
generation adults of albipennis are somewhat larger than those of the first
generation.

The only published North American record of P. albipennis is from Bra-
dore Bay, Quebec, on the easternmost point adjoining Labrador; no dates
or number of specimens were given (Moore, 1950). Carvalho (1958) did not
list a New World record for albipennis in his catalogue of world Miridae.
We found that Moore’s record was based on specimens taken during August
2-8, 1930, by the coleopterist W. J. Brown, who spent the summer of that
year collecting at Thunder Bay and at Bradore Bay ‘‘on the western ex-
tremity of the Straits of Belle Isle’’ (Brown, 1932). Most of his material is
housed in the Canadian National Collection in Ottawa, with the remainder
of the series in the H. H. Knight collection at the National Museum of
Natural History, Washington, D.C. Knight’s collection also contains 32
specimens of P. albipennis from Cook Co. in northeastern Minnesota; this
record, actually predating that from Bradore Bay, was never published. The
data for this early collection of P. albipennis are: Grand Marais, Minnesota,
Aug. 13, 1922, H. H. Knight.

On May 30, 1981, we found a large population of P. albipennis; adults
and a few fourth- and fifth-instar nymphs were feeding on Artemisia cam-
pestris L. growing in dry, sandy soil along a railroad south of Yaphank. One
of us (AGW) and T. J. Henry returned to the same site on August 29 and
found adults and instars II-V. In 1982 we found two adults on A. campestris
in Suffolk Co., near the Suffolk County Air Force Base, NW of Quogue.

It appears that this species breeds throughout the summer and that instead
of the two annual generations reported for Europe there are at least three
on Long Island. We consider P. albipennis to have been introduced from
Europe with man’s commerce. This species can easily be overlooked and
may be more widespread in eastern North America. Crossley (1980), in
explaining the lack of records for Yorkshire, England, pointed out that be-
cause these bugs develop on “‘plants of waste places they often escape the
attention of entomologists!”’

As we were collecting mirids from Artemisia campestris, we suspected
we might be dealing with a European species. We were not aware of any
phylines associated with Artemisia spp. in the eastern U.S. (they are com-
/ mon on these plants in the western states), and Long Island is especially

698 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

liable to accidental introductions with commerce. We described the Long
Island mirid to T. J. Henry, who suggested that it might represent Psallus
waldeni Knight, a species whose identity has long remained in doubt. Knight
(1923) described waldeni from two females taken at New Haven, Connect-
icut, on May 30, 1911. The Connecticut record is cited by Blatchley (1926)
and listed by Carvalho (1958), but no further records are available. If this
species has been collected since 1911, it apparently has not been correctly
identified. After comparing our specimens with Knight’s holotype, Henry
concluded that even though only females of waldeni were available, the
Long Island material was conspecific. We then borrowed European speci-
mens of P. albipennis. After examining and comparing the male genitalia of
the Yaphank specimens with those of the borrowed European specimens,
we concluded that our material was conspecific with the Old World species.
Therefore, we propose a NEw SYNONYMY; Psallus waldeni Knight (1923)
is a junior synonym of P. albipennis (Fallén).

As Henry (1981) pointed out, the generic limits of Plagiognathus and
Psallus are poorly defined; the North American species of both genera are
badly in need of revision. At the time Knight (1923) described waldeni, he
separated the two genera on the basis of simple setae (Plagiognathus) vs.
tomentose or scalelike setae, plus simple setae (Psallus). Because waldeni
has sericeous setae, especially on the head and pronotum, he placed the
new species in Psallus. Because of this setal type, albipennis will key to
Psallus in Knight (1923, 1941) and in some of the European literature. Wag-
ner (1975) uses scalelike setae to separate Plagiognathus from Psallus but
places albipennis in the former genus. The mirid specialists L. A. Kelton
and T. J. Henry feel that albipennis properly belongs in Psallus, and they
have allowed us to formalize this generic transfer. We therefore recognize
the new combination, Psallus albipennis (Fallén).

Adults of P. albipennis are characterized by the following brief descrip-
tion, drawn from the European literature and from specimens at hand.

Adult.—Small (2.8—3.3 mm), elongate-oval, dorsal habitus as in Fig. 2,
color highly variable, usually grayish white but varying from pale to brown-
ish black. Pubescence consisting of dense, long, silvery white setae.

Head dark in front. Antenna with segment I dark or black; segment II
dark or black at base, paler at apex; segments III and IV pale.

Pronotum transverse, blackish; lateral margins slightly rounded, brown
or grayish white. Scutellum blackish.

Hemelytra usually dark. Cuneus pale at base. Wing membrane dusky,
veins and spot below cuneus white or pale.

Hindfemur, and usually also fore- and midfemora, black or blackish brown,
yellow brown at apex. Fore-, mid-, and hindtibiae yellowish, with row of
black spots, each bearing a black seta. All tarsi brownish.

The male genitalia of P. albipennis are characterized as follows: The right

VOLUME 84, NUMBER 4 699

Cc
aa
TaN
Fig. 5. Male genitalia of Psallus albipennis. a, Right paramere. b, c, Opposing views of

left paramere. d, Aedeagus. e, Close-up of apex of aedeagus. (All structures redrawn from
Wagner, 1941.)

700 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

a ON ON
Sy x
C2 a |
a ‘ ae eg ‘~
/ ‘ \
j _+ \
5 lial < ah - Ver
( \ Va eo ee mf ~& » Lg
7 / ~~ |
~ = |
| ¥ \ |
Ae | — ik
Y |
age a / Te
-

Fig. 6. Dorsal habitus of Psallus albipennis nymph, fifth instar.

paramere is moderately large with a small, but prominent subapical protu- |
berance (Fig. 5a), while the left paramere is shaped as in Fig. 5b,c. The
aedeagus is quite long and broadly curved (Fig. 5d); the apical appendage
is short, evenly curved and directed laterally, and the apex of the membra-
nous appendage is pointed and caudally directed (Fig. Se). Psallus albipen-
nis does not key readily to any species in Knight (1941), but will key to

VOLUME 84, NUMBER 4 701

waldeni n. sp. in Knight (1923); waldeni Knight is a junior synonym of P.
albipennis.

Fifth-instar nymph.—(In alcohol), Fig. 6. Length 2.1 mm; background
color pale cream white; head, pro-, meso-, and metanotum, wing pads, and
basal abdominal segments moderately, densely clothed with long, semi-erect
pale setae; apical abdominal segments clothed dorsally with long, semi-
erect, pale, brownish setae. Head: Length 0.30 mm, width across eyes 0.60
mm. Rostrum: Length 1.0 mm, extending to metacoxae; pale white, apex
infuscated. Antenna: slightly brownish, inner margin of segment I at apex
with large dark spot bearing 2 erect dark setae; basal “% of segment II
mottled with small brownish spots; all segments clothed with fine recumbent
setae; segment I, length 0.20 mm; II, 0.48 mm; III, 0.40-0.45 mm; IV, 0.30
mm. Pronotum: Length 0.27-0.30 mm, median width 0.60—0.70 mm, anterior
and posterior margins broadly rounded, each with a very small dark spot
bearing a single seta; surface uniformly cream white. Meso- and metanotum:
Length along median line 0.40-0.60 mm, median width across wing pads 1.0
mm, uniformly pale cream white, apices slightly infuscated, extending near-
ly to apex of abdominal segment IV. Abdomen broadly tapered to rounded
apex, uniformly pale cream white. Dorsal abdominal gland opening small
and inconspicuous along suture between terga 3 and 4; opening (Type 3)
with sinuate sclerotized bar.

Fore-, mid-, and hindfemora pale, each with a large, dorsal, subapical
dark spot bearing a single dark seta, and a smaller, apical, dark spot bearing
2 setae. Fore-, mid-, and hindtibiae pale white, slightly infuscated apically,
with 2 rows of large dark spots along dorsal face bearing dark setae. Fore-,
mid-, and hindtarsi brownish. Venter pale cream white throughout.

Material examined.—8 fifth-instar nymphs, collected from foliage of Ar-
temisia campestris, growing along Rt. 21 south of Yaphank (Suffolk Co.),
Long Island, on May 30, 1981. Determined by association with adults.

ACKNOWLEDGMENTS

We thank L. A. Kelton (Biosystematics Research Institute, Agriculture
Canada, Ottawa) for suggesting the transfer of albipennis from Plagiogna-
thus to Psallus, for allowing us to formalize this new combination, and for
bringing our attention to records of albipennis from Minnesota. We also are
grateful to V. R. Vickery (Lyman Entomological Museum, Ste. Anne de
Bellevue, Quebec) for his personal communications regarding the status of
the G. A. Moore collection, and to T. J. Henry (Systematic Entomology
Laboratory, USDA, Washington, D.C.) for comparing our Long Island ma-
terial with the holotype of Psallus waldeni Knight, for suggesting the pos-
sibility of the new synonymy proposed herein, and for his comments on the
generic transfer for albipennis. Our gratitude is extended to J. A. Slater

702 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(University of Connecticut, Storrs) for the loan of European specimens of
P. albipennis. Robert J. Hill (York College of Pennsylvania, York) provided
the identification of Artemisia campestris. Finally, we are grateful to the
Planting Fields Arboretum, Oyster Bay, Long Island, for permission to col-
lect on the arboretum grounds, to Gene Clarke, arboretum archivist, for
information on the history of the arboretum, and to J. F. Stimmel (Penn-
sylvania Department of Agriculture, Harrisburg) for photographing adults
of both phyline species.

LITERATURE CITED

Akingbohungbe, A. E. 1974. Nymphal characters and higher classification analysis in the
Miridae (Hemiptera: Heteroptera) with a subfamily key based on the nymphs. Can.
Entomol. 106: 687-694.

Aukema, B. 1981. A survey of the Dutch species of the subgenus Hylopsallus of Psallus
(Hemiptera-Heteroptera, Miridae). Tijdschr. Entomol. 124: 1-25.

Bailey, L. H. 1935. The standard cyclopedia of horticulture. Vol. II, Macmillan Co., New
York. Pp. 1201-2421.

Blatchley, W. S. 1926. Heteroptera or true bugs of eastern North America with especial
reference to the faunas of Indiana and Florida. Nature Publ. Co., Indianapolis, Ind.
1116 pp.

Brown, W. J. 1932. Additional notes on the Coleoptera of the north shore of the Gulf of St.
Lawrence. Can. Entomol. 64: 198-209.

Butler, E. A. 1923. A biology of the British Hemiptera-Heteroptera. H. F. & G. Witherby,
London. 682 pp.

Carvalho, J.C. M. 1958. Catalogue of the Miridae of the world. Pt. IJ. Subfamily Phylinae.
Arch. Mus. Nac., Rio de J. 45: 1-216.

Crossley, R. 1980. Some interesting insects at Tag Lock, Elland. Naturalist (Leeds) 105(955):
159-160.

Ehanno, B. 1965. Notes ecologiques sur les Miridae (Insecta-Heteroptera) observes en Bre-
tagne sur le chene. Vie Milieu 16: 517-533.

Hedrick, U. P. 1950. A history of horticulture in America to 1860. Oxford University Press,
New York. 551 pp.

Henry, T. J. 1981. A new eastern United States Psallus Fieber (Heteroptera: Miridae) from
Physocarpus (Rosaceae). Proc. Entomol. Soc. Wash. 83: 399-402.

Hoebeke, E. R. 1980. A mirid bug (Psallus variabilis (Fallen)). Coop. Plant Pest Rpt. 5(33):
628.

Kelton, L. A. 1980. The insects and arachnids of Canada. Part 8. The plant bugs of the
prairie provinces of Canada (Heteroptera: Miridae). Biosystematics Research Institute,
Ottawa, Publ. 1703, 408 pp.

Knight, H. H. 1923. Family Miridae (Capsidae), pp. 422-658. Jn Britton, W. E., ed., The
Hemiptera or sucking insects of Connecticut. Conn. State Geol. Nat. Hist. Surv. Bull.
34.

—. 1927. On the Miridae in Blatchley’s ‘‘Heteroptera of Eastern North America.”’ Bull.

Brooklyn Entomol. Soc. 22: 98-105.

1941. The plant bugs, or Miridae, of Illinois. Ill. Nat. Hist. Surv. Bull. 22: 1-234.
Kullenberg, B. 1944. Studien tiber die Biologie der Capsiden. Zool. Bidr. Upps. 23: 1-522.
Moore, G. A. 1950. Check-list of Hemiptera of the Province of Quebec. Contrib. Inst. Biol.

Univ. Montreal 26: 1-49. [Reprinted Nat. Can. (Que.) 77: 233-271.]

VOLUME 84, NUMBER 4 703

Reuter, O.M. 1878. Hemiptera Gymnocerata Europae. Hemipteres Gymnocerates d’ Europe,
du bassin de la Mediterranee et de |’ Asie russe. I. Acta Soc. Sci. Fenn. 13: 1-188.

Scudder, G. G. E. 1956. A contribution to a survey of the distribution of the Hemiptera-
Heteroptera of Wales. Entomol. Mon. Mag. 92: 54-64.

Southwood, T. R. E. and D. Leston. 1959. Land and water bugs of the British Isles. Frederick
Warne, London & New York. 436 pp.

Stichel, W. 1956. Illustrierte Bestimmungstabellen der Wanzen. II. Europa. 10. Heft, pp.
289-320.

Strawinski, K. 1964. Zoophagism of terrestrial Hemiptera-Heteroptera occurring in Poland.
Ekol. Pol. (A)12: 429-452.

Van Duzee, E. P. 1889. Hemiptera from Muskoka Lake District. Can. Entomol. 21: I-11.

1894. A list of the Hemiptera of Buffalo and vicinity. Bull. Buffalo Soc. Nat. Sci. 5:

167-204.

Wagner, E. 1941. Zwei neue deutsche Plagiognathusarten (Hem. Miridae). Stett. Entomol.
Ztg. 102: 248-257.

———. 1952. Die Tierwelt Deutschlands, 41. Teil: Blindwanzen oder Miriden. Verlag von
Gustav Fischer, Jena. 218 pp.

——. 1975. Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen
Inseln (Hemiptera, Heteroptera). Entomol. Abh. Mus. Tierk. Dresden 40(suppl.): I-
483.

Woodroffe, G. E. 1957. A preliminary revision of the British Psallus Fieber (Hem., Miridae),
with a description of a new species. Entomol. Mon. Mag. 93: 258-271.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 704-715

SYNOPSIS OF THE HILAROGRAPHINI (LEPIDOPTERA:
TORTRICIDAE) OF THE WORLD

JOHN B. HEPPNER

Department of Entomology, Smithsonian Institution, Washington, D.C.
20560.

Abstract.—A summary of the world Hilarographini (Lepidoptera: Tortri-
cidae: Chlidanotinae) is provided, with distribution notes, host records, and
a world checklist.

The Hilarographini were formerly a group of genera in Glyphipterigidae
sensu Meyrick (1914) but have been shown in recent years to belong to the
Tortricidae (Heppner, 1977, 1978). The tribe was described by Diakonoff
(1977a) and placed in the Chlidanotinae. The genital characters of Hilaro-
graphini, as well as other characters, show relationships to other Chlida-
notinae, in fact, bridging previous gaps between Schoenotenini and Polyor-
thini. The latter tribe may even require redefinition inasmuch as one of the
main distinctions of the Polyorthini (split valvae with valval coremata, in
the male genitalia) is also found among many Hilarographini species. Tor-
tricoid abdominal apodemes, among other characters, distinguish the Hila-
rographini from Glyphipterigidae, and the naked haustellum clearly distin-
guishes the tribe from Choreutidae, a family also recently segregated from
Glyphipterigidae and one with many species superficially resembling Hilaro-
graphini species.

The tribe currently includes 78 species in 9 genera. Although 9 genera are
now in the tribe, it is possible that Mictopsichia Hubner (possibly also
Embolostoma Diakonoff and Mictocommosis Diakonoff) should be placed
in the tribe Archipini of the subfamily Tortricinae (by genital characters),
but this requires further study. The distribution of the 78 included species
can be seen by referring to the map (Fig. 1). Each dot represents the type-
locality of a described species of Hilarographini except for a few localities
that have a number of species described from approximately the same area
and where all the dots cannot be included. Multiple type-localities are found
in the following areas: Assam, India (8 spp.); Java, Indonesia (8 spp.); Sri
Lanka (4 spp.); Surinam (5 spp.); Amazonas, Brazil (4 spp.). Table | gives
a summary of the species of Hilarographini in each faunal region of the

VOLUME 84, NUMBER 4 705

world. Most species are currently known only from the type-locality, but
many species should have a wider distribution in adjacent tropical forests
or other suitable habitats (encircled on map).

Host records for Hilarographini include only those for 5 species: Thau-
matographa caminodes (Meyrick), from rhyzomes of Cardamon sp. (Zin-
giberaceae); Thaumatographa cubensis Heppner, from Pinus cubensis Gri-
sebach (Pinaceae); Thaumatographa eremnotorna (Diakonoff and Arita),
from the cambium of Pinus densiflora Siebold and Zuccerini (Pinaceae);
Thaumatographa oenobapta Diakonoff, from the inflorescence of /xora sp.
(Rubiaceae); and Thaumatographa regalis (Walsingham), from the cambium
of Pinus sabiniana Douglas and Pinus ponderosa Douglas (Pinaceae). Other
species like Thaumatographa youngiella (Busck) and Thaumatographa
jonesi (Brower) have been collected in or near pine forests. The Neotropical
species are diurnally active, and it may be that most members of the tribe
are diurnally active.

The checklist format includes the original generic combination at the end
of each citation, with the type-locality noted to the far right. Localities in
brackets are presumed type-localities where the original author did not spe-
cifically indicate the locality.

GENERIC SYNOPSIS OF HILAROGRAPHINI

Embolostoma Diakonoff, 1977 |
Mictopsichia Hubner, [1825] 11
Mictopsychia [sic] Riley, 1889, missp.
Mictropsichia [sic] Heppner, 1978, missp.
Mictocommosis Diakonoff, 1977 4
Trianassa Meyrick, 1905 4
Idiothauma Walsingham, 1897 4
Nexosa Diakonoff, 1977 4
Charitographa Diakonoff, 1979 |
s

Thaumatographa Walsingham, 1897 3
Tharmatographa [sic] Diakonoff, 1977, missp.
Hilarographa Zeller, 1877 17
TORTRICIDAE: CHLIDANOTINAE
HILAROGRAPHINI
Embolostoma Diakonoff, 1977b: 51 (Type-species: E. plutostola Diakonoff,
1977). .
plutostola Diakonoff, 1977b: 52. Indonesia (Java)

Mictopsichia Hiibner, [1825]: 374 (Type-species: M. hubnerana [sic] Hub-
ner, [1825], missp. [=Phalaena (Tortrix) hubneriana Stoll, 1787}).
Mictopsychia [sic] Riley, 1889: 158, missp.

706 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

SCALE ON EQUATOR

° 1000 2000 3000 MILES
t n h

+ T i 1
° {000 2000 3000 4000 KILOMETERS

MERCATOR PROJECTION

=

Fig. 1. Distribution of Hilarographini by type-localities (shaded areas are known distribu-
ests on land areas).

VOLUME 84, NUMBER 4 707

oO EAST LONGITUDE

tions of North American species; encircled areas represent approximate limits of tropical for-

708 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 1. Faunal region distribution of Hilarographini. N = Nearctic; NL = Neotropical;
P = Palearctic; E = Ethiopian; O = Oriental; A = Australian; OC = Oceania.

Genus N NL P E O A OC Total
Embolostoma l 1
Mictopsichia 11 11
Mictocommosis l 2 l 4
Trianassa 3 | 4
Idiothauma 3 1 4
Nexosa 3 l 4
Charitographa | 1
Thaumatographa 3 I 5 16 > 2 32
Hilarographa 17 17
Faunal totals 3 29 vi 5 25 6 3 78

Mictropsichia [sic] Heppner, 1978: 53, missp.
callicharis Meyrick, 1921b: 477. Brazil (Amazonas)
durranti Walsingham, 1914: 304. Brazil (Para)
fuesliniana (Cramer, 1781: 163) ([Phalaena (Tortrix)]). [Surinam |
fueslynialis (Cramer, 1782: 249 [index])(Phalaena (Pyralis)), emend.
fueslyniana [sic] (Verloren, 1837: 138)(Pyralis), missp.
fuesslyana Walsingham, 1914: 304, emend.

gemmisparsana (Walker, 1863: 415) (Gauris). Brazil (Amazonas)
godmani Walsingham, 1914: 305. Mexico (Tabasco)
hubneriana (Stoll, 1787: 41) (Phalaena (Tortrix)). [Surinam]
hubnerana [sic] Hubner, [1825]: 374, missp.
superba Felder and Rogenhofer, 1875: 13.
miocentra Meyrick, 1920: 330. Brazil (Para)
ornatissima (Dognin, 1909: 94) (Gauris) Peru
pentargyra Meyrick, 1921b: 478. Peru
periopta Meyrick, 1913a: 99. Guyana
renaudalis (Stoll, 1787: 42) (Phalaena (Pyralis)). [Surinam]

Mictocommosis Diakonoff, 1977b: 8 (Type-species: Simaethis nigromacu-
lata Issiki, 1930).
argus (Walsingham, 1897: 54) (Mictopsichia), NEW COMBINATION.
Rep. Congo
microctenota (Meyrick, 1933: 370) (Mictopsichia), NEW COMBINATION.
Sierra Leone
nigromaculata (Issiki, 1930: 423) (Simaethis). Japan
takaonis (Matsumura, 1931: 1081) (Simaethis).
stemmatias (Meyrick, 192la: 178) (Mictopsichia), NEW COMBINATION.
Indonesia (Celebes)

VOLUME 84, NUMBER 4 709

Figs. 2-7. Examples of typical Hilarographini. 2, Mictopsichia hubneriana, 3 (7.5 mm),
Panama [USNM]. 3, M. fuesliniana, 6 (8 mm), Brazil [USNM]. 4, M. ornatissima, 3 (9.5
mm), Bolivia [ZMHB]. 5, Idiothauma africanum, 3 (6 mm), Spanish Guinea [ZMHB]. 6,
Thaumatographa zapyra, 2 (7 mm), New Guinea [USNM]. 7, Hilarographa swederiana, ¢
(9 mm), Costa Rica [USNM]. Lengths are forewing base to apex dimensions.

710 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Irianassa Meyrick, 1905: 609 (Type-species: J. sapphiropa Meyrick, 1905).
poecilaspis Meyrick, 1923: 616. India (Assam)
sapphiropa Meyrick, 1905: 609. Sri Lanka
speciosana (Pagenstecher, 1900: 225) (Grapholitha [sic}).
Papua New Guinea (Bismarck Is.)
alcyonopa Meyrick, 1926: 301.

uranopa Meyrick, 1927: 101. Samoa

Idiothauma Walsingham, 1897: 49 (Type-species: /. africanum Walsingham,
1897).

africanum Walsingham, 1897: 50. Rep. Congo

druidicum (Meyrick, 1909: 428) (Hilarographa), NEW COMBINATION.
India (Assam)

malgassicellum Viette, 1958: 112. Madagascar
rigatiellum (Ghesquiere, 1940: 30) (Hilarographa), NEW COMBINATION.
Zaire

Nexosa Diakonoff, 1977b: 12 (Type-species: Mictopsichia marmarastra
Meyrick, 1932).

aureola Diakonoff, 1977b: 15. Papua New Guinea
hexaphala (Meyrick, 1912a: 36) (Mictopsichia). Sri Lanka
marmarastra (Meyrick, 1932: 273) (Mictopsichia). Indonesia (Java)
picturata (Meyrick, 1912a: 35) (Mictopsichia). India (Assam)

Charitographa Diakonoff, 1979: 291 (Type-species: Hilarographa mikadonis
Stringer, 1930).

mikadonis (Stringer, 1930: 418) (Hilarographa). Japan

micadonis [sic] (Diakonoff and Arita, 1976: 193) (Hilarographa), missp.

Thaumatographa Walsingham, 1897: 52 (Type-species: Hilarographa zapyra
Meyrick, 1886).
Tharmatographa [sic] Diakonoff, 1977b: 51, missp.
aurosa (Diakonoff and Arita, 1976: 181) (Hilarographa). Japan
calathisca (Meyrick, 1909: 427) (Hilarographa), NEW COMBINATION.
India (Assam)
caminodes (Meyrick, 1905: 610) (Hilarographa), NEW COMBINATION.
Sri Lanka
ceramopa (Meyrick, 1920: 329) (Hilarographa), NEW COMBINATION.
India (Assam)
cirrhocosma (Meyrick, 1930: 1) (Hilarographa). Solomon Is.
citharistis (Meyrick, 1909: 428) (Hilarographa), NEW COMBINATION.
India (Assam)
cladara Diakonoff, 1977b: 33. Indonesia (Borneo)
cubensis Heppner, in press. Cuba
decoris (Diakonoff and Arita, 1976: 191) (Hilarographa).
USSR (Kuril Is.)

VOLUME 84, NUMBER 4 711

dolichosticha Diakonoff, 1977b: 32. Indonesia (Java)
eremnotorna (Diakonoff and Arita, 1976: 185) (Hilarographa). Japan
excellens Pagenstecher, 1900: 230. Papua New Guinea (Bismarck Is.)
pyranthis (Meyrick, 1907: 91) (Hilarographa).
ferox (Meyrick, 1921a: 179) (Hilarographa). Indonesia (Java)
hermatodes (Meyrick, 1909: 426) (Hilarographa), NEW COMBINATION.
Sri Lanka
jonesi (Brower, 1953: 96 (Hilarographa). USA (Massachusetts)
leucopyrga (Meyrick, 1912a: 36) (Hilarographa). Japan
ludens (Diakonoff, 1948: 200) (Hilarographa). Indonesia (Buru)
macaria Diakonoff, 1977b: 46. Indonesia (Java)

machaerophora (Diakonoff and Arita, 1976: 180) (Hilarographa). Japan

mechanica (Meyrick, 1909: 427) (Hilarographa), NEW COMBINATION.
India (Assam)

merinthias (Meyrick, 1909: 426) (Hilarographa), NEW COMBINATION.
India (Assam)

mesostigmatias Diakonoff, 1977b: 42. Taiwan
oenobapta Diakonoff, 1977b: 28. Indonesia (Java)
opistocapna Diakonoff, 1977b: 38. Indonesia (New Guinea)
phlox Diakonoff, 1977b: 40. Indonesia (Java)
regalis (Walsingham, 1881: 320) (Glyphipteryx [sic]). USA (California)

spermatodesma (Diakonoff, 1955: 17) (Hilarographa).
Indonesia (New Guinea)

tetralina (Meyrick, 1930: 1) (Hilarographa). Solomon Is.
tornoxena Diakonoff, 1977b: 50. Indonesia (Java)
~ undosa Diakonoff, 1977b: 45. Indonesia (New Guinea)

|

:

youngiella (Busck, 1922: 278) (Hilarographa). Canada (British Columbia)
olympica (Braun, 1923: 118) (Hilarographa).
zapyra (Meyrick, 1886: 286) (Hilarographa). Papua New Guinea

Hilarographa Zeller, 1877: 187 (Type-species: Phalaena (Tortrix) sweder-
iana Stoll, 1790: 75).

aenigmatica (Meyrick, 1912b: 682) (Cnephasia). Colombia
bellica Meyrick, 1912a: 37. Surinam
bryonota Meyrick, 1921b: 479. Peru
dulcisana (Walker, 1863: 415) (Gauris). Brazil (Amazonas)
dulciana Meyrick, 1913b: 24, emend.
| eriglypta Meyrick, 1921b: 478. Peru
euphronica Meyrick, 1920: 328. Brazil (Para)
hexapeda Meyrick, 1913a: 99. Guyana
methystis Meyrick, 1921b: 479. Peru
orthochrysa Meyrick, 1932: 274. Brazil (Santa Catarina)

plectanodes Meyrick, 1921b: 480. Peru

712 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

plurimana (Walker, 1863: 416) (Gauris). Brazil (Para)
quinquestrigana (Walker, 1866: 1796) (Carpocapsa). Brazil (Sao Paulo)
firmana (Felder and Rogenhofer, 1875: 5) (Carpocapsa).

refluxana (Walker, 1863: 416) (Gauris). Brazil (Rio de Janeiro)
ribbei (Zeller, 1877: 189) (Setiostoma). Panama
swederiana (Stoll, 1790: 75) (Phalaena (Tortrix)). [Surinam]
trabeana (Felder and Rogenhofer, 1875: 11) (Grapholitha [sic]).
thaliarcha Meyrick, 1920: 328. Brazil (Para)
xanthotoxa Meyrick, 1920: 329. Brazil (Amazonas)
ACKNOWLEDGMENTS

Preliminary research resulting in this summary was partially made pos-
sible by a grant from the National Science Foundation (76-12550 DEB) and
the University of Florida, Department of Entomology and Nematology
(IFAS), Gainesville, Florida (while at this institution), for study of collec-
tions at the British Museum (Natural History), London, England, in 1976—
77. | also acknowledge the support of the Smithsonian Institution (USNM).
Humboldt Universitat, East Berlin (ZMBH) provided some specimens.

LITERATURE CITED

Braun, A. F. 1923. Microlepidoptera: notes and new species. Trans. Am. Entomol. Soc. 49:
115-127.

Brower, A. E. 1953. Three new species of microlepidoptera (Olethreutidae, Glyphipterygidae
and Yponomeutidae). Ann. Entomol. Soc. Am. 46: 95-98.

Busck, A. 1922. Microlepidoptera from British Columbia. Can. Entomol. 53: 276-280.

Cramer, P. 1780-1782. De Uitlandsche Kapellen voorkomende in de drie waereld-deelen
Asia, Africa en America. Vol. 4. Baalde, Amsterdam. 252 pp., pls. 289-400.

Diakonoff, A. 1948. Fauna Buruana. Microlepidoptera II. Treubia 19: 197-219.

1955. Microlepidoptera of New Guinea. Results of the third Archbold Expedition
(American-Netherlands Indian Expedition 1938-1939). Pt. 5. Verh. Koninkl. Nederl.
Akad. Wetenschappen, Afd. Natuurk (Tweede Reeks) 50(3): 1-212.

—. 1977a. Description of Hilarographini, a new tribus in the Tortricidae (Lepidoptera).
Entomol. Ber. (Amsterdam) 37: 76-77.

—. 1977b. Rearrangement of certain Glyphipterygidae sensu Meyrick, 1913, with de-
scriptions of new taxa (Lepidoptera). Zool. Verh. (Leiden) 158: I-55.

—. 1979. Descriptions of new taxa of the so-called Glyphipterigidae auctorum (Lepi-
doptera). Zool. Meded. (Leiden) 54: 291-312.

Diakonoff, A. and Y. Arita. 1976. New and old species of Glyphipterigidae and a new species
of Acrolepiidae from Japan and the Far East (Lepidoptera). Zool. Meded. (Leiden) 50:
179-219.

Dognin, P. 1909. Hetéroceres nouveaux de l’ Amerique du Sud. Ann. Soc. Entomol. Belg.
53: 74-94.

Felder, R. and A. F. Rogenhofer. 1875. Jn, Felder C., R. Felder, and A. F. Rogenhofer,
Reise der Osterreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859
unter den Behilfen des Commodore B. von Wiillerstorf-Urbair. Zoologischer Theil.
Zweiter Band. Zweite Abtheilung: Lepidoptera. Vienna. 20 pp., pls. 108-140.

i tT ee

a

VOLUME 84, NUMBER 4 743

Ghesquiere, J. 1940-42. Catalogue raisonnés de la faune entomologique du Congo Belge.
Lépidopteres, microlépidopteres. Tervuren, Belgium. 240 pp., 6 pls.

Heppner, J. B. 1977. The status of the Glyphipterigidae and a reassessment of relationships
in yponomeutoid families and ditrysian superfamilies. J. Lepid. Soc. 31: 124-134.

—. 1978. Transfers of some Nearctic genera and species of Glyphipterigidae (auctorum)

to Oecophoridae, Copromorphidae, Plutellidae, and Tortricidae (Lepidoptera). Pan-Pac.

Entomol. 54: 48-55.

. Inpress. Revision of North American Hilarographini (Lepidoptera: Tortricidae), witha

new species from Cuba. Pan-Pac. Entomol.

Hiibner, J. 1816-[1825]. Verzeichniss bekannter Schmettlinge [sic]. Augsburg. 431 pp.

Issiki, S. T. 1930. New Japanese and Formosan microlepidoptera. Ann. Mag. Nat. Hist. (10)
6: 422-431.

Matsumura, S. 1931. 6000 illustrated insects of Japan-Empire. Japan. 1496 pp.; 191 pp.,
index.

Meyrick, E. 1880. Descriptions of Lepidoptera from the South Pacific. Trans. Entomol. Soc.
Lond. 1886: 189 -296.

1905. Descriptions of Indian micro-lepidoptera. I. J. Bombay Nat. Hist. Soc. 16

580-619.

1907. Descriptions of Australasian micro-lepidoptera. XIX. Plutellidae. Proc. Linn.

Soc. N. S. W. 32: 47-150.

1909. Descriptions of Indian micro-lepidoptera. IX. J. Bombay Nat. Hist. Soc. 19

410-437.

1912a. Glyphipterygidae. Exotic Microlepid. 1: 35-63.

1912b. Descriptions of South American micro-lepidoptera. Trans. Entomol. Soc.

Lond. 1911: 673-718.

1913a. Glyphipterygidae. Exotic Microlepid. 1: 98-104.

1913b. Carposinidae, Heliodinidae, Glyphipterygidae. Jn Wagner, H., ed., Lepidop-

terorum catalogus. Part 13. W. Junk, Berlin. 53 pp.

1914. Lepidoptera Heterocera. Fam. Glyphipterygidae. Jn Wytsman, M. P., ed.,

Genera insectorum. Fasc. 164. Brussels. 39 pp., 2 pls.

1920. Glyphipterygidae. Exotic Microlepid. 2: 325-338.

1921a. New microlepidoptera. Zool. Meded. (Leiden) 6: 145-202.

1921b. Glyphipterygidae. Exotic Microlepid. 2: 477-480.

1923. Glyphipterygidae. Exotic Microlepid. 2: 616-619.

1926. Glyphipterygidae. Exotic Microlepid. 3: 301-308.

1927. Micro- eEeeD ie In Insects of Samoa and other Samoan terrestrial Arthro-

alah Lepidoptera. Fasc. 2, Part 3, pp. 65-116.

1930. Ciohinenedae Exotic Microlepid. 4: I-7.

1932. Glyphipterygidae. Exotic Microlepid. 4: 273-276.

1933. Glyphipterygidae. Exotic Microlepid. 4: 370-372.

Pagenstecher, A. 1900. Die Lepidopterenfauna des Bismarck-Archipels. Mit Berticksichti-
gung der thiergeographischen und biologischen Verhiltnisse systematisch dargestellt.
Zweiter Theil: Die Nachtfalter. Zoologica (Stuttgart) 12(29): 1-268, 2 pls.

Riley, C. V. 1889. Two brilliant and interesting microlepidoptera new to our fauna. Proc.
Entomol. Soc. Wash. 1: 155-159.

Stoll, C. 1787-1791. Aanhangsel van het Werk, de Uitlandsche Kapellen, voorkomende in
de drie Waereld-deelen Asia, Africa en America, door den Heere Pieter Cramer, ver-
vattende naauwkeurige afbeeldingen van Surinaamsche Rupsen en Poppen; als mede
van veele zeldzaame en nieuwe ontdekte Uitlandsche Dag-en Nagt-Kapellen. [vol. 5].
Gravius, Amsterdam. 184 pp., 42 pls.

Stringer, H. 1930. New species of microlepidoptera in the collection of the British Museum.
Ann. Mag. Nat. Hist. (10)6: 415-422.

Pree tte

| i i | | |

714 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Verloren, H.

1837. Catalogus systematicus lepidopterorum, quae in opere Crameri descripta

sunt, secundum methodum Latreillii. J. Altheer, Utrecht. 280 pp.

Viette, P. E. L.
111-124.

1958. Microlepidoptera nouveaux de Madagascar. Rev. Fr. Entomol. 25:

Walker, F. 1863. List of the specimens of lepidopterous insects in the collection of the British
Museum. Part XXVIII. Tortricites & Tineites. British Museum, London. Pp. 288-561.

1866. List of the specimens of lepidopterous insects in the collection of the British

Museum. Part XXXV. Supplement.—Part 5. British Museum, London. Pp. 1535-2040.

Walsingham, T. de G. 1881.
1881: 301-325; pls. 35-36.

1897: 33-67; pls. 2-3.

On some North American Tineidae. Proc. Zool. Soc. Lond.
1897. Western equatorial African micro-lepidoptera. Trans. Entomol. Soc. Lond.

1914. Hemerophilidae, pp. 300-319. Jn Tineina, Pterophorina, Orneodina and Pyr-

alidina and Hepialina (part). Jn Godman, F. D., and O. Salvin, eds., Biologia Centrali-
Americana. Insecta. Lepidoptera-Heterocera. Vol. 4. London. 482 pp., 9 pls.

Zeller, P. C.

1877. Exotische microlepidoptera. Hor. Soc. Entomol. Ross. 13: 3-493, 6 pls.

ALPHABETICAL LIST OF SPECIES NAMES

aenigmatica (Meyrick), Hilarographa

africanum Walsingham, Idiothauma

alcyonopa Meyrick, Irianassa

argus (Walsingham), Mictocommosis

aureola Diakonoff, Nexosa

aurosa (Diakonoff and Arita), Thaumatogra-
pha

bellica Meyrick, Hilarographa

bryonota Meyrick, Hilarographa

calathisca (Meyrick), Thaumatographa

callicharis Meyrick, Mictopsichia

caminodes (Meyrick), Thaumatographa

ceramopa (Meyrick), Thaumatographa

cirrhocosma (Meyrick), Thaumatographa

citharistis (Meyrick), Thaumatographa

cladara Diakonoff, Thaumatographa

cubensis Heppner, Thaumatographa

decoris (Diakonoff and Arita), Thaumatogra-
pha

dolichosticha Diakonoff, Thaumatographa

druidicum (Meyrick), Idiothauma

dulciana Meyrick, Hilarographa

dulcisana (Walker), Hilarographa

durranti Walsingham, Mictopsichia

eremnotorna (Diakonoff and Arita), Thau-
matographa

eriglypta Meyrick, Hilarographa

euphronica Meyrick, Hilarographa

excellens Pagenstecher, Thaumatographa

ferox (Meyrick), Thaumatographa

firmana (Felder and Rogenhofer), Hilarogra-
pha

fuesliniana (Cramer), Mictopsichia

fueslyniana [sic] (Verloren), Mictopsichia

fueslynialis (Cramer), Mictopsichia

fuesslyana Walsingham, Mictopsichia

gemmisparsana (Walker), Mictopsichia

godmani Walsingham, Mictopsichia

hermatodes (Meyrick), Thaumatographa

hexapeda Meyrick, Hilarographa

hexaphala (Meyrick), Nexosa

hubnerana [sic] Htibner, Mictopsichia

hubneriana (Stoll), Mictopsichia

jonesi (Brower), Thaumatographa

leucopyrga (Meyrick), Thaumatographa

ludens (Diakonoff), Thaumatographa

macaria Diakonoff, Thaumatographa

machaerophora (Diakonoff and Arita), Thau-
matographa

malgassicellum Viette, Idiothauma

marmarastra (Meyrick), Nexosa

mechanica (Meyrick), Thaumatographa

merinthias (Meyrick), Thaumatographa

mesostigmatias Diakonoff, Thaumatographa

methystis Meyrick, Hilarographa

micadonis [sic] (Diakonoff and Arita), Char-
itographa

microctenota (Meyrick), Mictocommosis

mikadonis (Stringer), Charitographa

miocentra Meyrick, Mictopsichia

VOLUME 84, NUMBER 4

nigromaculata (Issiki), Mictocommosis
oenobapta Diakonoff, Thaumatographa
olympica (Braun), Thaumatographa
opistocapna Diakonoff, Thaumatographa
ornatissima (Dognin), Mictopsichia
orthochrysa Meyrick, Hilarographa
pentargyra Meyrick, Mictopsichia
periopta Meyrick, Mictopsichia

phlox Diakonoff, Thaumatographa
picturata (Meyrick), Nexosa
plectanodes Meyrick, Hilarographa
plurimana (Walker), Hilarographa
plutostola Diakonoff, Embolostoma
poecilaspis Meyrick, Irianassa
pyranthis (Meyrick), Thaumatographa
quinquestrigana (Walker), Hilarographa
refluxana (Walker), Hilarographa
regalis (Walsingham), Thaumatographa
renaudalis (Stoll), Mictopsichia

ribbei (Zeller), Hilarographa

715

rigatiellum (Ghesquiere), Idiothauma

sapphiropa Meyrick, Irianassa

speciosana (Pagenstecher), Irianassa

spermatodesma (Diakonoff), Thaumatogra-
pha

stemmatias (Meyrick), Mictocommosis

superba Felder and Rogenhofer, Mictopsi-
chia

swederiana (Stoll), Hilarographa

takaonis (Matsumura), Mictocommosis

tetralina (Meyrick), Thaumatographa

thaliarcha Meyrick, Hilarographa

tornoxena Diakonoff, Thaumatographa

trabeana (Felder and Rogenhofer), Hilarogra-
pha

undosa Diakonoff, Thaumatographa

uranopa Meyrick, Irianassa

xanthotoxa Meyrick, Hilarographa

youngiella (Busck), Thaumatographa

zapyra (Meyrick), Thaumatographa

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 716-725

TAXONOMY AND ETHOLOGY OF A NEW CENTRAL AMERICAN
SPECIES OF ROBBER FLY IN THE GENUS
GLAPHYROPYGA (DIPTERA: ASILIDAE)

Eric M. FISHER AND HENRY A. HESPENHEIDE

(EMF) Department of Entomology, University of California, Riverside,
California 92521; (HAH) Department of Biology, University of California,
Los Angeles, California 90024.

Abstract.—Glaphyropyga dryas, from Panama and Costa Rica, is de-
scribed and illustrated. The genus Glaphyropyga Schiner is characterized
by species with a very narrow face, a slender third antennal segment with
a short to long arista, lack of postscutellar pile, and possession of a gray
microvillous shadow in the anteroapical portion of the wing. Known species
of Glaphyropyga share the habitat preference of G. dryas and range from
eastern Mexico to southern Brazil. Adults of G. dryas are active during the
rainy season and inhabit the shaded forest understory, perching on twig tips
0.5 to 2.0 m above ground. Prey records include nine orders of insects, with
Diptera, Coleoptera, Hymenoptera, and Homoptera accounting for 94% of
recorded captures. The order Zoraptera is reported for the first time as prey
of an asilid. Size of prey ranges from 1.1 to 6.5 mm and averages 3.07 mm
for 101 measured items. Courtship consists of the male’s hovering and bob-
bing, with hindlegs extended below, to the side of the perched female. Cop-
ulating pairs assume a tail-to-tail position on a branch tip.

The tropical forests of Central America possess a very rich and diverse
robber fly fauna. Field studies by the authors on Barro Colorado Island,
Panama, indicate that more than 70 species of Asilidae occur at this small
(15.6 km*), tropical moist forest site (Hespenheide and Fisher, unpublished
data). About half of these species are undescribed—a legacy of both the
species richness of Neotropical forests and the very limited taxonomic at-
tention so far given to Central American robber flies. An undescribed species
of Glaphyropyga Schiner is one of the most abundant asilids on Barro Col-
orado Island. The purpose of this paper is to describe this species and to
present observations on the taxonomy and biology of it and of the genus
Glaphyropyga. The taxonomic portion of this paper was prepared by EMF,
the ethological portion primarily by HAH.

VOLUME 84, NUMBER 4 717

Glaphyropyga dryas Fisher, NEW SPECIES
Figs. 1-6

Diagnosis.—A small species, most closely related to G. himantocera
(Wiedemann), differing from that species as follows: Male with conspicuous,
dorsal-median, setigerous protuberance on epandrium; female with a group
of 6 apically convergent, black bristles at apex of tergum 7.

Holotype male.—Length 12.1 mm. Head yellowish brown anteriorly, black
dorsally and posteriorly. Face '/10 width of head at middle, very densely
silvery tomentose above, thinly yellowish gray tomentose below at base of
proboscis; front with tomentum yellowish brown laterally, black medially;
vertex dark brown tomentose, more yellowish posteriorly, a polished black
spot anterior to ocellar tubercle; occiput with tomentum mostly gray, dark
brown next to vertex; mystax silvery white, with 10 long, prominent bristles
on gibbosity, arranged in a triangular fashion—2 vertical rows of 3 slender
bristles above, which diverge ventrad, and 2 pairs of strong bristles below
on oral margin, the middle pair stoutest—plus 6 short, slender bristles on
oral margin; a few short, yellow hairs laterally on frons, a brown pair on
ocellar tubercle; slender, subproclinate occipital bristles yellowish dorsally,
white laterally and ventrally; beard white. Proboscis black, palpi yellow,
the sparse hair white.

Antenna (Fig. 1) 1.8 mm long, 3rd segment about 3% x as long as arista
plus microsegment; segments |, 2, and extreme base of 3 brownish yellow,
remainder blackish brown (including sub-apical ring on 2); strong ventroap-
ical bristle on | and setae on | and 2 yellowish white; segment 3 dark brown
tomentose dorsally, yellowish brown ventrally; arista glabrous.

Thorax black, yellowish brown posteroventrally on pleurae. Mesonotum
(Fig. 3) with tomentum dark brown on broad central stripe (slightly paler
along midline) and intermediate spots, golden brown on dorsocentral and
lateral vittae, grayish brown on humeri, postalar calli and posterior declivity
(latter with median, dark brown spot above scutellum), gray along extreme
lateral margins; long, stout bristles black, distributed as follows: 5 pair of
dorsocentrals, 2 pair of notopleurals, | pair of supraalars, | pair of postalars;
5 pair of slender dorsocentrals, 6 pair of anterior acrostichals and very
sparse bristles on anterior and posterior declivities all black and short (sub-
equal in length to antennal segment 2); sparse, scattered pile on humeri and
lateral margins white. Scutellum gray tomentose, more brownish centrally
on disk; very sparse, short diskal pile white, | pair stout, long marginal
bristles black. Pleurae and postscutellar slopes whitish gray tomentose, with
pronotum dorsally and anepisternum posterodorsally brown tomentose; pile
short, sparse, white; laterotergal fan of 6 to 8 stout bristles white.

Legs very pale brownish yellow, translucent, the foreleg almost white,
except these areas dark brown: Anterodorsal stripe and apical 4 hindfemur,

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

718

VOLUME 84, NUMBER 4 719

dorsum of basal 3 and entire apical “4 hindtibia, and apical /% of Sth segment
hindtarsus; very narrow anterodorsal stripe and apical '/; midfemur, and
segments 4 and 5 of midtarsus; segments 3 to 5 of foretarsus. Bristles and
pile nearly all whitish yellow on pale areas of legs and brown to black on
dark areas, but dorsal surfaces of all femora with short, sparse, recumbent
black hair and tibiae and tarsi with scattered, minute black setulae; pos-
teroventral surfaces of fore- and midfemora and foretibia with sparse, erect
pile about as long as width of midfemur; remainder of vestiture typical for
genus. Claws black, with extreme bases red. Coxae yellowish brown, with
whitish-gray tomentum and white pile.

Wings (see Fig. 2) 8.9 mm long; membrane very faintly infuscate, slightly
darker along posterior margin; gray shadow of microvilli extending across
anterior 4/s of apex of Ist submarginal cell (R2,3;). Halter brownish yellow,
edges of knob darker.

Abdomen black, narrow lateral and posterior margins of terga 1-5 yellow-
ish brown; terga 1-7 with black areas dark brown tomentose, paler areas of
terga 1-5 yellowish brown tomentose; sterna 1-6 grayish brown tomentose,
sternum 7 thinly so; narrow lateral margin of tergum 7 and all of tergum and
sternum 8 and genitalia polished black; short, recumbent dorsal pile black,
longer bristles yellow, ventral pile and bristles yellow. Genitalia as in Figs.
4-6: epandrium near middle with prominent dorsomedian protuberance,
which bears at apex a clump of about 6 short, stout bristles; apical /2 of
epandrium, beyond protuberance, angled slightly ventrad; gonopods with
basistyli rounded apically; hypandrium with apical margin slightly concave;
aedeagus reddish brown, visible basal portion with oval-shaped ventral
depression; bristles and pile whitish yellow, some recumbent pile at epan-
drium base black.

Allotype female —Body length 11.1 mm; wing length 9.3 mm. Similar to
male, differing as follows: Face with plane portion pale yellowish and gib-
bosity grayish white tomentose; 3 pair of dorsal bristles in mystax brown.
Antenna with basal 2 segments brown tomentose, with brown bristles and
black hair. Mesonotum with anterior acrostichal bristles shorter, subequal
to % length of antennal segment 2. Legs much darker, chiefly blackish
brown, with these areas yellowish: Ventral margins of basal % of all femora
(extending onto dorsum at bases of mid- and hindfemora), dorsum of mid-
tibia from before middle to apical 3/4, and foretibia except apical '/5 and
narrow posterior stripe; vestiture nearly all black, except large bristles at

—

Figs. 1-6. Glaphyropyga dryas. |, Antenna of female. 2, Lateral view of female. 3, Meso-
notum and scutellum. 4-6, male genitalia. 4, Lateral view (ep = epandrium; hy = hypandrium;
ba = basistylus; di = distylus; ae = aedeagus). 5, Dorsal view. 6, Ventral view. Scales in mm.

720 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

base of hindfemur and ‘‘cleaning pads’’ on fore- and hindlegs yellowish.
Coxae with some anterior bristles brown. Abdomen with segments 7 and 8
and narrow lateral margin of tergum 6 polished black; apex of tergum 7 with
file of 6 black bristles which converge and cross apically. Ovipositor with
short, yellow pile.

Distribution.—Known from the Isthmus of Panama and Cartago and Her-
edia Provinces, Costa Rica.

Type specimens.—Holotype d (USNM): Barro Colorado Island, Canal
Zone, Panama, 11/13 June 1976 (E. M. Fisher). Allotype 2 (USNM), same
data as for holotype. Paratypes (55 d, 113 2). PANAMA: Canal Zone:
Barro Colorado Island, 436, 63 2, 9 July to 11 Sept. 1974, 2/3 Aug. 1975,
8/20 June 1976, 10 May, 29 July 1977, 9 July to 12 Aug. 1978 (H. A. Hes-
penheide and/or E. M. Fisher); 9 km NW Gamboa, 3°, 4/12 Aug. 1975 (E.
M. and J. L. Fisher); Madden Forest, mi 3.5, 2 ¢, 4 July 1977 (H. A.
Hespenheide); Chiva Chiva Rd., 8 air km N Fort Clayton, 22, 23 July 1978
(E. M. Fisher, N. E. Woodley, A. Worthington); | km E Farfan, 4d, 20
2, 31 July, 11 Aug. 1975 (E. M. Fisher); 9 km NW Gatun, 292, 9 Aug. 1975
(E. M. Fisher); Devil’s Beach, 5 km W Fort Sherman, | 6, 2°, 9/10 Aug.
1975 (E. M. Fisher). Panama Prov.: Bayano R. Bridge on Hwy 2, nr Chepo,
12, 12 Sept. 1974 (H. A. Hespenheide); Pacora, Sta. C, 1¢, 12, 9 July
1951 (F. Miller); Cerro Azul, N of Tocumen, | 2, 7 June 1958 (W. J. Han-
son); Cerro Campana, 800-860 m, 4¢, 18 2, 26 June, 13 July 1977 (H. A.
Hespenheide), 17 July, 27 July, 19 Aug. 1978 (E. M. Fisher, N. E. Woodley).

Paratypes to be deposited in the National Museum of Natural History,
Washington, D.C. (USNM); California Academy of Sciences, San Francis-
co; Natural History Museum of Los Angeles County, California; American
Museum of Natural History, New York; University of California, Riverside;
British Museum (Natural History), London; Naturhistorisches Museum, Vi-
enna; the collections of the authors and Joseph Wilcox, and at other appro-
priate institutions and collections.

Other specimens examined.— COSTA RICA: Cartago: 5 km E Turriabla,
C.A.T.1.E., 66, 369, 21/22 July 1975 (E. M. and J: L. Fisher); Turriabla,
12,9 July 1965 (G. R. Noonan). Heredia: Finca La Selva, 3 km S Puerto
Viejo, 22, 23/25 July 1976 (E. M. Fisher, H. A. Hespenheide).

The Costa Rican specimens agree with the Panamanian material in all
respects but are not being made paratypes.

Discussion.—The specific name is derived from the Greek Dryas (f.—a
wood nymph) and refers to the habitat of this species.

Variation among available specimens of G. dryas is very slight. Most
notable are differences in body length, males ranging from 10.3 to 13.0 mm
long, females from 10.2 to 13.2 mm. Many males have the lateral margins
of tergum 6 narrowly polished, instead of entirely tomentose. About a quar-
ter of the female specimens have a faint trace of the oval white spot near

VOLUME 84, NUMBER 4 721

the apex of the first submarginal cell, a characteristic found in G. himan-
tocera and several other Glaphyropyga species; however it is never as dis-
tinct and opaque in G. dryas.

The shape of the male epandrium and the file of bristles on tergum 7 of
females are characteristics unique to G. dryas. The combination of the fol-
lowing characters are also useful for distinguishing this species from the
other known (described and undescribed) species of Glaphyropyga: Small
size (10.2 to 13.2 mm body length); form of the mystax (four stout bristles
in-line along oral margin, six slender bristles above, in two dorsally con-
verging rows of three each, leaving a triangular area in the center of the
gibbosity bare of long bristles); arista and microsegment together being less
than 4 the length of antennal segment 3; wing with shadow extending ex-
actly to posterior '/s of first submarginal cell and, in females, lacking a
conspicuous, oval, opaque white spot. Glaphyropyga dryas is most similar
to the Brazilian G. himantocera, which is a larger species (14.5-17.0 mm
long; Carrera, 1945) and has non-protuberant, non-angulate epandria (par-
allel sided in lateral view). Carrera (1950) described G. aristata from Barro
Colorado Island; it is similar in size and form to G. himantocera but has
the arista nearly as long as the third antennal segment.

GENERAL REMARKS ON THE GENUS GLAPHYROPYGA

Taxonomy.-—The genus Glaphyropyga was placed in the Asilinae by Hull
(1962) and the Apocleinae by Papavero (1973). It may be recognized by the
_ slender form of its species, their sparse pilosity and dark brown dorsal
| tomentum, the very narrow face with ventrally placed gibbosity and mystax,
the narrow and elongate third antennal segment, and the lack of postscu-
tellar pile. In addition, all known species of the genus have a gray micro-
villous shadow in the apical half of the subcostal cell of the wing (and usually
the marginal and first submarginal cells also).
Hull (1962) emphasized the very short arista of the type-species, G. hi-
mantocera, aS a means of recognizing the genus. Earlier Hull (1958) de-
| scribed the genus Opopotes to include a new species, O. attenuatus Hull,
| with an elongate arista (in describing Opopotes, Hull evidently overlooked
Carrera’s previous description of G. aristata, as this species obviously would
fit his concept of the new genus). Papavero (1973) listed Opopotes as a
synonym of Glaphyropyga; we agree with this synonymy. Tapinostylus En-
| derlein is also synonymous with Glaphyropyga (Hull, 1962; Martin and Pa-
pavero, 1970). Species of Glaphyropyga have also been incorrectly placed
_ in Senoprosopis Macquart, a quite unrelated genus, but one whose members
have a similarly slender habitus. Curran’s figures 53 and 137 (Curran, 1934:
170, 180), identified as Senoprosopis sp., are actually of G. dryas.
As now understood, Glaphyropyga contains species with an arista length
(including the short, basally attached microsegment) of about '/; to sub-

T22 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

equal the length of the third antennal segment. Although the species of
Glaphyropyga vary as to length of arista, they are otherwise a taxonomically
and biologically cohesive and well characterized group. An arbitrary divi-
sion of those species with a longer arista into another genus is unwarranted.

Six species of Glaphyropyga are currently known in the literature, and
the genus is reported to range from Brazil to Costa Rica (Martin and Pa-
pavero, 1970). In addition to G. dryas, at least eight undescribed Middle
American species are on hand. One of these occurs in the cloud forests of
Hidalgo, Mexico, some 2000 km northwest of Costa Rica. We anticipate the
discovery of many additional species in the genus, especially in South Amer-
ica.

Biology.—Probably all species of Glaphyropyga (11 species have been
observed by us in the field) occur in well forested habitats at low to moderate
elevations. The genus is apparently restricted to older second growth and
‘‘primary’’ tropical forest, and is absent from badly disturbed forest and
early-successional second growth areas. Glaphyropyga species invariably
perch on slender, bare tips of branches, at heights of one to several meters.
They are seldom found in open, fully sunlit areas and instead prefer the
partial to complete shade of the forest understory. Edges of trails and open-
ings in the understory with abundant, but always indirect, light seem to be
optimal habitat. On very cloudy or rainy days they may be seen on perches
in clearings, and some species are quite active during rains.

ETHOLOGY OF GLAPHYROPYGA DRYAS

Microhabitat and foraging behavior.—Glaphyropyga dryas has been ob-
served over most of Barro Colorado Island, and is the most common robber
fly of the forest interior (Hespenheide and Fisher, unpublished data), ac-
counting for about 25% of all flies observed in trailside censuses. Collection
and observation records range from 10 May to 11 September and indicate
that this species is active only during the rainy season, which on Barro
Colorado Island is typically from early May through mid-December (Croat,
1978). For the small number of flies for which perch characteristics were
recorded, most were observed in closed-canopy situations, but a significant
proportion were also observed in the vicinity of sunflecks or at the shaded
margins of larger openings created by fallen trees. Of 40 flies for which
perch type was recorded, 36 were on the tips of bare twigs of understory
shrubs or fallen branches, 2 were on tips of leaves, and 2 on vine tips or
tendrils. Of 27 recorded perch heights, 3 were at or below about 0.5 m above
ground, 12 between 0.5 and 1.5 m, 10 between 1.5 and 2.0 m, and 2 above
2.0 m. These heights undoubtably reflect the bias of our observational meth-
od toward flies at or near eye level, but they do differ significantly from
those of some of the other typical forest understory robber fly species.
Because a portion of our recorded perch sites and heights were of unusual

VOLUME 84, NUMBER 4 123

Table 1. Sizes of prey taken by Glaphyropyga dryas.

eee
Number of prey

Taxon Taken Measured Mean (mm) +SD Range
Coleoptera 32. 31 2.63 1.053 1.1-5.9
Diptera 36 28 2.86 1.048 1.4-5.3
Hemiptera 2 2 4.15 2.899 2.1-6.2
Homoptera 15 13 4.18 1.769 1.3-6.5
Hymenoptera 21 20 3.28 1.192 1.7-5.4
Isoptera Il I 2.70 — —
Lepidoptera 4 4 3.65 1.493 1.7-5.2
Psocoptera l 1 1.90 — _
Zoraptera l I 1.20 — —

Total 113 101 3.07 1.308 1. 1-6.5

or extreme situations, the overall qualitative microhabitat of the species can
be characterized as shaded forest understory, with preferred perches being
twig tips between 1.5 and 2.0 m above ground.
_ In addition to foraging sites, we also collected 113 prey items from or
together with flies. These represented nine orders of insects, of which Dip-
tera (36 prey, 32%), Coleoptera (32, 29%), Hymenoptera (21, 18%), and
_ Homoptera (15, 14%) predominated. The following taxa were included among
these: COLEOPTERA: Cerambycidae (1), Chrysomelidae (3), Curculioni-
| dae (1, Cossinae), Phalacridae (1), Rhynchitidae (1), Scarabaeidae (1), Scol-
| ytidae (8), Staphylinidae (5), undetermined (11). DIPTERA: Ceratopogonidae
| (1), Culicidae (1), Drosophilidae (2), Empididae (2), Mycetophilidae (5),
Phoridae (5), Psychodidae (1), Sciaridae (15), Sphaeroceridae (1), Stratio-
myidae (1), Tipulidae (1), undetermined (1). HEMIPTERA: Cydnidae (1),
Lygaeidae (1). HOMOPTERA: Achilidae (2), Cercopidae (1), Cicadellidae
(5), Delphacidae (2), Kinnaridae (?, 2), Meenoplidae (?, 1), Fulgoroidea (1),
undetermined (1). HYMENOPTERA: Formicidae (19, all reproductives, in-
| cluding the following taxa: Azteca sp. d (1), Myrmelachista sp. 3 (2),
Pachycondyla sp. 3 (3), Paratrechina sp. 2° (1); Ponerinae genus Ad (1),
kgenus Bo (1), genus Co (2); Solenopsis sp. Ad (1), sp. B@ (1), sp. C
2 (1), sp. D@ (1)), Tiphiidae (1), undetermined (1). ISOPTERA: undeter-
)mined (1). LEPIDOPTERA: undetermined moths (4). PSOCOPTERA: un-
determined (1). ZORAPTERA: Zorotypidae (1). Of these, the zorotypid is
especially noteworthy, apparently being the first record of this infrequently-
collected group of insects as prey of an asilid. Sizes of these prey are pre-
sented in Table 1.
Courtship and mating behavior.—Courtship behavior was observed by
both of us in detail on separate occasions, by EMF on 12 June 1976 between
10:20 and 10:40 h on a sunny day, and by HAH on 23 June 1977 somewhat

724 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

before noon, in both instances along William Morton Wheeler trail between
the 1400 and 1600 m posts. Courtship was also informally observed and
photographed by HAH on other occasions in early July 1977.

Display sites were at typical forging sites, in the understory of closed-
canopy forest and (on 23 June 1977) at a shaded broken-canopy site at a
partially-overgrown treefall. In all cases females were observed perched on
twig tips at heights of 1.5 to 2.0 m above ground. Males hover to the right
or left side of the female at the same height, on an axis perpendicular to
her, and at a distance of I-3 cm. Although generally constant in position,
the male moves slowly toward and away from the female over a range of
1-3 cm and oscillates vertically over a range of a few mm with an intensity
that varies from slow (usually) to very rapid or agitated. While hovering the
male holds the fore- and midlegs folded in toward the thorax, fully extends
the hindlegs ventrally and occasionally rubs the hindtarsi together. The po-
sition of the hindlegs is significant with respect to their coloration, especially
as seen by the female; only the whitish-yellow color would be visible to the
female as the dark areas would be oriented away from her. Because the
forelegs would also show only yellow and all the coxae are very pale, the
male would be perceived as a bobbing, pale object against the relatively dim
and therefore dark understory background.

Copulation is attempted by the male | to 3 times per minute by striking
the female from the side and attempting to couple. No successful copulation
was seen by us. Females would either leave the perch, and thus end the
courting sequence, or fly out 2-3 cm toward the male in a manner similar |
to a sally for a prey item, although over a much shorter distance than typical.
After a few to several minutes of unsuccessful courtship, the male flies off
to a nearby perch by means of an erratic, bobbing flight, although with no |
audible sounds. After an interval of a few minutes, the male may again court
the same female. |

On 12 June 1976, a pair of G. dryas was seen in copula shortly afterwards |
and close by the pair observed courting, but evidently not the same indi- |
viduals. The two flies were in a tail-to-tail orientation with the female at the
tip of the branch facing outwards and the male behind her and facing away.

ACKNOWLEDGMENTS

We are very grateful to the following organizations for their generous
support of our field work in Central America: National Science Foundation
(award DEB76-10109 to HAH); The Explorers Club (Exploration Fund grant |
obtained with the assistance of Donald Patterson and the California Acad- |
emy of Sciences); and the Department of Biology, California State Univer-
sity, Long Beach. We are also indebted to the staffs of the Smithsonian |
Tropical Research Institute in Panama and the Organization for Tropical |
Studies in Costa Rica for their assistance and use of facilities. We thank the |

VOLUME 84, NUMBER 4 725

following individuals and institutions for the loan of certain specimens used
in this study: California Academy of Sciences (Paul Arnaud, Jr.); Natural
History Museum of Los Angeles County (Charles Hogue, Roy Snelling); and
Joseph Wilcox, Anaheim, California. Roy Snelling kindly identified the ants.

LITERATURE CITED

Carrera, M. 1945. Estudio sobre os generos Glaphyropyga e Senoprosopis con descricao de
novo genero e novas especies. Pap. Avulsos Zool. (Sao Paulo) 5: 175-192.

—. 1950. Uma nova especies de Glaphyropyga da zona do canal no Panama (Diptera,
Asilidae). Dusenia 1: 27-32.

Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford Univ. Press, Stanford, Calif.

943 pp.

Curran, C. H. 1934. The families and genera of North American Diptera. New York.
512 pp.

Hull, F. M. 1958. More flies of the family Asilidae (Diptera). Ann. Mag. Nat. Hist. (12) 10:
884-895.

—. 1962. Robber flies of the world. The genera of the family Asilidae. U.S. Natl. Mus.
Bull. 224 (Pt. 2): 433-907.

Martin, C. H. and N. Papavero. 1970. Family Asilidae. Jn A catalogue of the Diptera of the
Americas south of the United States. Mus. Zool. Univ. Sao Paulo (35b): 1-139.
Papavero, N. 1973. Studies of Asilidae (Diptera) systematics and evolution. I. A preliminary

classification in subfamilies. Arg. Zool. (Sao Paulo) 23: 217-274.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 726-728

A NEW SPECIES OF TRIOXYS (HYMENOPTERA: APHIDITDAE)
PARASITIC ON A PECAN APHID

PETR STARY AND PAUL M. MARSH

(PS) Institute of Entomology, Czechoslovak Academy of Science, Ceske

Budejovice, Czechoslovakia; (PMM) Systematic Entomology Laboratory, |
IIBIII, Agricultural Research Service, USDA, Beltsville Agricultural Re- |

search Center, Beltsville, Maryland, 2070S.

Abstract.—Trioxys monelliopsis, n. sp., a parasitoid of Monelliopsis ni-

gropunctata (Gran.) on pecan, is described from Georgia. This is the first |

record of parasitism of this aphid in the field.

Parasitoids of aphid pests on pecan were first studied in connection with |

biological control of Chrompahis juglandicola (Kaltenbach) on walnut. The
imported parasitoid Trioxys pallidus (Haliday) was found to parasitize Mo-
nellia costalis (Fitch) in the laboratory but not in the field (Schlinger et al.,
1960; van den Bosch et al., 1962).

The parasitoid complex associated with the foliar-feeding aphids Monellia |

costalis, Monelliopsis nigropunctata (Granovsky), and Tinocallis caryae-
foliae (Davis) on pecan was studied by Tedders (1977, 1978). Tedders suc-

cessfully reared in the laboratory two introduced species, Trioxys pallidus |
and T. complanatus Quilis, but neither parasitoid species became estab- |

lished in the field. However, an unknown Trioxys species was found to

parasitize M. nigropunctata in the field. Subsequent examination has shown |

that it represents a new, apparently indigenous species, which is described
below. Thus, M. nigropunctata has been found to be parasitized by three

Trioxys species in the United States: T. pallidus and T. complanatus in the |

laboratory (establishment not yet documented), and 7. monelliopsis, n. sp.,
in the field.

Trioxys (Trioxys) monelliopsis Stary and Marsh, NEW SPECIES

This species is easily distinguishable from other Nearctic species of Triox- |

ys by characters on the prongs of the last abdominal sternum in the female.
These prongs are hairless dorsally and bear a single claw-shaped apical
bristle (Figs. 5, 6). Recently, a closely related species of Trioxys, a parasit-
oid of a Myzocallis species, has been described from Mexico (Stary and

VOLUME 84, NUMBER 4 727

Figs. 1-6. Trioxys (T.) monelliopsis, 2 paratype. 1, Forewing. 2, Tergum |. 3, Mesonotum.
4, Propodeum. 5, Genitalia. 6, Apex of the prong, detail.

Remaudiere, in press). This species is distinguishable from 7. monelliopsis
by the shape of the apical bristle on the prongs. The bristle is ovoid-shaped
to droplike in the Mexican species and claw-shaped in 7. monelliopsis. In
Stary’s (1978) key to European species, 7. monelliopsis will run to couplet
14 but is distinguished from 7. curvicaudus Mackauer and T. phyllaphidis
Mackauer by the hairless dorsal surface on the abdominal prongs.
Female.—Eyes large. Gena equal to '/7 of eye length. Tentorial index
| (i.e., tentorio-ocular line over inter-tentorial line, relative length) 0.3. An-
_tenna |1-segmented, slightly thickened to apex, reaching to about middle of
abdomen. Mesonotum (Fig. 3) with sparse hairs. Propodeum (Fig. 4) dis-
tinctly areolated, central areola somewhat irregular in shape. Forewing (Fig.
1) with pterostigma almost 3x as long as wide, metacarpus equal to about
Y. pterostigma length, and radial vein reaching about %4 of its possible length.

728 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Tergum | (Fig. 2) twice as long as wide across spiracles. Spiracular tubercles
distinct, situated before middle. Distance between spiracles and apex of
tergum | distinctly longer than width across spiracles. Genitalia (Fig. 5) with
prongs slightly arcuate, hairless dorsally, and with | claw-shaped apical
bristle (Fig. 6).

Head brown, clypeus and mouthparts yellow brown. Scape brown, yel-
lowish at apex. Pedicel and flagellomere | yellow, 2 mostly yellowish, rest
brownish and darkened to apex. Thorax brown. Wings hyaline, venation
brownish. Legs brown, trochanters, base of tibiae and tarsi (except dark
specimens) yellowish. Abdomen brown, tergum | and triangular spot at base
of tergum 2 yellowish, ovipositor sheaths light brownish, prongs yellowish.

Body length about 1.4 mm.

Male.—Unknown.

Holotype female.— Byron, Georgia, 12-VIII-1975, W. L. Tedders, ex
Monelliopsis nigropunctata (Gran.) on pecan. Deposited in the National
Museum of Natural History, Washington, D.C. (USNM).

Paratypes.—2 2, same data as holotype, deposited in USNM and in col-
lection of P. Stary, Czechoslovakia.

Remarks.—The studies of Tedders (1977, 1978) show that this species is
apparently specific to M. nigropunctata as it was not reared from other |
aphid species on pecan despite intensive rearing activities. Furthermore, it
seems to be a rare species on the target host in the observed area. The
mummy is yellowish white. Of the three mummies available, two were alate
adults and one was a fourth-instar immature aphid.

ACKNOWLEDGMENTS

We are indebted to W. L. Tedders (SE Fruit and Tree Nut Laboratory,
USDA, Byron, Ga.) for supplying us material and information on the para- |
sitoid species.

LITERATURE CITED

van den Bosch, R., E. I. Schlinger, and K. S. Hagen. 1962. Initial field observations in |
California on Trioxys pallidus (Haliday), a recently introduced parasite of the walnut
aphid. J. Econ. Entomol. 55: 857-862.
Schlinger, E. I., K. S. Hagen, and R. van den Bosch. 1960. Imported French parasite of
walnut aphid established in California. Calif. Agric. 14: 3-4.
Stary, P. 1978. Parasitoid spectrum of the arboricolous callaphidid aphids in Europe (Hy-
menoptera, Aphidiidae; Homoptera, Aphidoidea, Callaphididae). Acta Entomol. Bo-
hemoslov. 75: 164-177.
Stary, P. and G. Remaudiere. Jn press. New genera, species, and host records of aphid |
parasitoids (Hym., Aphidiidae) from Mexico. Ann. Soc. Entomol. Fr. (N.S.).
Tedders, W. L. 1977. Trioxys pallidus and Trioxys complanatus as parasites of Moenllia
costalis, Monelliopsis nigropunctata and Tinocallis caryaefoliae. Ann. Entomol. Soc.
Am. 70: 687-690.
——. 1978. Important biological and morphological characteristics of the foliar-feeding
aphids on pecan. U.S. Dep. Agric. Tech. Bull. 1579, 29 pp.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 729-741

THE MAYFLIES OF NORTHEASTERN MINNESOTA
(EPHEMEROPTERA)'

T. M. Lacer, M. D. JOHNSON, AND W. P. MCCAFFERTY

(TML) 636 State Street, Menasha, Wisconsin 54952; (MDJ) P.O. Box 368,
Teton Village, Wyoming 83025; (WPM) Department of Entomology, Purdue
University, West Lafayette, Indiana 47907.

Abstract.—Analysis of the Ephemeroptora of northeastern Minnesota is
based primarily on extensive sampling of larvae from St. Louis and Lake
counties. Of 54 identifiable and nominal species, 33 represent new state

records. Several notable range extensions are documented. The fauna has

a strong affinity with that of eastern Canada; other faunistic relationships
are also present. In Minnesota, Baetis macani bundyae Lehmkuhl larvae
inhabit streams and complete their life cycle one and a half months earlier
than larvae in the northern extreme of their range where they inhabit tundra

_ ponds; larvae and adults are similar in size throughout their range.

cS — a —

The Ephemeroptera of Minnesota, especially in the northeastern region,
are largely unknown. Although Daggy surveyed the mayfly fauna throughout
the state (Daggy, 1938, 1941), he published only a small part of the data
(Daggy, 1945). Thomas Say, who accompanied an expedition on the Min-
nesota and Rainy rivers in the 1820’s, was probably the first to report on
the state’s mayflies (Say, 1823). Burks (1953) documented Minnesota dis-
tributions for species found in Illinois. Additional records occur in system-
atic works (Banks, 1910; Needham et al., 1935; Lewis, 1974; McCafferty,
1975; Edmunds et al., 1976; Berner, 1978; Bednarik and McCafferty, 1979;
Morihara and McCafferty, 1979b; Pescador and Berner, in press) and in
ecological studies of certain species (Waters, 1966; Fremling, 1973; Waters
and Crawford, 1973; Hall et al., 1975).

In this paper we document the mayflies of northeastern Minnesota, pre-
dict those species likely to occur in the northern region of the state, show
faunal relationships to other areas of North America, and discuss the effi-
cacy of factors which may influence the faunal distributions. Larvae were

1 Part of the Minnesota Regional Copper-Nickel Study, Environmental Quality Board, St.
Paul, Minnesota.

730 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

extensively sampled in St. Louis and Lake counties located in northeastern
Minnesota (Fig. 1) during 1976 and 1977 by the following persons: M. D.
Johnson, T. M. Lager, J. L. McCulloch, M. W. Mischuk, and S. N. Wil-
liams. Dr. L. Berner identified adult mayflies collected by the late W. V.
Balduf, University of Illinois, at Eagles Nest Lake from 1953 to 1959 and
made the data available to us. All collections from Eagles Nest Lake are
credited to Dr. Balduf.

DESCRIPTION OF STUDY AREA

The Precambrian Shield, composed of metamorphic rocks and covered
by glacial till and drift, extends into this region (Wright, 1972). The glacial
history is evident in the generally flat but locally rugged topography. Many
lakes formed in the glacial ice scoured troughs, and numerous streams drain
surrounding ridges and lowlands. The Laurentian Divide partitions the area |
into northern and southern sections (Fig. 1). Streams in the northern section
flow north to Hudson Bay via the Nelson System and those in the southern
half are a part of the St. Lawrence System (Danks, 1979).

Within the study area, streams up to fourth order occur. The South Ka-_ |
wishiwi River, an atypical fifth order stream, is largely a chain of lakes or |
lake-like reaches connected by rapids. Most waters in this region are poorly |
buffered, particularly those in bog drainages. The alkalinity of the South
Kawishiwi River, which receives water from most lakes and streams north
of the divide ranged from 9 mg/l to 11 mg/l (as CaCOs) (U.S. Geological
Survey, 1978). South of the divide the alkalinity of the St. Louis River |
downstream from the confluence with the Partridge River ranged from 42
mg/l to 59 mg/l.

The forest is a part of the Great Lakes—St. Lawrence forest region, which
is a transitional zone between the coniferous and deciduous forests (Scud-
der, 1979). The region extends from New Brunswick, Canada, to Lake of |
the Woods on the United States and Canada border about 210 km west of |
the study area. Portions of northern Wisconsin and Michigan are also in-
cluded. The dominant vegetation of northeastern Minnesota is characterized
by quaking aspen (Populus tremuloides) alder (Alnus rugosa and A. crispa),
white pine (Pinus strobus), red pine (P. resinosa), and white cedar (Thuja
occidentalis) (Swain, 1980).

The mayfly faunal list is presented systematically by family and alpha-
betically by genus and species within each family. New state records are
indicated with an asterisk (*); new state records that were previously re-
corded by Daggy (1938; 1941) but unpublished are indicated with two as-
terisks (**). Collection sites are indicated by numbers corresponding to
those listed below and plotted in Fig. 1. Predictions of mayfly species are
listed below the lists of collected species. Voucher specimens are deposited
at the University of Minnesota, St. Paul, Minnesota.

VOLUME 84, NUMBER 4 731

CaN
(of0 >

2 |

} Pea Qo

Fi ie EIS

© Polio a ban Ve

(28) @ ee i
Gi) HOYT LAKES d (21)
hg 5 ry mi 5 KM
saps
2 ? =

Fig. 1. Study area in St. Louis and Lake counties of northeastern Minnesota with site
numbers plotted.

COLLECTION SITES

1. Eagles Nest L., St. Louis Co.: T.62N., R.12W., S.26—28 and 33-35.

2. Shagawa R., St. Louis Co.: T.63N., R.12W., S.26.

. L. White Iron, St. Louis Co.: T.62N., R.12W., S.11 and 14.

i bear isiang k:. St. Louis.Co.: T-G2ZN.. RulZW., S.23.

5-9. South Kawishiwi R., Lake Co., 5: T.63N., R.11W., S.3; 6: T.63N.,
RoW... 8.20: 7: .0.62N., KR: TTIW. -S.312 8: TO2N Ries
97T.62N-. R.10W:, 9.6.

WwW

732 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

10. ‘Filson Cr:, Lake’Co.; Ty-62N., Rel1W.,°8:24.

PLAT wodumnamed str.) Lake Co T.62N.9R TW oS33"

12. Isabella R., Lake Co:: T.61N.,.R29W., S:6.

13. Embarrass’ R.; St. outs Co? T:c0NGeRe lise: S225,

142. Birch’ Ro St. Louis Cox’ T.6INe, RWS.

15. DunkaiR:4St) Louis: Co:: T:60Ns, RA2W2S:9?

16. An unnamed str., St. Louis Co.: T.61N., R.12W., S.36.

id. (Birch L., Lake :Co:: I261N.; Ro TIWasS.30:

18—20. Stony R:, Lake Co. 18: T.61N., R.11W., S.30; 19: T.60N., R. TW?
S.8; 20: T.60N., R.10W., S.28.

21 ANip Crake Comer o0N: Rom Wa 'S.34-

22. Denley Cry VakeiCoi-6IN Re MWe: 28:

23.Nira Crs LakeiCox, 161IN- AR A0W, S38.

24. Snake Cr., Lake Co.: T.61N., R.9W., S.30.

25=26:-StLous R.;'St: Louis Coy25: T-58Nte R21SW:2S)22- 26st
KR I2Wes 2

27. L. Seven Beaver, St. Louis Co P.58N., R.12W., 8:14.

28. Partndge R.7 St-_Lbouis!Conr Pe SssNeaR Wee Ses.

29: Colby, St, Wouis'Co. 9 PS8NIeR 14a Wonses:

LisT OF SPECIES
Family Siphlonuridae
Genus Ameletus Eaton

Ameletus was not taken, however, A. browni McDunnough, A. lineatus

Traver, A. ludens Needham, and A. walleyi Harper are expected.
Genus Parameletus Bengtsson

This genus was not found; expected species are P. croesus McDunnough

and P. midas McDunnough.
Genus Siphlonurus Eaton

Siphlonurus alternatus (Say): 5.

Larvae tentatively identified as §. marshalli Traver were found; expected
species are S. quebecensis (Provancher), S. rapidus (McDunnough), and S.
typicus Eaton.

Family Metretopodidae

Genus Siphloplecton Clemens

Siphloplecton interlineatum (Walsh): 17, 27.
Expected: §$. basale (Walker).

VOLUME 84, NUMBER 4 733

Family Baetidae
Genus Baetis Leach

Baetis brunneicolor McDunnough: 13, 16, 21.

*Baetis flavistriga McDunnough: 2, 5, 13, 18, 20, 25, 28.
*Baetis intercalaris McDunnough: 25, 28.

*Baetis macani bundyae Lehmkuhl: 11.

*Baetis pygmaeus (Hagen): 2, 13, 15.

Baetis tricaudatus Dodds: 20, 24.

Expected: B. frondalis McDunnough and B. hageni Eaton.

Genus Callibaetis Eaton

*Callibaetis ferrugineus (Walsh): 1.

Expected: C. brevicostatus Daggy, C. fluctuans (Walsh), and C. skoki-
anus Needham.

Genus Centroptilum Eaton
Larvae were not identifiable to species. Expected species are C. album

McDunnough, C. bellum McDunnough, C. convexum Ide, and C. rufostri-
gatum McDunnough.

Genus Cloeon Leach

**Cloeon simplex McDunnough: 1.
Expected: C. insignificans McDunnough, C. mendax (Walsh), C. minus
McDunnough, and C. rubropictum McDunnough.

Genus Heterocloeon McDunnough

**Heterocloeon curiosum (McDunnough): 8.

Genus Paracloeodes Day

Although P. minutus (Daggy) inhabits the Mississippi River in southern
Minnesota, the lack of large rivers in the study area may prevent its pres-
ence.

Genus Pseudocloeon Klapalek

Pseudocloeon anoka Daggy: 25.
**Pseudocloeon carolina Banks: 15.
**Pseudocloeon cingulatum McDunnough: 15.
**Pseudocloeon dubium (Walsh): 13, 25.
**Pseudocloeon parvulum McDunnough: 8, 4.
Expected: P. punctiventris McDunnough.

734 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Family Oligoneuriidae
Genus Isonychia Eaton
Larvae were not identifiable to species. Expected species are J. rufa
McDunnough, /. sadleri Traver, and I. sicca (Walsh).
Family Heptageniidae

Genus Arthroplea Bengtsson

*Arthroplea bipunctata McDunnough: 27, 29.

Genus Epeorus Eaton
Larvae were not identifiable to species, however, E. pleuralis (Banks)
and E. vitreus (Walker) are expected.
Genus Heptagenia Walsh
This genus was not collected, however, H. flavescens (Walsh) and H.
pulla (Clemens) are expected.
Genus Leucrocuta Flowers
Leucrocuta hebe (McDunnough): 9, 15.
Expected: L. umbratica (McDunnough).
Genus Nixe Flowers
Nixe was not found; N. lucidipennis (Clemens) and N. rusticalis (Mc-
Dunnough) are expected.
Genus Pseudiron McDunnough
Although Pseudiron was not found, P. centralis McDunnough may occur
here.
Genus Rhithrogena Eaton
No larvae were identifiable to species; R. impersonata (McDunnough),
R. jejuna Eaton, and R. pellucida Daggy are expected.
Genus Stenacron Jensen

**Stenacron candidum (Traver): 15.
Stenacron interpunctatum (Say): 5, 28.
Stenacron minnetonka (Daggy): 13, 28.

Genus Stenonema Traver

Stenonema exiguum Traver: 13, 14, 15, 18, 20.
Stenonema femoratum (Say): 6, 9, 12, 29.

ms

VOLUME 84, NUMBER 4 735

Stenonema integrum (McDunnough): 25.

Stenonema modestum (Banks): 10, 13, 14, 20, 22, 25, 28.
Stenonema pulchellum (Walsh): 9, 13, 28, 20.
Stenonema terminatum (Walsh): 12, 13.

Stenonema vicarium (Walker): 7, 12, 13, 15, 18, 24, 26.
Expected: S$. mediopunctatum (McDunnough).

Family Leptophlebiidae
Genus Choroterpes Eaton

**Choroterpes basalis (Banks): 20.

Genus Habrophlebiodes Ulmer
Although this genus was not collected, H. americanus (Banks) is ex-
pected.
Genus Leptophlebia Westwood
| **T eptophlebia cupida (Say): 1.
Expected: L. nebulosa (Walker).
Genus Paraleptophlebia Lestage

**Paraleptophlebia guttata (McDunnough): 15.
_ **Paraleptophlebia mollis (Eaton): 15, 20, 25.
**Paraleptophlebia praepedita (Eaton): 11, 15, 20.
| **Paraleptophlebia volitans (McDunnough): 20.
Expected: P. adoptiva (McDunnough), P. debilis (Walker), P. moerens
(McDunnough), P. ontario (McDunnough), and P. strigula (McDunnough).

Family Potamanthidae

Genus Potamanthus Pictet

This genus was not found but P. myops (Walsh), P. rufous Argo, and P.
| verticis (Say) are expected.

Family Palingeniidae
Genus Pentagenia Walsh

Although P. vittigera (Walsh) is known from Minnesota, the lack of large
rivers with clay banks in the study area evidently precludes its presence.

Family Ephemeridae
Genus Ephemera Linnaeus

Ephemera simulans Walker: 15, 20.

736 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Genus Hexagenia Walsh
Hexagenia bilineata (Say): 13.
Hexagenia limbata Serville: 1, 3.
Expected: H. atrocaudata McDunnough, H. munda Eaton, and H. rigida
McDunnough.
Genus Litobrancha McCafferty
*Litobrancha recurvata (Morgan): 24.
Family Polymitarcyidae
Genus Ephoron Williamson
*Ephoron leukon Williamson: 19.
Expected: Ephoron album (Say).
Genus Tortopus Needham and Murphy
Tortopus was not collected, although the range of 7. primus (Mc-
Dunnough) may encompass northeastern Minnesota, the absence of large
rivers having clay banks may preclude its presence.
Family Ephemerellidae
Genus Attenella Edmunds
**A trenella attenuata (McDunnough): 15, 20.

Genus Dannella Edmunds

**Dannella simplex (McDunnough): 15.

Genus Drunella Needham

Drunella was not found, however, D. cornuta (Morgan), D. cornutella
(McDunnough), D. lata (Morgan), and D. walkeri (Eaton) are expected.

Genus Ephemerella Walsh

* Ephemerella invaria (Walker): 18.
**Ephemerella needhami McDunnough: 15.
*Ephemerella rotunda Morgan: 12, 13, 18.
Ephemerella subvaria McDunnough: 5, 7.

Genus Eurylophella Tiensuu

**Furylophella bicolor (Clemens): 15, 18, 25.

*Eurylophella minimella (McDunnough): 15.

**Furylophella temporalis (McDunnough): 5, 9, 24.

*Eurylophella versimilis (McDunnough): 29.

Expected: E. bartoni (Allen) and E. lutulenta (Clemens). Eurylophella

VOLUME 84, NUMBER 4 737

funeralis (McDunnough) and E. prudentalis (McDunnough) were found in
the study area by Daggy (1941).

Genus Serratella Edmunds
**Serratella deficiens (Morgan): 18, 20, 25.
*Serratella serrata (Morgan): 15.
**Serratella sordida (McDunnough): 15, 20.

Family Tricorythidae
Genus Tricorythodes Ulmer

No larvae were identifiable to species, however, T. atratus McDunnough
is expected.

| Family Caenidae
Genus Brachycercus Curtis

Brachycercus larvae were not identifiable to species but B. lacustris
(Needham) is expected.

Genus Caenis Stephens

Caenis simulans McDunnough: |.
Expected: C. forcipata McDunnough, C. jocosa McDunnough, C. lati-
_ pennis Banks, and C. tardata McDunnough.

Family Baetiscidae

Baetisca laurentina McDunnough: 9, 13, 15, 18, 19, 25.
Expected: B. lacustris McDunnough and B. obesa (Say).

|
}
Genus Baetisca Walsh
| FAUNISTICS SUMMARY

| Fifty-four nominal Ephemeroptera species were confirmed from the study
area; 33 are new state records, 20 of which were recorded by Daggy (1938,
1941) but are unpublished. As a result of earlier studies, including Daggy’s
surveys, 97 species are presently known from Minnesota. The total number
for the northern region should eventually number at least 140; southern
areas will yield additional numbers.
Of the 54 species, 46 have strong geographic affinities with eastern Can-
ada, and most of these also occur in north central and/or northeastern United
States. A total of 41 of these species range into southeastern United States;
within the southeast 20 appear restricted to Appalachian areas and 21 extend
on to the coastal plain. Only six species, Baetis macani bundyae, Cloeon
simplex, Pseudocloeon anoka, Siphloplecton interlinatum, Stenacron can-

——

738 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

didum, and Stenacron minnetonka, show no affinities with far eastern Can-
ada. Baetis m. bundyae and C. simplex are Canadian species from north
central and western regions, respectively; the other four species are pri-
marily midwestern species.

Only four species from the study area remain unknown from Canada:
Callibaetis ferrugineus, Hexagenia bilineata, Stenacron minnetonka, and
Stenonema integrum. Another group of four species, Baetis tricadatus,
Caenis simulans, Ephemera simulans, Hexagenia limbata, are widely dis-
tributed North American species that range even into southwestern United
States.

New central states distributions are reported for B. m. bundyae, Para-
leptophlebia volitans, and Serratella serrata. Baetis m. bundyae was pre-
viously known only from eastern Northwest Territories, Canada (Lehmkuhl,
1973: Cobb and Flannagan, 1980) and S. serrata was thought restricted to
the Appalachians (Allen and Edmunds, 1963). Ranges are extended west-
ward for Eurylophella minimella, Eurylophella versimilis, and S. serrata
and northwestward for Ephemerella needhami.

Use of the dominant vegetative-growth forms as a predictor of species
presence appears to function for some species (Scudder, 1979; Ross, 1963).
The study area which is located in an ecotome between the coniferous and
deciduous forests was expected to have faunal elements common to these
forest types. Approximately 57% of the mayfly fauna were distributed within
these forested areas. A large portion of the fauna (43%) is distributed beyond
the boundaries of the dominant vegetative-growth form on to the south-
eastern coastal plain and southwestern regions of the United States, and
even into the tundra of northern Canada.

Borders of drainage systems in northeastern Minnesota do not delinate
distributional boundaries for mayflies and did not function effectively as a
biogeographic barrier to dispersal. Only one species, B. m. bundyae, was
restricted to the Hudson Bay drainage system and only five other species
are known to be restricted to regions south of the Laurentian Divide at this
time. These five species, Eurylophella versimilis, Hexagenia bilineata,
Pseudocloeon anoka, Stenacron minnetonka, and Stenonema integrum, may
occur to the north in Canada where few studies have been conducted.

The efficacy of the Laurentian Divide as a barrier to dispersal may have
been circumvented while glaciers receded and waters that now flow north
were continuous with southern river systems. This hypothesis has been
suggested to explain some present day disjunct distributions of mayflies
(Ide, 1955; Lehmkuhl, 1972, 1976) and other aquatic insects (Laufersweiler
and Flannagan, 1981). The relatively short distance across the divide (Fig.
1) and the similarity of habitats on both sides undoubtedly increases the
probability of successful aerial dispersal between drainage systems. Never-
theless, drainage system boundaries may be effective barriers to those species

VOLUME 84, NUMBER 4 739

restricted to large river environments since these species do not inhabit
lower order streams near the boundaries.

Minnesota provides a good setting for the study of factors that may affect
the distribution of aquatic insects since a number of distributional barriers
and indicators converge within the state. The Mississippi River System,
which drains much of central and all of southern regions of the state, borders
the two northern drainage systems previously identified. Two major forest
types (coniferous and deciduous) converge in the northeast. The prairie,
which covers much of the south central and western sections, meet the
deciduous forest along a diagonal line from the northwest to the southeast
corners of the state. In addition some areas of the southeast are unglaciated.
Perhaps future studies will develop a more complete understanding of the
effects of climate, drainage system dynamics and glacial events of the pres-
ent distribution of aquatic insects.

Since B. m. bundyae was found at a latitude much farther south than
expected and in a distinctly different habitat, some biological notes are
presented here on this species. In northern Canada this species inhabits
edges of shallow tundra ponds (Lehmkuhl, 1973). In Minnesota larvae were
found on wood, rock, sand, and detritus substrata in two first order streams.
One stream flows among alder and tamarack [Larix laricina (Du Roi) K.
Koch] and the other occurs in a cleared area; both are less than | m wide
and about 0.1—0.3 m deep.

Specimens in the following stages of development were collected in 1977:
two larvae with partially developed wing pads, 20 June (3.9-4.4 mm); four
larvae with well developed but not darkened wing pads, 20 June (4.3-6.3
mm); two exuviae from mature larvae, 23 June (5.9-6.1 mm); one reared
male, 23 June (6.2 mm); and one reared male, 29 June (6.4 mm). On 18 June
1981 five female larvae with well developed and darkened wing pads (6.5—
7.0 mm) were collected. Mature larvae and adults in Canada are similar in
size (S-7 mm and 5-6 mm, respectively) (Lehmkuhl, 1973; Morihara and
McCafferty, 1979a). The temporal pattern of life cycles differ between north-
ern and southern range extremes. In Canada larvae hatch in early July and
adults emerge in early August (Lehmkuhl, 1973) while in Minnesota adults
emerge in late June and larvae are presumed to hatch in May. Baetis m.
bundyae is likely to have one generation per year and pass the winter in the
egg state in Minnesota as in Canada since the streams it inhabits are reduced
to pools in late summer and must surely completely freeze in winter.

So

——

ACKNOWLEDGMENTS

We thank A. F. Bednarik, L. Berner, and D. K. Morihara for verifications
of Stenonema, Baetisca, and Baetis, respectively. We appreciate the use
of the late W. V. Balduf’s data provided by L. Berner and thank E. F. Cook
for archiving the study specimens. R. Hrdlick and P. Cleveland, The Insti-

rc

740 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tute of Paper Chemistry, assisted with the preparation of the figure and
typing, respectively.

LITERATURE CITED

Allen, R. K. and G. F. Edmunds, Jr. 1963. A revision of the genus Ephemerella (Ephem-
eroptera: Ephemerellidae). VI. The subgenus Serratella in North America. Ann. Ento-
mol. Soc. Am. 56: 583-600.

Banks, N. 1910. Notes on our eastern species of the mayfly genus Heptagenia. Can. Ento-
mol. 42: 197-202.

Bednarik, A. F. and W. P. McCafferty. 1979. Biosystematic revision of the genus Stenonema
(Ephemeroptera: Heptageniidae). Can. Bull. Fish. Aquat. Sci. 201, (vi) + 73 pp.

Berner, L. 1978. A review of the mayfly family Metretopodidae. Trans. Am. Entomol. Soc.

104: 91-137.
Burks, B. D. 1953. The mayflies or Ephemeroptera of Illinois. Bull. Ill. Nat. Hist. Surv. 26,
(iii) + 216 pp.

Cobb, D. G. and J. F. Flannagan. 1980. The distribution of Ephemeroptera in northern
Canada, pp. 155—166. Jn Flannagan, J. F. and K. E. Marshall, eds., Advances in Ephem-
eroptera Biology. Plenum Publ. Co., New York. 556 pp.

Daggy, R. H. 1938. Studies on the biology of some Minnesota mayflies (Ephemeroptera)
with special reference to immature stages. Univ. Minn., Master’s Thesis (unpublished).
81 pp..

—. 1941. Taxonomic and biological investigation of Minnesota mayflies (Ephemerop-
tera). Univ. Minn. Ph.D. Thesis (unpublished). (viii) + 331 pp.

———. 1945. New species and previously undescribed naiads of some Minnesota mayflies
(Ephemeroptera). Ann. Entomol. Soc. Am. 38: 373-396.

Danks, H. V. 1979. Physical basis for Canadian regional diversity, pp. 7-30. In Danks, H.
V., ed., Canada and its insect fauna. Mem. Entomol. Soc. Can. No. 108, 573 pp.
Edmunds, G. F., Jr., S. L. Jensen, and L. Berner. 1976. The mayflies of North and Central

America. Univ. Minnesota Press, Minneapolis. (x) + 330 pp.

Fremling, C. R. 1973. Environmental synchronization of mass Hexagenia bilineata (Ephem-
eroptera) emergences from the Mississippi River. Verh. Internat. Verein. Limnol. 18:
1521-1526.

Hall, R. J., L. Berner, and E. F. Cook. 1975. Observations on the biology of Tricorythodes
atratus McDunnough (Ephemeroptera: Tricorythidae). Proc. Entomol. Soc. Wash. 77:
34-49.

Ide, F. P. 1955. Two species of mayflies representing southern groups occurring at Winnipeg,
Manitoba (Ephemeroptera). Ann. Entomol. Soc. Am. 48: 15-16.

Laufersweiler, P. M. and J. F. Flannagan. 1981. Present recorded distributions and origins
of the Ephemeroptera, Plecoptera, and Trichoptera of Manitoba. Program and Abstracts
of N. Am. Benthol. Soc. 23rd Ann. Mtg. 63 pp.

Lehmkuhl, D. M. 1972. Baetisca (Ephemeroptera: Baetiscidae) from the western interior of
Canada with notes on the life cycle. Can. J. Zool. 50: 1015-1017.

——. 1973. A new species of Baetis (Ephemeroptera) from ponds in the Canadian arctic,
with biologial notes. Can. Entomol. 105: 343-346.

——. 1976. Additions to the taxonomy, zoogeography, and biology of Analetris eximia
(Acanthametropodinae: Siphlonuridae: Ephemeroptera). Can. Entomol. 108: 199-207.

Lewis, P. A. 1974. Taxonomy and ecology of Stenonema mayflies (Heptageniidae: Ephem-
eroptera). U.S. Environ. Prot. Agency, Environ. Monitor. Ser., EPA-670/4-74-006,
(vii) + 81 pp.

VOLUME 84, NUMBER 4 74]

McCafferty, W. P. 1975. The burrowing mayflies (Ephemeroptera: Ephemeroidae) of the
United States. Trans. Am. Entomol. Soc. 101: 447-504.

Morihara, D. K. and W. P. McCafferty. 1979a. Subspecies of the transatlantic species Baetis
macani (Ephemeroptera: Baetidae). Proc. Entomol. Soc. Wash. 81: 34-37.

1979b. The Baetis larvae of North America (Ephemeroptera: Baetidae). Trans. Am.

Entomol. Soc. 105: 139-221.

Needham, J. G., J. R. Traver, and Y.C. Hsu. 1935. The biology of mayflies with a systematic
account of North American species. Comstock Publ. Co., Ithaca, (xvi) + 759 pp.
Pescador, M. L. and L. Berner. /n press. The mayfly family Baetiscidae (Ephemeroptera).

Part II. Biosystematics of the genus Baetisca. Trans. Am. Entomol. Soc.

Ross, H. H. 1963. Stream communities and terrestrial biomes. Arch. Hydrobiol. 59: 235-
242.

Say, T. 1823. Descriptions of insects belonging to the order Neuroptera Linn. Latr., collected
by the expedition authorized by J. C. Calhoun under the command of Major S. E. Long.
Godman’s Western Quarterly Reporter 2(2): 160-165.

Scudder, G. G. E. 1979. Present patterns in the fauna and flora of Canada, pp. 87-179. In
Danks, H. V., ed., Canada and its insect fauna. Mem. Entomol. Soc. Can. No. 108,
573 pp.

Swain, A. M. 1980. Landscape patterns and forest history in the Boundary Waters Canoe
Area, Minnesota: A pollen study from Hug Lake. Ecology 61: 747-754.

U.S. Geological Survey. 1978. Water resources data for Minnesota Vol. 1, Great Lakes and
Souris-Red-Rainy River basins. Report No. USGS/WRD/HD-79-028, (viii) + 300 pp.

Waters, T. F. 1966. Production rate, population density, and drift of a stream invertebrate.
Ecology 47: 595-604.

Waters, T. F. and G. W. Crawford. 1973. Annual production of a stream mayfly population:
A comparison of methods. Limnol. Oceanogr. 19: 286-296.

Wright, H. E., Jr. 1972. Physiography of Minnesota, pp. 561-578. Jn Sims, P. K. and G. B.
Morey, eds., Geology of Minnesota: A centennial volume. Univ. Minn., Minn. Geol.
Surv. St. Paul, 623 pp.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 742-745

NOTES ON THE ETHOLOGY OF NEOSCLEROPOGON ELONGATUS
(DIPTERA: ASILIDAE) IN SOUTH AUSTRALIA'”

ROBERT J. LAVIGNE

Entomology Section, University of Wyoming, Box 3354, University Sta-
tion, Laramie, Wyoming 82071.

Abstract.—Notes are presented on the predatory and mating behavior of
Neoscleropogon elongatus (Macquart) in South Australia. Prey taken by
N. elongatus represented three orders, Diptera, Hymenoptera, and Heter-
optera-Hemiptera, with Diptera and Hymenoptera accounting for 88% of
the prey. Mating, without prior courtship, occurred in the tail-to-tail posi-
tion, typical of Dasypogoninae.

The taxonomy of Neoscleropogon elongatus (Macquart) is somewhat
confused. It was first described by Macquart in 1847 in the genus Dasypo-
gon. White (1917) regarded it as belonging to the genus Stenopogon and
recorded it from Tasmania in January and February. Hardy (1926) illustrated
the genitalia of S. elongatus from New South Wales specimens, but later
(1930) indicated that this drawing referred to S. fraternus Bigot. However,
in 1934, he established S$. elongatus as a synonym of S. fraternum (mis-
spelled). As a result of his visit to Australia, Hull (1962) again separated the
two species and placed both in the genus Neoscleropogon. Therefore, if
White and Hardy were correct in their identifications, N. elongatus has
been recorded, thus far, only from Tasmania and N. fraternus from New
South Wales.

Data included herein on the behavior of N. elongatus is admittedly in-
complete. However, since the behavior of Australian asilids is virtually
undocumented, these notes may be of some interest.

Behavioral observations were made on this species, at Sandy Creek, near
Gawler, South Australia, intermittently between November 28 and Decem-

' Published with the approval of the Director, Wyoming Agricultural Experiment Station, as
Journal Article No. JA 1128.

°* This research was supported in part by National Science Foundation Travel Grant INT 78-
17131, in part by the Waite Agricultural Research Institute, Adelaide, Australia, and in part
by the University of Wyoming.

VOLUME 84, NUMBER 4 743

ber 12, 1978, in a harvested oat field and the weedy area adjoining it. Sal-
vation jane (Echium plantagineum L.) was common in the field of oats,
while the weedy area was dominated by a wallaby grass (Danthonia genic-
ulata J. M. Black), a spear grass (Stipa semibarbata R. Br.), Lipidosperma
viscidum R. Br., Neurachne alopecuroidea R. Br., and Baeckea behrii
(Schlidl.) F. Muell. The weather was typical for a South Australian spring,
with several days of rain during the period and temperatures ranging from
16° to 28°C.

Other locations in South Australia where N. elongatus was collected are
Aldinga Beach, 22.1.79, and Ferries-McDonald Conservation Park, south of
Monarto, 12.xii.78. The habitat in both locations was composed of native
vegetation, e.g., Mallee scrub, and beach scrub community, respectively.

Methods used to observe N. elongatus were the same as those used in
describing the behavior of Neoitamus vittipes (Macquart) (Lavigne, 1982).

FORAGING AND FEEDING BEHAVIOR

Asilids were active whenever it was not raining and temperatures ex-
ceeded 21°C. Orientation flights were long (3 to 7 m) undulating excursions
with the asilid often zigzagging through the vegetation 15 to 46 cm above
the ground. Neoscleropogon elongatus foraged from oat stubble and dried
plant stalks at heights ranging from 12 cm to 0.75 m. Successful flights after
prey were initiated from distances of 30 to 36 cm, although rapid flights of

2 m after prey were commonly observed. Potential prey were sometimes

chased over long distances. In some instances the robber fly would veer

| away when close. Insufficient data are available for establishing any pattern.
Foraging by N. elongatus occurred as early in the day as 0945 h and as late
as 1920 h.

Prey were collected in the air, and, on the single occasion when the author
was close enough to observe, the prey was impaled during manipulation
following the landing. The same type of manipulation, that of holding the
vegetation with one foretarsus and using the other five to revolve the prey,
was seen during the feeding sequence on a honey bee. This behavior is
similar to that exhibited by Diogmites angustipennis Loew in western North
America (Lavigne and Holland, 1969). On the single occasion when prey
release, following feeding, was observed, the asilid pushed a tachinid off its
proboscis with both foretarsi.

Based on 25 measured prey of N. elongatus, the ‘‘preferred”’ prey length
was 8.3 mm: the prey size varied from 2.8 to 15.0 mm. Measurements of

four female and five male asilids suggest that females are slightly smaller
than males (17.9 vs. 20.0 mm; range, male: 18.5-21.0 mm, female: 16.3-
20.0 mm), an unusual situation. However, males chose smaller prey (18.5—
21.0 mm, < 7.6 mm) than did females (16.3-20.0 mm, x 9.3 mm). Since it is
| unusual for female asilids to be smaller than males, it is probable that had

|

744 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON |

a longer series of asilids been collected, the reverse situation would have
been found, thus explaining the smaller prey size for males. The number of |
prey collected from males (9) was slightly more than one-half that collected
from females (14). Males showed a slight “‘preference’’ for Hymenoptera,
whereas females concentrated on both Hymenoptera and Diptera. The species
appears to be stenophagic, as defined by Lavigne and Holland (1969), taking
as prey representatives of only three insect orders, Diptera (40%), Hyme-
noptera (48%), and Hemiptera-Heteroptera (12%). Even though there were
ca. 15 beehives in the adjoining paddock, honey bees were not being taken
excessively in terms of their abundance; only 11% of the prey were honey
bees. Despite the fact that bush flies were very conspicuous, these insects
were largely ignored by the asilids, presumably because of their small size.
The mean predator to prey ratio was 2.3.

Herein is a list of prey taken by N. elongatus. Specific identifications
were made, where possible, but because of the state of knowledge in some
groups, only genus and/or family names are included for some specimens.
The number of records and sex of the predator are indicated in parenthesis
following the prey record. All prey records are from Sandy Creek, unless
otherwise noted.

DIPTERA, Asilidae: Bathypogon sp., 12.xii.78 (2) (Ferries-McDonald |
Consv. Pk.); Muscidae: Musca vetustissima Walker, 7.xii.78 (6), 8.xii.78 |
(?); Syrphidae: Eristalis tenax L., 1.xii.78 (2); Tachinidae: Chaetophthal- |
mus sp., 7.xii.78 @), 8.xii.78 G, 3 2). HEMIPTERA-HETEROPTERA, |
Alydidae: Riptortus sp., 28.xi.78 (6); Lygaeidae: Nysius vinitor Bergroth, |
1.xii.78 (2); Miridae: Creonotiades dilutus Stal, 28.x1.78 (6). HYMENOP-
TERA, Apidae: Apis mellifera L., 28.xi.78 (2 2), 8.xii.78 ); Halictidae:
Lasioglossum sp., 1.xii.78 (6), 7.xii.78 ( , 2); Tiphiidae: Thynninae, 28.x1.78
(6, 2), 7.xii.78 @), 8.xii.78 G, 2). An additional unidentified Diptera and
an unidentified Hymenoptera also served as prey at Ferries-McDonald Con-
servation Park.

MATING

Only one complete mating was observed (28.xi.78). In this instance there
was no courtship. A female flying a zigzag pattern through the vegetation |
at 1240 h was accosted by a male rising out of the understory. Coupling —
took place at a height of ca. 45 cm and the pair landed on an oat stalk at
approximately the same height (temperature 28°C). Upon landing the pair
took a tail-to-tail position. The female was feeding on a tiphiid wasp when
encountered by the male. She subsequently dropped the prey during a flight
to a new resting place. Separation occurred at 1253 h after the female pushed
at the genitalic junction with her hindtarsi.

Three other mated pairs were encountered 1101 h (24°C), 7.xii.78; 1223 h
(26°C), 7.xii.78 and 1200 h (26.5°C), 8.xii.78. Remaining times in copula were

VOLUME 84, NUMBER 4 745

6 min., 17 min., and 10 min., respectively. Near the end of each mating,
the female would alternately raise and lower her hindlegs several times.
This action was accompanied by the female pushing against the genitalic
connection with her hindtarsi, ca. two minutes prior to separation. At sep-
aration the male released his claspers and flew off, leaving the female on
site.

ACKNOWLEDGMENTS

I acknowledge, with great appreciation, the taxonomists at CSIRO, Di-
vision of Entomology, Canberra, who identified the prey insects: D. H.
Colless (Diptera) and I. D. Naumann (Hymenoptera). G. F. Gross of the
Entomology Department, South Australian Museum, Adelaide, is thanked
for his identification of Hemiptera-Heteroptera. I express my appreciation
to Gregory Daniels, Department of Entomology, University of Queensland,
St. Lucia, for his identification of N. elongatus and Bathypogon sp. E. D.
Symon, Agronomy Department, Waite Agricultural Research Institute, Uni-
versity of Adelaide, was most helpful in identifying botanical specimens that
characterized the environments in which N. elongatus was collected. I ad-

| ditionally thank Peter Miles and staff at Waite Institute for providing me
with laboratory space, supplies, and transportation. I especially thank my
wife, Judy, for her time and effort as an unpaid research associate.

LITERATURE CITED

|
/Hardy,G. H. 1926. A new classification of Australian robber flies belonging to the subfamily
| Dasypogoninae (Diptera: Asilidae). Proc. Linn. Soc. N.S.W. 51(Pt. 3): 305-312.
1930. Fifth contribution towards a new classification of Australian Asilidae (Diptera).
Proc. Linn. Soc. N.S.W. 55(Pt. 3): 249-260.
1934. The Asilidae of Australia. Part II]. Ann. Mag. Nat. Hist. (10)14: 1-35.
Hull, F. M. 1962. Robber flies of the world. The genera of the family Asilidae. Pt. 1. U.S.
Natl. Mus. Bull. 224: 1-430.
Lavigne, R. J. 1982. Ethology of Neoitamus vittipes (Diptera: Asilidae) in South Australia.
Proc. Entomol. Soc. Wash. 84: 617-627.
Lavigne, R. J. and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming
robber flies (Diptera: Asilidae). Univ. Wyo. Agric. Exp. Stn. Sci. Monogr. No. 18,
61 pp.
White, A. 1917. The Diptera—Brachycera of Tasmania. Part 3. Families Asilidae, Bomby-
liidae, Empididae, Dolichopodidae and Phoridae. Pap. R. Soc. Tasmania 1916: 148-266.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 746-752

OCONOPERLA, A NEW GENUS OF NORTH AMERICAN PERLODINAE
(PLECOPTERA: PERLODIDAE)'

BILL P. STARK AND KENNETH W. STEWART

(BPS) Department of Biology, Mississippi College, Clinton, Mississippi
39058; (KWS) Department of Biological Sciences, North Texas State Uni-
versity, Denton, Texas 76203.

Abstract.—Oconoperla weaveri, a new genus and species of Perlodinae, is
described from one adult male and five pre-emergent nymphs collected in
mountainous North and South Carolina. Oconoperla appears to be most
closely related to Malirekus Ricker. Descriptions are supported by SEM
photomicrographs and original drawings.

The most recent study of Nearctic Perlodinae (Ricker, 1952) recognized
13 subgenera of Isogenus. All of these have subsequently been given generic
rank by Illies (1966) and Zwick (1973). This study reports the discovery of
an adult male and five nymphs of a previously undescribed genus in this
group. Nymphs were associated by dissecting adult genitalia from speci-
mens which died during emergence. Methodology follows Stark and Stewart
(1981).

Oconoperla Stark and Stewart NEW GENUS

Type-species.—Oconoperla weaveri, n. sp.

Description.—Size: Medium sized, 12-14 mm in length.

Mesosternum: Y-ridge arms attached to posterior corners of furcal pits;
transverse ridge present. Stem of Y-ridge short (Fig. 4).

Male 10th tergum: Cleft extends to near anterior margin; hemitergal lobes
rounded and covered apically with dense patch of long thin setae (Fig. 1).

Epiproct: Bulbous apically and covered dorsolaterally with small scale-
like setae. Dorsal sclerite slender, almost needle-like; not reaching tip (Figs.
1, 2). Ventral sclerite well developed, sharply upturned over apex; 2 ven-
trally directed teeth at midlength (Fig. 1).

Lateral stylets: Not exceeding apex of epiproct in length; most heavily

' Study supported by NSF Grant #DEB 78-12565.

VOLUME 84, NUMBER 4 747

LVL ie .
LUA Vs
GL SA Wy any

~

.
bean Ep ICL

ew

.

Nee

.
s
s
..

Figs. 1-4. Oconoperla weaveri adult structures. 1, Male terminalia, lateral (E = epiproct;

LS = lateral stylet). 2, Epiproct, dorsal. 3, Female subgenital plate with vagina (dorsal aspect)
inset. 4, Mesosternal grooves.

sclerotized along dorsal margin; ventral margin abruptly narrowed near apex
to form truncate tip (Fig. 1).

Ventral lobes: Absent.
Female subgenital plate: Slightly produced and densely hairy; posterior
_ margin truncate and emarginate mesally (Fig. 3).
_ Nymphal lacinia: Bicuspid; shorter subapical tooth strongly divergent
from apical tooth. Apical tooth with groove along basal half of inner surface;
slender, delicate sensillum located at base of tooth. Three or 4 prominent

setae along inner margin of lacinia followed by row of smaller setae; base
with patch of small setae near margin (Fig. 9).

748 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Oconoperla weaveri, nymphal habitus.

VOLUME 84, NUMBER 4 749

Figs. 6-9. Oconoperla weaveri, SEM photomicrographs of nymphal structures. 6, Abdom-
inal tergum 8, 200. 7, Detail of tergal setae, 1000. 8, Right mandible, 150. 9, Lacinia,
) 170x.

Nymphal mandibles: Four major teeth; innermost tooth tridentate. Ser-
rations absent. Marginal setal fringe covering ca. 2 of inner margin; ventral
median setae forming an irregular linear patch; 4 prominent setae near base
of apical tooth. Linear patch of stout spines near base of inner tooth on left
mandible.

Submental gills: Not extending beyond lateral margins of submentum.

Nymphal pronotum: Posterolateral margins notched (Fig. 5).

Distribution Southern Appalachian Mountains.

Etymology.—The prefix, Ocono, is derived from Oconee, the South Car-
olina county in which the holotype was collected.

Diagnosis. —Oconoperla nymphs are distinguished from those of sympat-
ric perlodinae genera (Cultus, Diploperla, Helopicus, Hydroperla, Isoge-
noides, Malirekus, Remensus, and Yugus) by the notched lateral pronotal
margin and the densely hairy habitus (Fig. 5). Yugus and Malirekus nymphs
bear the closest general resemblance to those of Oconoperla, but in both
these genera the lacinia bears a short second tooth which arises unusually
close to the apex of the major tooth, and a tuft of 4-5 large setae on a small

750 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

knob near the base of the second tooth. The lacinia of Oconoperla bears a
longer, more divergent and proximally located second tooth and lacks a
tufted knob on the inner margin (Fig. 9).

Five of the sympatric genera listed above have lateral stylets but only
two of these (Isogenoides and Hydroperla) also share the presence of spines
on the ventral epiproct sclerite with Oconoperla. Males of these two genera
are additionally distinguished from those of Oconoperla by the presence of
long submental gills.

The affinities of this unique genus will be clearer once eggs and adult
females are available for study. Male genitalic structure is suggestive of a
close relationship with the Malirekus-Yugus group of genera, but the ap-
parent absence of synapomorphies in nymphal mouthparts (e.g., tufted la-
cinial knob, grooved inner margin of major lacinial tooth, isolated setal row
near base of apical mandibular tooth) between Oconoperla and Malirekus-
Yugus group suggests the relationship is a remote one.

Oconoperla weaveri Stark and Stewart, NEW SPECIES
Figs. 1-13

Male.—Forewing length 14 mm; body length 11 mm. General color dark |
brown. Dorsum of head brown except for small pale spet posterior to ocelli.
Pronotum brown with narrow yellow median stripe. Genitalia described
above.

Female.—External genitalia from pre-emergent nymph described above.
Vagina membranous; lateral margins sinuate. Spermatheca slender, mem-
branous, bearing ca. 7 accessory glands in apical % (Fig. 3).

Nymph (Fig. 5).—General color brown patterned with yellow; membra-
nous areas with purple-red pigment in living specimens. Head brown except |
for pale M-line and 2 pair small circular spots; smaller pair forward of |
M-line, larger pair lateral to ocelli and forward of ecdysial suture. Thoracic
terga brown with scattered circular pale areas. Abdominal terga brown with
double row pale spots on Ab 3-7 and mesal single row pale spots on Ab 6—
9; additional expanded pale areas on Ab 6-10. Body densely hairy; occipital
spinule row absent. Legs without fine silky setal fringe; femora without large
socketed spinelike setae (Figs. 10, 11). Abdominal terga with dense mesal
patch of short, thick curled setae (Figs. 6, 7).

Types.—Holotype ¢ (USNM #100106) from Oconee Co., S.C., 0.5 mi E
Hwy 107, Tamassee Rd., small spring at headwaters of Wash Branch, 22-
V-1981, B. Stark and J. S. Weaver, deposited in the National Museum of
Natural History, Washington, D.C. Paratypes: NORTH CAROLINA: Ma- |
con Co., Dirty John Crk, Wayah Bald, 18-IV-81, J. S. Weaver and R. Hol-
zenthal, 2 nymphs (NTSU, BPS); Rattlesnake Spring, Wayah Bald, 18-IV-
1981, J. S. Weaver and R. Holzenthal, 1 nymph (BPS); Rattlesnake Spring,

VOLUME 84, NUMBER 4 751

PEO 8 THE

4

AM
SL UE ate Wi (KAN Life

re md

Figs. 10-13. Oconoperla weaveri, SEM photomicrographs of nymphal structures. 10, Fore-
leg, anterior aspect, 58x. 11, Femur, anterior aspect, 125x. 12, Antenna, basal segments,
100. 13, Detail of antennal setation, 1000.

Wayah Bald, 26-V-1981, B. Stark and J. S. Weaver, 2 nymphs (USNM,
BPS).

Etymology.—This species is named in honor of John S. Weaver, III.
_ Ecological note—Nymphs of O. weaveri were found under rocks in splash
Zones of small spring seeps. Additional rarely collected species taken at
Wayah Bald with Oconoperla include Viehoperla ada (Needham and Smith),
Isoperla distincta Nelson, and Beloneuria georgiana (Banks).

ACKNOWLEDGMENTS

| We are grateful to John S. Weaver, III, for his assistance in fieldwork

, and for his gift of additional specimens used in this study. Sarah Faison of
the University of Mississippi Dental School assisted in preparation of SEM
photomicrographs and Ellen Hart Navoor prepared the full nymphal illus-
tration under our direction.

752 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

LITERATURE CITED

Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich, 82. Walter de Gruyter and
Co., Berlin. 632 pp.

Ricker, W. E. 1952. Systematic Studies in Plecoptera. Indiana Univ. Publ. Sci. Ser. 18: 1I-
200.

Stark, B. P. and K. W. Stewart. 1981. The Nearctic genera of Peltoperlidae (Plecoptera). J.
Kans. Entomol. Soc. 54: 285-311.

Zwick, P. 1973. Insecta: Plecoptera. Phylogenetisches system und Katalog. Das Tierreich,
94. Walter de Gruyter and Co., Berlin. 465 pp.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 753-760

LARVAL DEVELOPMENT IN BLISTER BEETLES OF THE
GENUS LINSLEYA (COLEOPTERA: MELOIDAE)

RICHARD B. SELANDER

Departments of Genetics and Development and of Entomology, Univer-
sity of Illinois, Urbana, Illinois 61801.

Abstract.—Twenty T, larvae of Linsleya convexa (LeConte) failed to feed
on pollen taken from the comb of the honeybee. Two of nine T, larvae given
eggs of the acridid grasshopper Melanoplus differentialis (Thomas) com-
pleted the developmental pattern T,-FG,_;-C, in 23 and 27 days. After
being chilled at 5°C for two months to break diapause, one of the larvae
reached instars SG,, Pz, and A, in 19, 24, and 34 days, respectively.
The other larva molted to instar C, after chilling but failed to complete

- ecdysis.

A previous contention that L. sphaericollis (Say) regularly follows the
_abbreviated ontogenetic pattern T,-FG,_;-P,-A,;, with diapause in instar FG,
_is regarded as unfounded.

| Some anatomical features of the larva and pupa of L. convexa are de-
| scribed. It is concluded that, while Lins/eya does not belong in the Lyttina,
| the evidence for placing it in the Epicautina is, as a whole, equivocal.

Following MacSwain’s (1951) proposal, based on external anatomy of the
first-instar (triungulin) larva, that the genus Linsleya MacSwain be assigned
to the subtribe Epicautina, rather than the Lyttina, several workers have

addressed the question of whether species of this genus are, like most species
| of the genus Epicauta Dejean, predators of the eggs of acridid grasshoppers
(Selander, 1964; Selander and Pinto, 1967; Church and Gerber, 1977). The
question is certainly an important one systematically, since an affirmative
| answer would provide strong support for MacSwain’s hypothesis.
Selander (1964) called attention to a report of Criddle (1931), in which
Linslelya sphaericollis (Say) was cited as one of several ‘‘enemies”’ of grass-
| hoppers in Canada, and of Romanov (1954), in which the larva of this species
was stated to be predaceous on grasshopper eggs in Manitoba. Unfortu-
nately, neither report gives any indication whatsoever regarding the evi-
dence for the reputed predation of grasshopper eggs by larvae of L. sphaer-
icollis. On the other hand, Church and Gerber (1977) reported that “‘none

a

754 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

of several hundred meloids’’ taken from egg pods of Melanoplus sanguinea
(Fabricius), M. bivittatus (Say), and Camnula pellucida Scudder in the
Canadian prairies “‘in recent years’’ proved to represent Linsleya sphaeri-
collis, even in instances where adults of this species were abundant in the
area in which grasshopper eggs were collected. Moreover, Selander and
Pinto (1967) reported failure of 24 larvae of Linsleya convexa (LeConte) to
feed on eggs of Melanoplus differentialis (Thomas) in an attempted labo-
ratory rearing.

At this point one might suspect that the reports of Criddle and Romanov
were perhaps suppositional and that grasshopper eggs are not, in fact, the |
prey-type of larvae of Linslelya. Yet Church and Gerber (1977) and Peterson
(cited by them) were able to induce a small percentage of larvae of Linsleya
sphaericollis to feed on the eggs of Melanoplus in the laboratory and in one
instance actually obtained a complete rearing to the adult stage.

In this article I report an experimental rearing of Linsleya convexa in |
which first-instar larvae were given, as prospective food, either pollen pro-
visioned by the honeybee (Apis mellifera Linnaeus) or eggs of the grass- |
hopper Melanoplus differentialis. In addition, | describe some of the ana-
tomical features of the larva and pupa of L. convexa.

AN EXPERIMENTAL REARING LINSLEYA CONVEXA

Materials and methods.—I follow Selander and Mathieu (1964) in referring |
to the four phases of the larval stage of Meloidae as triungulin, first grub,
coarctate, and second grub, commonly denoted by the symbols T, FG, C, }
and SG, respectively, with numerical subscripts to indicate instar, as nec- |
essary.

The experiment utilized 29 T larvae of Linsleya convexa eclosing from |
an egg mass laid by a female that was part of a group of adults collected at
Fort Davis, Jeff Davis County, Texas, 3/10 August 1968, on Chilopsis lin- |
earis and Tetraclea coulteri. The eggs, laid 13 August at the collection site,
were placed in a 3-dram vial and held at ambient temperature, 100% RH, |
in darkness until 15 August, when temperature was stabilized at 27°C. The
egg mass contained 30 eggs, all of which hatched on 5 September (24 days). |
Larvae remained under the conditions of incubation before use in the ex- |
periment. )

Twenty larvae were given pollen at the age of seven days (12 September); |
nine were given grasshopper eggs at the age of 10 days (15 September). One |;
larva from the lot died at the age of 8 or 9 days, before entering the exper- |
iment. f
Pollen was removed from a comb of the honeybee shortly before use, |
mixed with distilled water, and worked into small balls, as described by |
Selander and Mathieu (1964). The amount of water was varied to produce |

VOLUME 84, NUMBER 4 755

five balls each of four consistencies (from ‘‘rather dry’’ to ‘‘soupy’’). Each
ball was placed on the inner surface of a cotton-plugged glass tube (4 mm
inner diameter), and a Linsleya larva was deposited on one of the cotton
plugs near the pollen ball.

Melanoplus differentialis eggs were nine months old. They had been in-
cubated initially at 27°C, 100% RH for 1-2 months in order to allow them to
develop to the diapause state and then held at 5°C until use. Larvae of
Linsleya were given about 40 Melanoplus eggs each in individual cotton-
plugged glass tubes 8 mm inner diameter. Allotments were random selec-
tions of eggs from 50 egg pods.

Larvae receiving pollen were placed at 27°C, 100% RH, in darkness.
Those receiving Melanoplus eggs were placed initially under the same con-
ditions, but on day 8, surviving, unfed larvae were transferred to 35°C.
Larvae that completed feeding in instar FG; were transferred to a sand/soil
mixture (moistened with 10% water by volume). Subsequent treatment of
larvae is described in the section on results.

Results.—All 20 larvae of Linsleya convexa given pollen died in the T
phase, evidently without feeding. Seventeen became mired in the pollen and
apparently drowned on day 1: two did so on day 2. On day 3 the lone
survivor was transferred to a larger tube (8 mm inner diameter) with a fresh
pollen ball of medium consistency in which a small amount of honey had
‘been incorporated. This larva survived 31 days in rearing. The length of the
| survival period might be interpreted as evidence that the larva was obtaining
some nourishment from the pollen ball. However, some of the T larvae of
\L. convexa studied by Selander and Pinto (1967) lived as long as 23 days at
27°C without feeding, and some of those of L. sphaericollis studied by
Church and Gerber (1977) survived without food for 4-7 weeks on moist soil
at 20°C.

Seven of the nine larvae of L. convexa given Melanoplus eggs died in the
T phase, without feeding, in a mean of 13.1 (1.78) days (range 8-22). One
larvae (#1, a female), after transfer to 35°C, began feeding on an egg on day
110; reached instars FG,, FG,, FG,, and FG, on days 11, 13, 15, and 17,
respectively; was transferred to sand/soil on day 22; and ecdysed to C, on
day 27. A day later it was returned to 27°C. Another larva (#2, a male),
which remained at 27°C, began feeding on day 2; reached instars FG,, FG,
FG,, and FG, on days 5, 7, 9, and 12, respectively; was transferred to sand/
soil on day 17; and ecdysed to instar C, on day 23.

In the FG phase both larvae produced a feces of paste-like consistency
which was spread over the grasshopper eggs and on the sides of the glass
ubes as droplets. In FG, both larvae excavated a cell in the sand/soil in
which they became motionless two days before ecdysis occurred.

On day 176 of the experiment the two C larvae, which were evidently in

756 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

diapause, were transferred to 8 mm inner diameter tubes and placed at 15°C,
100% RH, in darkness. After 60 days they were returned to 27°C, again at
100% RH in darkness.

Larva #1 broke diapause and entered instar SG; 19 days after chilling
ended (day 255), pupated five days later (day 260), and reached the adult
stage 10 days after that (day 271). The adult was perfectly formed and of
exceptionally large size (length, to end of elytra, 15 mm; cf. Selander, 1955).
Larva #2 was killed 871 days after chilling ended (day 1107), at which
time I found that it had molted, at some time during the post-chilling period,
to instar C; inside the C, skin. The occurrence of two consecutive instars
in the coarctate larval phase was recorded previously in Pyrota palpalis
Champion by Selander and Mathieu (1964).

Discussion.—The results of the experiment are consistent with the rearing
attempts of Church and Gerber (1977) in two major respects. First, it is now
established that the larva of Linsleya convexa, like that of L. sphaericollis,
is capable of developing to the adult stage on a diet of the eggs of Melan-
oplus grasshoppers. Second, as in the case of Linsleya sphaericollis, only
a small proportion of L. convexa larvae provided with Melanoplus eggs in
the laboratory respond positively to them.

In Church and Gerber’s (1977) rearing of Linsleya sphaericollis on the
eggs of Melanoplus sanguinipes and M. bivittatus, 16(4.3%) of 370 T larvae
fed, 12 (3.2%) reached FG,, and 8 (2.2%) reached FG;. Five of the FG;
larvae “‘lived 4 to 10 weeks at 20°C after they stopped feeding, but remained
unchanged’’; the fate of the remaining three was not mentioned. Peterson, _
in an unpublished study cited by Church and Gerber, obtained 31 FG larvae
from 240 T larvae given food (12.9%); the percentage of success might have
been higher had he reared each of the larvae individually, rather than placing
1-5 in a vial. Two of the FG larvae lived until at least day 76 of his rearing.
In addition, a dead adult was found on day 76 among material kept at 29°C,
having “‘evidently pupated directly from the fifth instar [FG,]’’ (Church and
Genber1977).

A possible explanation for the poor feeding response obtained in the lab- |
oratory for both species of Linsleya is suggested by the exceptionally long |
survival period of T larvae under starvation, mentioned above. Comparable }
longevity is charactersitic of T larvae of Epicauta pennsylvanica (DeGeer)
from Illinois and Mississippi (Selander, unpublished data). Moreover, larvae
of this species, like those of Linsleya, are very unlikely to feed in the first
few weeks following eclosion. In this case it can be shown that the newly
eclosed T larvae are in a behavioral diapause state. A procedure that I have }
found effective in rearings of E. pennsylvanica is to chill newly eclosed T }
larvae for two months at 5°C. It might be interesting to see the effect of this |
or a similar treatment on the feeding response of T larvae of Linsleya.

VOLUME 84, NUMBER 4 757

Church and Gerber (1977) concluded that their and Peterson’s FG; larvae
of Linsleya sphaericollis that survived for several weeks were in diapause
when they died and that “‘hypnothecal’’ (C) and ‘‘non-vorant’’ (SG) phases
of the larval stage “‘probably seldom or never occur in this species.”’ Fur-
ther, they speculated that in nature larvae overwinter in instar FG,, pupate
directly from that instar in spring, and emerge as adults a few weeks later.
Not only that, since the FG; larvae are ‘‘much less resistant to desiccation
than [coarctate larvae] and likely could not survive more than one winter,
a l-year life cycle is indicated’’!

_ This is highly imaginative ecology and, for all I know, may be true. But
on the basis of the actual evidence, it is hardly to be taken seriously. The
failure of the larvae of L. sphaericollis to develop beyond the FG phase
may have been simply the result of lack of access to soil of proper moisture
content (whether any soil was available to them in Church and Gerber’s
rearing is not stated). Although some Lyttini can complete the FG phase
without access to moist soil, in my experience it is an absolute necessity
for species of Epicauta. Nor can the possibility of disease or nutritional
inadequacy of the food be ruled out. But in any event, arrested development
is, by itself, hardly proof of diapause, especially when it terminates in death.
| If, as Church and Gerber report, Peterson’s adult did not pass through
‘the C phase of larval development, this in itself may be taken as good
evidence for regarding Linsleya as an epicautine genus, since the abbrevi-
ated pattern T-FG-P-A has been recorded previously only in members of
the genus Epicauta (Selander and Weddle, 1969). However, no species of
‘the subfamily Meloinae is known to diapause in the FG larval phase and no
}species of Meloidae is known to follow only the abbreviated ontogenetic
pattern, and it was at best ingenuous, considering the fragmentary nature
of their information, for Church and Gerber to suggest that Linsleya sphaer-
icollis does so.

| ANATOMY

The triungulin (T,) larva of Linsleya convexa was described and illustrated
iby MacSwain (1956). Some anatomical features of the FG., FG;, C,;, and
SG, larva and of the pupa are noted below. The descriptions are limited to
characters which, in my experience, vary significantly among genera and
other higher taxa of Meloidae. Except in the case of the C, larva, they are
based on exuvia.

FG, larva.—Cuticle sparsely but conspicuously clothed throughout with
relatively long, slender setae. Spiracles lateral. Labrum rectangular, with
anterior margin straight and with 8 setae in median transverse row. Mandible
)Fig. la) with a series of 9-10 vestigial, blunt teeth on mesodorsal margin,
jterminating basad in a large, prominent, triangular tooth; mandibular setae

:
|

758 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Linsleya convexa. a, Right mandible of FG, larva, dorsal view. b, Right mandible
of FG, larva, dorsal view. c, Right mandible of SG; larva, dorsal view. d, Left maxilla of FG.
larva, ventral view. e, Right leg II of FG, larva, anterior view. f, Tibia and tarsungulus III of |
FG, larva, anterior view.

equal in length. Antennal segment III about 3/10 as long as II, with terminal
seta 2!/s x as long as III; II bent, with sensory appendix rounded, button-like,
lateral. Maxilla (Fig. 1d) with mala prominent, glabrous ventrally, with sev-
eral setae dorsally; palpus retaining orbicular form much as in T,. Labial
palpus with segment II tapered, as long as I. End of abdomen with a row
of 4 long setae of equal length. Leg (Fig. le) elongate, rather heavy, witha |
definite pattern of setation; coxa prominent; tibia about 1'/;x as long as
femur; tarsungulus 4 as long as tibia, bearing a single seta.

FG, larva.—Cuticle relatively densely, very conspicuously clothed with
setae, most of which are heavy and rather short. Mandible (Fig. 1b) much
more massive than in FG,, lacking teeth. Antennal segment III less than 4
as long as II, less than % as wide, with terminal seta about as long as II];
II */s as wide as long, with sensory appendix terminal, slightly wider than
III. Maxilla with mala massive, sparsely setate ventrally, densely so dorsally; |

VOLUME 84, NUMBER 4 759

| Fig. 2. C, larva of Linsleya convexa.

_palpus not so orbicular as in FG,. Labial palpus less elongate than in FGs.
| Leg (Fig. If) heavy, relatively short, densely setate; tarsungulus well de-
' veloped, bearing 2 setae.

C, larva (Fig. 2).—Subnavicular, slightly curved, lacking lateral ridge on
‘abdomen. Cuticle uniform light brown in color, uniformly pebbled, com-
/ pletely lacking striae. Mandible stubs acute. Spiracular cones small, not at
all bulbous or sagged posteriad; spiracular openings very small. Dorsal line
| of dehiscence well developed and complete on thoracic segments and ab-
)dominal segments I-VI.

SG,.—Similar to FG;. Mandible (Fig. 1c) much wider, with a chisel-like
flange on mesal margin. Antenna, maxillary palpus, and labial palpus some-
what shorter; antennal segment III flattened, at least in exuvia hardly dis-
tinguishable as a separate segment. Leg lacking tarsungulus.

Pupa (P,).—Propping spines conspicuous, each with a spinelike seta at
‘apex; 4 small spines on head; 10 (6 very large) on pronotum; none on meso-

and metathorax; 4 (2 large) on abdominal segment I, 6 on II-VII, 4 (small)
on VIII.

Remarks.—With respect to the conspicuous setation of the body and the
leg structure in FG, and the distribution of propping spines in the pupa
(absent on mesothorax, few in number and strictly dorsal on abdomen),
Linsleya convexa is more epicautine than lyttine. However, it differs strik-
ingly from Epicauta, and at the same time agrees with the Lyttina and
Pyrotina, in most of the characters of the coarctate larva mentioned above.

In view of the paucity of published work on the comparative anatomy of
immature meloids other than the triungulin larva, it is difficult to interpret
ithe curious mixture of lyttine and non-lyttine characteristics found in Lin-

760 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

sleya. It is, however, apparent that there is little about the genus that is
specifically epicautine. Since grasshoppers eggs are the typical prey-type of
larval Mylabrina (MacSwain, 1956), as well as of Epicauta, even positive
proof that Linsleya utilizes this prey-type in nature would not constitute
definitive evidence that the genus is epicautine. Pinto (1974) claimed to find
special similarity in patterns of courtship of Linsleya and Epicauta, but his
treatment was based on a decidedly restricted survey of the range of be-
havioral variation in the tribe Lyttini, and I fail to find his argument con-
vincing. Indeed, it would appear that the only unequivocal basis for includ-
ing Linsleya in the Epicautina is the characteristic originally cited by
MacSwain (1951): the presence of lanceolate setae on the legs of the T larva.
An alternative interpretation, and one apparently not considered heretofore,
would be to place Linsleya in a separate subtribe, allied to the Epicautina
and Mylabrina, at least phenetically, on bionomic grounds.

ACKNOWLEDGMENTS

The drawing of the coarctate larva was done by Alice Prickett. Field work
was supported by a grant from the National Science Foundation.

LITERATURE CITED

Church, N. S. and G. H. Gerber. 1977. The development and habits of Linsleya sphaericollis
(Coleoptera: Meloidae). Can. Entomol. 109: 375-380.

Criddle, N. 1931. Grasshopper control in Canada east of the Rocky Mountains. Can. Dep.
Agric. Bull. 143(n.s.), 18 pp.

MacSwain, J. W. 1951. A new genus of Meloidae from North America. Pan-Pac. Entomol.
27: 58.

———. 1956. A classification of the first instar larvae of the Meloidae (Coleoptera). Univ.
Calif. Publ. Entomol. 12, 182 pp.

Pinto, J. D. 1974. Courtship behavior in Linsleya compressicornis and its taxonomic signif-
icance (Coleoptera: Meloidae). Pan-Pac. Entomol. 50: 1-8.

Romanov, W. 1954. Predators of grasshoppers. Proc. Entomol. Soc. Manit. 9: 14-18.

Selander, R. B. 1955. The blister beetles genus Linsleya (Coleoptera, Meloidae). Am. Mus.
Novit. 1730, 30 pp.

——. 1964. The systematic position of the genus Linsleya (Coleoptera: Meloidae). Proc.
Entomol. Soc. Wash. 66: 216.

Selander, R. B. and J. M. Mathieu. 1964. The ontogeny of blister beetles (Coleoptera, Me-
loidae). I. A study of three species of the genus Pyrota. Ann. Entomol. Soc. Am. 57:
711-732.

Selander, R. B. and J. D. Pinto. 1967. Sexual behavior in blister beetles (Coleoptera: Me-
loidae) I. Linsleya convexa. J. Kans. Entomol. Soc. 40: 396-412.

Selander, R. B. and R. C. Weddle. 1969. The ontogeny of blister beetles (Coleoptera, Me-
loidae). Il. The effects of age of trungulin larvae at feeding and temperature on devel-
opment in Epicauta segmenta. Ann. Entomol. Soc. Am. 62: 27-39.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 761-774

THE PERLODINAE OF VIRGINIA, USA
(PLECOPTERA: PERLODIDAE)

Boris C. KONDRATIEFF AND J. REESE VOSHELL, JR.

_ Department of Entomology, Virginia Polytechnic Institute and State Uni-
versity, Blacksburg, Virginia 24061.

Abstract.—The taxonomy and distribution of the adult Perlodinae of Vir-
ginia, USA, are reviewed. The ten species occurring in the state are illus-
trated, and keys separating adult males and females are provided. Biological
notes are also given for each species.

|

_ Ten species of Perlodinae were recorded by Kondratieff and Voshell
| (1979a) from Virginia. Ricker (1952) revised the species of Perlodinae of

North America and subgenera of the world. He placed the species that we
_ discuss in this paper in several subgenera of the genus /sogenus. Illies (1966)
elevated all Ricker’s subgenera to full generic rank. Since Ricker’s work
two new species that occur in the eastern United States have been described
(Stark and Gaufin, 1974; Kondratieff and Voshell, 1979b). The purpose of
| this paper is to provide comprehensive and up-to-date taxonomic descrip-
tions and keys to the species of Perlodinae that occur in Virginia and sur-
rounding states. In addition, we have included biological information on the
species.

Virginia (Fig. 1) is divided into five physiographic provinces (Hoffman
1969). These provinces range in elevation from sea level to 1743 m. This
allows for a considerable faunal diversity to exist. Generally the members
| of the Perlodinae are restricted to clear, cool, fast flowing montane streams.
Of the ten species of Perlodinae that occur in Virginia, seven are apparently
restricted to the montane Ridge and Valley, and Blue Ridge provinces. Only
eo duplicata (Banks), Helopicus subvarians (Banks), and Remenus
bilobatus (Needham and Claassen) have a wider distribution that includes
a Piedmont Plateau Province.

Adults are not commonly collected in the field, but may be collected
occasionally by sweeping riparian vegetation. In this study we obtained
most adult specimens by rearing field collected nymphs.

The following key, primarily using genitalic characters, will separate males

762 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

POWELL CLINCH N.FK S.FK Tenn = “ = Ine ee
HOLSTON HOLSTON ROANOKE MEHERRIN CHOWAN

+4 Sh ag, v/ > VAD) SV wae E
= OY, ES Sa Sh SG fa Gay Qa ~ ON aes i 74 (EX eu
Da Osama -} 2 OA > A = JA iy Sy Ly ce: [es "

= AS TKS = > Z t ==

a a
DISMAL SWAMP N.C

Fig. 1. Physiographic provinces of Virginia (according to Hoffman, 1969). CP = Coastal
Plain; PP = Piedmont Plateau; BR = Blue Ridge; RV = Ridge and Valley; AP = Appalachian
Plateau.

of the Virginia Perlodinae. Females are separable using head pattern com-
bined with the form of the subgenital plate.

KEY TO THE SPECIES OF VIRGINIA PERLODINAE

I. Tenth tergum completely cleft, subgenital plate absent from 8th
Stérmum,miale ss! 2 0) 2040 Ae ie had SEE TRE ES eae i
~ Tenth tergum entire; subgenital plate present on 8th sternum,
females ee OS, BER Go. a 11
pss A median mesosternal ridge present (Fig. 9); submental gills
long;epiproct hooked ‘at‘apex (Fig? 12) 82222". ee
sid 0 SyeNE ROG SECC: Ces RRC euch ate ee Isogenoides hansoni (Ricker)
- Median mesosternal ridge absent; submental gills short or ap-
parently absent; epiproct not hooked at apex 4.22. J. 2222 s2ene 3
3) Lateral stylets present (Figs-37'30) ©) .2.2 2 a eee 4
- [ateralstylets absent (Figs? 63°19) Se eee ee 8
4. Lateral stylets exceeding apex of epiproct by more than % of
their length?) 02S) es RE A 5
- Lateral stylets exceeding apex of epiproct by no more than 2
of their length? 52. 5 SA A eee 7
Ss Apex of lateral stylets distinctly 2-pronged (Fig. 34) ...........
Sa Moiese ee Diploperla morgani Kondratieff and Voshell
_ Apex of lateral stylets acute or somewhat truncate (foot-shaped) ..
oe seid ale a ale Fed eee lates Sees ene ene ate et SIRNA Ce St tet a 6
6. Apex of lateral stylets somewhat truncate or foot-shaped (Fig.
SDiniivacs Be Res tA ee Diploperla duplicata (Banks)

763

VOLUME 84, NUMBER 4

A
l2

Figs. 2-13. 2-4, Cultus decisus. 5-8. Helopicus subvarians. 9-13, Isogenoides hansoni. 2,
5, 10, Adult head and pronotum. 3, 6, 11, Male terminalia, dorsal. 4, 8, 13, Female subgenital

plate, ventral. 7, 12, Epiproct, lateral. 9, Mesosternum.

764

8(3).

12(11).

13(12).

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Apex of lateral stylets acute (Bie: 37) 5. 2 va.<0 ae oe eee
Pceasaecage eee K wreak Re Diploperla robusta Stark and Gaufin
Lateral stylets broad, tips distinctly hooked; epiproct covered
with golden-brown setae (Figs: 15; 16) .:...5-2-5... 4. cee
Ee Ae EE IE iF cee, 5 Malirekus hastatus (Banks)
Lateral stylets slender, tips straight; epiproct not covered with
setae (Pie. 3).8 .c.5.025 5 eee ee ee Cultus decisus (Walker)
Epiproct terminating in a reversible threadlike *‘lash’’; total body
length.8=9 mm (Fig. 26) 2.00.25 tad so: woe oe See ee
fe reste. EO se Remenus bilobatus (Needham and Claassen)
Epiproct not terminating in a threadlike structure but apex bul-
bous.or acute; body length.15—17 mm. %° 2... :27.- = - oe eee 9
Apex of epiproct bulbous, subcylindrical (Fig. 23) ..........
EGE aE, EK, ee is es Yugus bulbosus (Frison)
Apex.of epiproct acute: .. ..vmtecasctSn, a:8 ..2%cc0s ee ee 10
Pronotum without lateral yellow spots (Fig. 5) ..............
er a) RCT Ea Helopicus subvarians (Banks)
Pronotum with lateral yellow spots (Fig. 18) ................
Bee iret Ae ES ORS aro ee eee CeCe Yugus arinus (Frison) |
Mesosternal ridge on mesothorax present (Fig. 9); submental

CUNSMIONCS = 0b Ales joe Isogenoides hansoni (Ricker)

Mesosternal ridge on mesothorax absent; submental gills short |
of apparently absent® ...- 0.2 ee See ee 125 |
Subgenital plate with a deep medial emargination or rounded |

notch (Figs) 17,24): 3.40 ye. See eee 138}
Subgenital plate entire or slightly emarginate (Figs. 28, 35) .... 14 |
Dorsal head pattern distinct, ocellar triangle with a spear shaped
pale area (Fig. 14); subgenital plate with wide shallow rounded
median notch (Fick). 2... eee Malirekus hastatus (Banks) |
Dorsal head pattern indistinct, with diffuse pale area (Fig. 22);
subgenital plate with deep median notch, broadest distally (Fig.

DATE. ye 01S NEE YE RON een Re See Yugus bulbosus (Frison) |!
Subgenital! plate slightly emarginate ...2.2....- 55 -e2 eee 15 |
Subgenitalplate entire’ 42%) Sieaeere es oo ee eee 16 |

Dorsal head pattern with dark transverse band across ocelli
(Fig. 33); pronotum without lateral yellow spots; subgenital plate
somewhat truncate and usually slightly emarginate (Fig. 35). .
rs DENS ey eee Diploperla morgani Kondratieff and Voshell |
Dorsal head pattern with two dark patches posterior to ocelli
(Fig. 18); pronotum with lateral yellow spots; subgenital plate
rounded and usually expanded laterally and narrowly emargin-
ate: (Fige'21)* 223 Lee eee Yugus arinus (Frison)

VOLUME 84, NUMBER 4 765

ir

Figs. 14-24. 14-17, Malirekus hastatus. 18-21, Yugus arinus. 22-24, Y. bulbosus. 14, 18,
20, Epiproct, lateral. 17,

22, Adult head and pronotum. 15, 19, 23. Male terminalia, dorsal, 16,
21, 24, Female subgenital plate, ventral.

766 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

16. Total body length 8-9 mm; dorsal head pattern with ocellar
triangle uniformly pigmented and with areas of pigment gently
curving ‘anteriorly (Fig..25). oo. Tae ae ae ee ee

aay a ero ene Pere a Remenus bilobatus (Needham and Claassen)

- Total body length 15-17 mm; dorsal head pattern with ocellar
triangle either partially unpigmented (Figs. 2, 5), uniformly pig-
mented with diffuse pigment forming pointed areas anteriorly
(Fig. 29) or with a pale spot in front of anterior ocellus (Fig.

36) irerdin ds Tk See See RNS at ck os itd Se eee 17

17. Apical margin of subgenital plate narrowly rounded; dorsal head
pattern with a dark M-shaped band connecting ocelli ........

Sate. ee eRe. St eae TREO aga. 2 te Cultus decisus (Walker)

- Apical margin of subgenital plate broadly rounded; dorsal head

pattern without a dark M-shaped band connecting ocelli ...... 18
18. Dorsal head pattern with a cone-shaped pale area in the ocellar

triangle (Fig. 5); subgenital plate broadly rounded to slightly

truncate (Figs. 8). ....$2..4) sae Helopicus subvarians (Banks)

- Dorsal head pattern with ocellar triangle either uniformly pig-
mented with diffuse pigment forming pointed areas anteriorly
(Fig. 29) or with a pale spot in front of anterior ocellus (Fig.
36) wsheae bakes ss dees acces ease das dos Ore 19

193 Dorsal head pattern with ocellar triangle uniformly pigmented
with diffuse pigment forming pointed areas anteriorly (Fig. 29);
subgenital plate with sides usually subparallel (Fig. 32); general
body color light brown to yellow ... Diploperla duplicata (Banks)

- Dorsal head pattern with a pale spot in front of anterior ocellus
(Fig. 36); subgenital plate with sides usually broadly rounded
(Fig. 38); general body color dark brown »- 20. 4ees. sass eee
ebohatenae hime he SRR RO SEIN 1 Sesh Diploperla robusta Stark and Gaufin

Cultus decisus (Walker)
Figs. 2—4

Perla decisa Walker, 1852: 170. Type-locality, St. Martin’s Falls, Albany
River, Ontario, Canada.

Isogenus (Cultus) decisus, Ricker 1952: 97.

Cultus decisus (Walker), Illies, 1966: 356.

Male.—Length of forewing 12-13 mm; length of body 12-13 mm. General
body color in alcohol brown. Head and prothorax marked with pale yellow
(Fig. 2). Wings hyaline; veins dark brown. Lateral stylets slender and acute;
epiproct prominent and swollen near tip (Fig. 3).

Female.—Length of forewing 13-14 mm; length of body 13-14 mm. Color

VOLUME 84, NUMBER 4 767

Figs. 25-35. 25-28, Remenus hilobatus. 29-32, Diploperla duplicata. 33-35, D. morgani.
25, 29, 33, Adult head and pronotum. 26, 30. Male terminalia, dorsal. 27, Epiproct, lateral. 28,
32, 35, Female subgenital plate, ventral. 31, 34, Lateral stylet, lateral.

768 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

38
36

Figs. 36-38. Diploperla robusta. 36, Adult head and pronotum. 37, Lateral stylet, lateral.
38, Female subgenital plate, ventral.

pattern and external morphology similar to male. Subgenital plate produced
to 10th sternum (Fig. 3); apical margin narrowly rounded.

Nymph.—Described by Claassen (1931) as Perla verticalis Banks.

Number of adults examined.—32 46, 89.

Geographic Range.—Eastern North America.

Distribution in Virginia.—Counties: Carroll, Craig, Floyd, Giles, Mont-
gomery, Smyth, Washington.

In Virginia, adults were collected during May, usually from medium size
2nd and 3rd order montane streams. Adults are attracted to light.

Helopicus subvarians (Banks)
Figs. 5-8

Perla subvarians Banks, 1920: 317. Type-locality, Great Falls, Va.
Hydroperla subvarians, Frison, 1942: 292.

Isogenus (Helopicus) subvarians, Ricker, 1952: 103.

Helopicus subvarians, Ulies, 1966: 361.

Male.—Length of forewing 15—16 mm; length of body 16—17 mm. General
body color brown. Head and prothorax marked with pale yellow (Fig. 5).
Wings hyaline with some tinting of membrane; veins black. Lateral stylets
absent (Fig. 6). Epiproct tapering to an acute apex (Fig. 7).

Female.—Length of forewing 19-20 mm; length of body 19-20 mm. Color
pattern and external morphology similar to male. Subgenital plate rounded
to slightly truncate produced over % of sternum 9 (Fig. 8).

Nymph.—Described by Ricker (1952).

VOLUME 84, NUMBER 4 769

Number of adults examined.—19 4, 349.

Geographic range.—Eastern North America.

Distribution in Virginia.—Counties: Alleghany, Appomattox, Caroline,
Carroll, Fairfax, Floyd, Giles, Grayson, Hanover, Henry, Louisa, Mont-
gomery, Nottoway, Pulaski, Roanoke, Rockbridge, Smyth, Washington,
Wythe.

This common species apparently prefers large streams with cobble and
coarse pebble substrate, having been collected from 4th to 6th order rivers
from the Ridge and Valley Province into the Piedmont Province. Adults
may be collected from March to May. A large ovipositional flight of several
hundred females was observed 2 May 1980 at the Appomattox River from
5:45 PM to 6:30 PM. Females were coming from nearby trees and descend-
ing 8 to 10 m in a “‘fluttering glide’’ each dipping its abdomen in riffle
areas and releasing an egg pack of 75 to 100 eggs.

Isogenoides hansoni (Ricker)
Figs. 9-13

Isogenus Usogenoides) hansoni Ricker, 1952: 111. Type-locality, Broad-
head Creek, Analomink, Pa.
Isogenoides hansoni, Illies, 1966: 365.

Male.—Length of forewing 15—16 mm; length of body 16-17 mm. General
body color in alcohol black. Head and prothorax marked with yellow (Fig.
10). Submental gills long. Wings hyaline to slightly smoky; veins black.
Median mesosternal ridge present. Lobes of 10th tergum yellow. Lateral
stylets slender, slightly flattened with acute tip not exceeding epiproct (Fig.
11). Epiproct with tip recurved with apical hook (Fig. 12).

Female.—Length of forewing 20-21 mm; length of body 21-22 mm. Color
pattern and external morphology similar to male. Subgenital plate produced
4 to % length of 9th sternum (Fig. 13); apical margin broadly rounded to
arcuate.

Nymph.—Described by Ricker (1952).

Number of adults examined.—12 4, 25°.

Geographic range.—Eastern North America.

Distribution in Virginia —Counties: Giles, Madison, Nelson, Smyth,
Washington.

This is one of the rarest Perlodinae in the state. A large population occurs
in Little Stony Creek, St. Rt. 460 in Giles County. Other nymphs and adults
were collected from clear, cool, and fast flowing Ridge and Valley 2nd to
3rd order ‘“‘trout’’ streams. Adults were active from April to June.

Malirekus hastatus (Banks)
Figs. 14-17

Isogenus hastatus Banks, 1920: 314. Type-locality, Andrews, NC.

770 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Isogenus (Malirekus) hastatus, Ricker, 1952: 117.
Malirekus hastatus, Mlies, 1966: 369.

Male.—Length of forewing 18-19 mm; length of body 18-20 mm. General
body color in alcohol dark brown. Head and prothorax marked with pale
yellow (Fig. 14). Wings slightly smoky, veins dark brown. Lateral stylets
hooked at tip; epiproct densely covered with golden-brown setae (Figs. 15,
16).

Female.—Length of forewing 19-20 mm; length of body 19-20 mm. Color
pattern and external morphology similar to male. Subgenital plate produced
24 length of 9th sternum and deeply emarginate medially (Fig. 17).

Nymph.—Described by Claassen (1931).

Number of adults examined.—21 6, 24°.

Geographic range.—Eastern North America.

Distribution in Virginia—Counties: Albemarle, Alleghany, Augusta, Giles,
Grayson, Floyd, Highland, Madison, Montgomery, Nelson, Patrick, Roa-
noke, Rockingham, Smyth, Tazewell, Washington, Wythe.

This species is very abundant in Ist to 3rd order streams in the Ridge and
Valley and the Blue Ridge Provinces of Virginia. Adults are active from
April to early June.

Yugus arinus (Frison)
Figs. 18-21

Diploperla arina Frison, 1942: 309. Type-locality, Balsam, N.C.
Isogenus (Yugus) arinus, Ricker, 1952: 124.
Yugus arinus, Illies, 1966: 380.

Male.—Length of forewing 16-17 mm; length of body 17-18 mm. General
body color in alcohol brown. Head and prothorax marked with pale yellow
(Fig. 18). Wings hyaline; veins dark brown. Lateral stylets absent (Fig. 19).
Epiproct with tip slender and acute, transversely flattened (Fig. 20).

Female.—Length of forewing 19-20 mm; length of body 18-19 mm. Color
pattern and external morphology similar to male. Subgenital plate produced
*4 length of 9th sternum (Fig. 21); with a shallow apical emargination.

Nymph.—Described by Frison (1942).

Number of adults examined.—10¢, 21°.

Geographic range.—N.C., Tenn., Va.

Distribution in Virginia.—10¢, 202, Giles Co., Little Stony Creek, U.S.
460, Pembroke, 11 April 1979, July, B. C. Kondratieff; 12, Rappahan-
nock Co., Shenandoah National Park, Big Devils Stairs, 2 May 1976, O. S.
Flint, Jr.

This very rare stonefly has been collected in April and May from small,
clear, cool fast-flowing streams. Other Perlodidae collected in association

|

VOLUME 84, NUMBER 4 771

with Y. arinus were Cultus decisus, Isogenoides hansoni, and Isoperla na-
mata Frison. Female subgenital plate form varies from Frison’s (1942) figure
to Fig. 21, with the latter apparently more common.

Yugus bulbosus (Frison)
Figs. 22-24
Diploperla bulbosa Frison, 1942: 307. Type-locality, Greenbrier Cove, Tenn.
Isogenus (Yugus) bulbosus, Ricker, 152: 124.
Yugus bulbosus, Ullies, 1966: 380.

Male.—Length of forewing 16-17 mm; length of body 16-17 mm. General
body color in alcohol brown. Head marked with some yellow (Fig. 22).
Wings hyaline; veins brown. Lateral stylets absent. Epiproct with tip swol-
len, subcylindrical (Fig. 23).

Female.—Length of forewing 16-17 mm; length of body 17-18 mm. Color
pattern and external morphology simlar to male. Subgenital plate produced
24 length of 9th sternum (Fig. 24) with a deep medial emargination.

Nymph.—Described by Frison (1942).

Number of adults examined.—6d, 42.

Geographic range.—N.C., Pa., Tenn., Va., W. Va.

Distribution in Virginia.—Counties: Grayson, Patrick, Smyth, Tazewell.

Nymphs have been collected and reared from spring-fed intermittent
streams in Tazewell County (Burkes Garden) to several large, permanent,
clear, cool and fast-flowing streams in Grayson, Patrick, and Smyth coun-
ties. Adults are active during April and May. Adults and nymphs of Y.
arinus and Y. bulbosus are very distinct and readily separable.

Remenus bilobatus (Needham and Claassen)
Figs. 25-28
Perla bilobata Needham and Claassen, 1925: 95. Type-locality, Old Forge,
N:Y.

Isogenus (Remenus) bilobatus, Ricker, 1952: 123.
Remenus bilobatus, Ulies, 1966: 376.

Male.—Length of forewing 8-9 mm; length of body 8-9 mm. General body
color in alcohol light brown. Head and prothorax marked darker brown (Fig.
25). Wings hyaline; veins light brown. Lateral stylets absent. Epiproct ter-
minating in a long threadlike lash (Figs. 26, 27).

Female.—Length of forewing 10-11 mm; length of body 9-10 mm. Gen-

, eral color pattern and external morphology similar to male. Subgenital plate

produced 4 length of 9th sternum (Fig. 28); apical margin broadly rounded,
Nymph.—Described by Claassen (1931) and Frison (1942).

| Number of adults examined.—7d,4°.

Wiz PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Geographic range.—Eastern North America.

Distribution in Virginia——Counties: Bland, Culpeper, Floyd, Grayson,
Montgomery, Roanoke, Rockbridge, Smyth, Wythe.

This uncommon small species has been collected in late June and early
July from small spring-fed head-water streams to medium-size 5th and 6th
order rivers.

Diploperla duplicata (Banks)
Figs. 29-32

Perla duplicata Banks, 1920: 316. Type-locality, Newington, Fairfax
Co3iVa:

Isogenus (Diploperla) duplicatus, Ricker, 1952: 100. In part.

Diploperla duplicata, Mllies, 1966: 359. In Part.

Diploperla duplicata, Stark and Gaufin, 1974: 435.

Male.—Length of forewing 10-11 mm; length of body 9-10 mm. General
body color in alcohol yellow; head and prothorax darker yellow-brown (Fig.
29). Wings hyaline; veins yellow-brown. Tips of lateral stylets foot-shaped
(Figs. 30, 31).

Female.—Length of forewing 12-13 mm; length of body 11-12 mm. Color
pattern and external morphology similar to male. Subgenital plate 74 length
of 9th sternum (Fig. 32); sides somewhat subparallel, apical margin rounded.

Nymph.—Described by Kondratieff et al. (1981).

Number of adults examined._5d, 6°.

Geographic range.—Ga, S.C., Tenn., Va., W. Va.

Distribution in Virginia.—Counties: Bedford, Fairfax, Henry, Patrick,
Pittsylvania, Smyth.

This species has been collected in April and May from small creeks to
Sth order rivers from Ridge and Valley to the Piedmont Plateau Province.
Two of the males were collected in a blacklight trap.

Diploperla morgani Kondratieff and Voshell
Figs. 33-35

Diploperla morgani Kondertieff and Voshell, 1979b: 451. Type-locality,
Sinking Creek, Giles Co., Va.

Male.—Length of forewing 12-13 mm; length of body 12-13 mm. General
body color in alcohol dark yellow; head and prothorax darker brown (Fig.
33). Wings hyaline; veins yellow brown. Tips of lateral stylets 2-pronged
(Fig. 34). Epiproct reduced.

Female.—Length of forewing 16-17 mm; length of body 16—17 mm. Color
pattern and external morphology similar to male. Subgenital plate 74 length
of 9th sternum (Fig. 35); apical margin somewhat truncate and slightly emar-
ginate medially and occasionally entire.

VOLUME 84, NUMBER 4 773

Nymph.—Described by Kondratieff et al. (1981).

Number of adults examined.—Besides type-series, 53. 8°.

Geographic range.—Va., W. Va.

Distribution in Virginia—Specimens in addition to types: 36,59 (reared),
numerous nymphs, Montgomery Co., Mill Creek, Co. Rt. 785, 15-25 April
1980. B. C. Kondratieff; 1d, 2 2° (reared) Smyth Co., North Fork Holston
River, Co. Rt. 620 near junction Co. Rt. 716, 28 April 1981, B. C. Kondra-
tieff; 1d, Smyth Co., North Fork Holston River, St. Rt. 42, 9 May 1981,
B. C. Kondratieff; 1 2 (reared), 1 nymph, Wythe Co., Reed Creek, St. Rt.
11, 14 May 1977, R. F. Kirchner.

This rare species is known from several 3rd to 4th order clear, cool streams.
located in the Ridge and Valley Province. Mill Creek supports good trout
populations and has unusually high benthic diversity. Additional nymphs and
adults have been examined from several West Virginia localities.

Diploperla robusta Stark and Gaufin
Figs. 36-38

Diploperla robusta Stark and Gaufin, 1974: 434. Type-locality, Union Town-
ship, Tuscarawas Co., Ohio.

Isoperla duplicata, Frison, 1935: 499.

Isogenus (Diploperla) duplicatus, Ricker, 1952: 100. In part.

Diploperla duplicata, Ulies, 1966: 359. In part.

Male.—Length of forewing 12-13 mm; length of body 12—13 mm. General
body color in alcohol dark yellow to brown. Head and prothorax darker
brown (Fig. 36). Wings hyaline; veins brown. Tips of lateral stylets rounded
to acute (Fig. 37). Epiproct reduced.

Female.—Length of forewing 14-15 mm; length of body 14-15 mm. Color
pattern and external morphology similar to male. Subgenital plate */; length
of 9th sternum (Fig. 38); sides broadly rounded, apical margin rounded.

Nymph.—Described by Frison (1935) as /soperla duplicata (Banks) and
by Kondratieff et al. (1981).

Number of adults examined.—6d, 9°.

Geographic range.—Ind., Ky., Ohio, Pa., Va. W. Va.

Distribution in Virginia—Counties: Bedford, Craig, Giles, Montgomery,
Wythe.

This species has been collected from late April to mid-June from Ist to
4th order streams in the Ridge and Valley and Blue Ridge Provinces.

ACKNOWLEDGMENTS
We thank Ralph F, Kirchner, Huntington, West Virginia for reviewing
our manuscript and providing additional records. The following persons also
made specimens available for study: Oliver S. Flint, Jr., Richard L. Hoff-
man, Michael Kosztarab, and Joe L. Despins.

774 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The base map of Virginia was kindly provided by R. E. Jenkins, Roanoke
College. It is a modification of the U.S.G.S. State of Virginia, scale 1:
500,000 map, 1957 edition, and was compiled by N. M. Burkhead, D. J.
Jenkins, and R. E. Jenkins, with financial support from the Virginia Com-
mission of Game and Inland Fisheries. We especially thank Penelope F.
Kondratieff for the illustrations.

LITERATURE CITED

Banks, N. 1920. New neuropteroid insects. Bull. Mus. Comp. Zool. 64: 299-362.

Claassen, P. W. 1931. Plecoptera nymphs of America (north of Mexico). Thomas Say Found.
3: 1-199.

Frison, T. H. 1935. The stoneflies, or Plecoptera, of Illinois. Ill. Nat. Hist. Surv. Bull. 20:
281-471.

———. 1942. Studies of North American Plecoptera, with special reference to the fauna of
Illinois. Ill. Nat. Hist. Surv. Bull. 22: 235-355.

Hoffman, R. L. 1969. The insects of Virginia. No. 1. Pt. II. The biotic regions of Virginia.
Va. Polytech. Inst. Res. Div. Bull. 48: 23-62.

Illies, J. 1966. Katalog der rezenten Plecoptera. Das Tierreich. 82. Walter de Gruyter and
Co., Berlin. 632 pp.

Kondratieff, B. C. and J. R. Voshell, Jr. 1979a. A checklist of the stoneflies (Plecoptera) of
Virginia. Entomol. News 90: 241-246.

—. 1979b. A new species of Diploperla (Plecoptera: Perlodidae) from Virginia. Ann.
Entomol. Soc. Am. 72: 451-453.

Kondratieff, B. C., R. F. Kirchner, and J. R. Voshell, Jr. 1981. The Nymphs of Diploperla
(Plectopera: Perlodidae). Ann. Entomol. Soc. Am. 74: 428-430.

Needham, J. G. and P. W. Claassen. 1925. A monograph of the Plecoptera or stoneflies of
America North of Mexico. Thomas Say Found. 2: 1-397.

Ricker, W. E. 1952. Systematic studies in Plecoptera. Ind. Univ. Publ. Sci. Ser. 18: 1-200.

Stark, B. P. and A. R. Gaufin. 1974. The genus Diploperla (Plecoptera: Perlodidae). J. Kans.
Entomol. Soc. 47: 433-436.

Walker, F. 1852. Phryganides—Perlides, pp. 136-192. In Catalogue of the specimens of neu-
ropterous insects in the collection of the British Museum, I.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 775-780

EVIDENCE FOR REPRODUCTIVE ISOLATION BETWEEN XESTIA
ADELA FRANCLEMONT AND XESTIA DOLOSA FRANCLEMONT
(LEPIDOPTERA: NOCTUIDAE)

ANNE HUDSON

Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario
KIA 0C6.

Abstract.—Males of the sibling species Xestia adela Franclemont and X.
dolosa Franclemont respond preferentially to conspecific virgin females in
numbers that indicate the presence of specific differences in sex phero-
mones. No fertile eggs were obtained from laboratory crosses between the
species, although some mating occurred.

The results of pheromone studies within the species complex that includes
the Eurasian Xestia c-nigrum (L.) and the North American Xestia adela
Franclemont and Xestia dolosa Franclemont require some further investi-
gation when considered with relationships proposed on the basis of mor-
phological and genetic characters (Franclemont, 1980; Hudson and Lefko-
vitch, 1982).

Bestman et al. (1979) have shown that_X. c-nigrum (as Amathes c-nigrum)
in the vicinity of Frankfurt, West Germany are attracted to cis-7-tetra-
decenyl acetate. Similarly in Hokkaido, Japan, Fujimara (1976) and Hirai
(1976) found that cis-7-TDA attracted males of X. c-nigrum (also as A. c-
nigrum), whereas the trans isomer did not. Subsequently, Hirai (1976) ex-
tracted material from 2000 virgin females and found that the active factor
eliciting responses from antennal receptors was indeed cis-7-TDA. North
American representatives of the species, formerly known as large and small
forms of A. c-nigrum, were found by Roelofs and Comeau (1971) to be
differentially attracted to the two isomers, the large form (now X. dolosa)
to cis-7-TDA and the small form (now X. adela) to trans-7-TDA. Further
trials did not confirm these results, and it is possible that at this time some
contaminants were present in the synthetic attractants. European X. c-ni-
grum and North American X. adela have been shown to have more char-
acters in common than _X. adela and X. dolosa (Franclemont, 1980; Hudson,
1981; Hudson and Lefkovitch, 1982); specimens of A. c-nigrum obtained
from Osaka, Japan also appear to be morphologically and genetically closer

776 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

to X. adela (Hudson, unpublished data), so that such interchange of behav-
iour towards the isomers of 7-TDA would be unexpected.

The purpose of the present study was to determine if the sex attractants
for X. adela and X. dolosa are different by observing the attraction of males
from wild populations to traps baited with virgin females, and to examine
the possibility of interbreeding between laboratory colonies of the two
species.

MATERIAL AND METHODS

The moths used in field and laboratory experiments were second and third
generation insects reared from founder females obtained from North Gower
(NG) and Harrow (HA), Ontario. Several lines of each species were crossed
in the second generation. The moths were maintained in an incubator at
75°F and approximately 70% RH, with a photoperiod of 16 hours. Larvae
were reared on an artificial diet as described by Hinks and Byers (1976).

In pheromone experiments Pherocon IC (Zoecon) sticky traps were set
up near Mallorytown, Ontario in an area close to the St. Lawrence Seaway
where both X. adela and X. dolosa are known to occur. Twenty traps (8
baited with X. adela females; 8 with X. dolosa females; and 4 blanks) were
maintained during the period August 28 to September 24, 1980, when the
second brood of the wild populations were flying. Sixteen traps, including
the 4 blanks, were attached at a height of approximately | m along a fence
separating fields of oats and buckwheat from a wooded area; the remainder |
were hung from branches of small trees in the wooded area. The positions
of the traps were randomized. Baited traps contained two 3-day old virgin
females, each individually housed in a plastic mesh cage. The females were
replaced every third or fourth day and the trapped males were identified by
electrophoresis (Hudson and Lefkovitch, 1980) and by dissection (Hudson,
1981); sample collections have been retained in the Canadian National Col-
lection of insects in Ottawa.

Mating experiments were carried out in wooden frame, plastic-mesh cages
30 x 30 x 17 cm in a rearing room maintained at 70°F and 70% RH with a |
16 hour photoperiod. Five males and five females were used in all combi-
nations, they were put into the cages immediately following eclosion until
a substantial number of eggs had been laid in the control cage, then females |
were examined for spermatophores and in most cases for the presence of
sperm in the spermatheca.

An attractancy index (A) was calculated substituting A for I (isolation
index) in the formulae of Wasserman and Koepfer (1977):

No. conspecific ¢ attracted — No. other species 6 6 attracted
(CS) (OS)

ne AO), ee : ms
Total no. dd (both species) attracted (CS + OS)

VOLUME 84, NUMBER 4 177

Table 1. Observed numbers of males caught in female-baited traps at Mallorytown, August-
September, 1980. Expected numbers in parentheses.

? Bait

3 Caught X. adela X. dolosa Blank Total

X. adela 741 (660.9) 103 (183.1) 0 844

X. dolosa 6 (86.1) 104 (23.9) 0 110
Total 747 207 0 954 x”, = 388.4

Attractancy index (A) 0.76 + 0.022 0.89 + 0.044

The standard error was calculated as

2 Cas |.
CS + OS

A value of A > 2 X standard error is considered to indicate significant con-
specific attraction. A positive index indicates a preference by males for
conspecific females, a negative index that no preference is shown.

RESULTS

Attraction of field population males to virgin females.—The number of
males caught in traps baited with virgin females of each species (Table 1)
indicates significant conspecific attraction (chi-square = 388.4). Expressed
as an attractancy index (A) and standard error (SE), from the formulae
described in the previous section, for X. adela A = 0.76 + 0.022 and for
X. dolosa A = 0.89 + 0.044; these are significant (2 x SE < A).

The number of X. adela males collected remained fairly constant over
the total time period, for example 18.0% (153) were collected on August 28
and 18.7% (158) on September 16. The total number of X. dolosa males
collected was lower, but the numbers increased towards the end of the
period, 10.5% (11) were collected on August 28 and 42% (44) on September
16. Traps hung along the fence bordering the oat field collected 80% (678)
of all X. adela males and 72% (80) of all X. dolosa males, the remainder
were collected in the tree traps.

Interspecific breeding experiments.—Xestia adela adapts very easily to
laboratory rearing and 83% of the females in control cages were found to
contain from 1-4 spermatophores. Mating success was unaffected by cross-
ing NG and HA populations. Xestia dolosa intraspecific crosses were less
successful and only 38% of the females contained spermatophores at the
time of dissection. Interpopulation crosses beween X. dolosa males and
females produced infertile eggs only, but only two trials were made because
of a scarcity of X. dolosa NG at that time.

778 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Crossing experiments between X. adela and X. dolosa.

Max. No. % No. Females
Species combination Infertile Spermatophores that had
SS x SY No. Trials Fertile Eggs Eggs per Female Spermatophores
adela x adela 10 aati + 4 85
dolosa X dolosa 14 + tate 2 38
adela x dolosa 12 - + 3 15
dolosa X adela 11 ~— ++ | 5
1 ++ = over 200 eggs/2; + = less than 200 eggs/2 ; — = no eggs.

When X. adela males were caged with X. dolosa females, spermatophores
were deposited and sperm were found in the spermatheca of one female,
but no fertile eggs were laid (Table 2). Multiple mating was demonstrated
by one female that had three spermatophores. In this combination 15% of
the total number of females were found to have spermatophores either com-
pletely or partially deposited (with part extruding from the ductus bursae).
Several pairs were observed in copula and three of these were unable to
separate.

In crosses between X. dolosa males and X. adela females only 5% (2) of
the total number of females (55) had one spermatophore each. There was
no evidence of sperm in the spermathecae of these females and no fertile
eggs were obtained, although many infertile eggs were laid.

DISCUSSION

The results of attractancy tests using virgin females strongly indicate that
the sex pheromones of X. adela and X. dolosa are different, but that some
cross attractancy does occur. The highest proportion of cross attractancy
was between X. adela males and X. dolosa females, but some also occurred
in the reciprocal combination, particularly towards the end of the collecting
period when there were more X. dolosa males flying.

No fertile eggs were obtained from interspecific no-choice crosses set up
in the laboratory. The evidence that mating occurred between both com-
binations of males and females and yet only infertile eggs were produced
indicates that circumvention of the long-distance (anemotactic) responses
(Roelofs and Cardé, 1977) did not induce hybridization, and it probably
means that both premating and post-mating isolation mechanisms are acting
together to maintain the taxonomic discreteness of the species.

A measure of genetic differentiation between X. adela and X. dolosa and
an European population of X. c-nigrum was obtained from an electropho-
retic study of allozyme frequencies (Hudson and Lefkovitch, 1982). Dis-
tances between the species were calculated from the formulae of Nei (1972)
as genetic distance (D) and suggested a closer relationship between X. c-

VOLUME 84, NUMBER 4 779

nigrum and X. adela (D = 0.104) and between X. c-nigrum and X. dolosa
(D = 0.260) than between X. adela and X. dolosa (D = 0.319). These re-
lationships could be interpreted as the result of two separate introductions
of X. c-nigrum, one of which, as X. dolosa, remained restricted to the
eastern and central regions of N. America, and the other, as X. adela,
became distributed transcontinentally. Where the two species are sympatric
reproductive isolation has been achieved in part by pheromone differentia-
tion, possibly brought about by the presence of different proportions of
attractant components; this could explain the cross-attractancy observed in
our experiment.

The larger number of X. adela males attracted to X. dolosa females could
be interpreted as a one-sided mating preference, but is more likely to be
due to the larger number of X. adela males in the field population during
the observation period.

ACKNOWLEDGMENTS

I thank J. R. Byers and J. D. Lafontaine for helpful discussions and J.
W. van der Meer and W. H. Forrest for excellent technical assistance.

LITERATURE CITED

Bestmann, H. J., O. Vostrowsky, H. Platz, T. Brosche, K. H. Koschatzky, and W. Knauf.
1979. Pheromone XXIII. (Z)-7-Tetradecenylacetat, ein sexual-lockstoff fiir mannchen
von Amathes c-nigrum (Noctuidae; Lepidoptera). Tetrahedron Lett. (6) 497-500.

Franclemont, J. G. 1980. ‘‘Noctua c-nigrum”’ in eastern North America, the description of
two new species of Xestia Hiibner (Lepidoptera: Noctuidae: Noctuinae). Proc. Entomol.
Soc. Wash. 82: 576-586.

Fujimura, T. 1976. Mating behavior and sex attractant of the spotted cutworm moth, Amathes
c-nigrum L. (Lepidoptera: Noctuidae). Jpn. J. Appl. Entomol. Zool. 20: 133-138.
Hinks, C. F. and J. R. Byers. 1976. Biosystematics of the genus Euxoa (Lepidoptera: Noc-
tuidae) V. Rearing procedures and life cycles of 36 species. Can. Entomol. 108: 1345-

1357.

Hirai, Y. 1976. Cis-7-tetradecen-l-ol acetate: a sex pheromone component of the spotted
curworm moth, Amathes c-nigrum, p. 165. In Kond, T. and S. Ishii, eds., Proc. of a
Symposium on Insect pheromones and their applications. Japan Plant Protection As-
sociation.

Hudson, A. 1981. Distinguishing characters of the reproductive system and genitalia of Xestia
dolosa and Xestia adela (Lepidoptera: Noctuidae). Proc. Entomol. Soc. Wash. 83: 413-
420.

Hudson, A. and L. P. Lefkovitch. 1980. Two species of the Amathes c-nigrum complex

(Lepidoptera: Noctuidae) distinguished by isozymes of adenylate kinase and by selected

morphological characters. Proc. Entomol. Soc. Wash. 82: 587-598.

1982. Allozyme variation in four Ontario populations of Xestia adela and Xestia
dolosa and in a British population of Xestia c-nigrum (Lepidoptera: Noctuidae). Ann.
Entomol. Soc. Am. 75: 250-256.

Nei, M. 1972. Genetic distances between populations. Am. Nat. 106: 283-292.

Roelofs, W. L. and R. T. Cardé. 1977. Responses of Lepidoptera to synthetic sex pheromone
chemicals and their analogues. Annu. Rev. Entomol. 22: 377-405.

780 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Roelofs, W. L. and A. Comeau. 1971. Sex attractants in Lepidoptera. Proc. Int. Congr.

Pesticide Chem., IUPAC, Tel Aviv, Israel 2: 91-1 12F
Wasserman, M. and H. R. Koepfer. 1977. Character displacement for sexual isolation be-
tween Drosophila mojavensis and Drosophila arizonensis. Evolution. 31: 812-823.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 781-785

A NEW LIRIOMYZA MINING LEAFLETS OF BLACK LOCUST
(DIPTERA: AGROMYZIDAE)

K. R. VALLEY

Bureau of Plant Industry, Pennsylvania Department of Agriculture, Har-
risburg, Pennsylvania 17110.

Abstract.—Liriomyza robiniae, new species, an early season leaflet miner
on black locust, Robinia pseudoacacia L., is described from Harrisburg,
Pennsylvania. Records of larval collections are given for New York, Penn-
sylvania, Virginia, and West Virginia, and some biological information is
presented.

The larva of Liriomyza robiniae, new species, forms a blotch mine (Fig.
1) on the leaflet of black locust, Robinia pseudoacacia L. (Faba-
ceae(=Leguminosae)). Adults were collected in late April in south central
Pennsylvania, and larvae were found in May in New York, Pennsylvania,
Virginia, and West Virginia. Only one generation is produced annually, the
pupae overwintering.

Apparently the only record of any agromyzid feeding on black locust is
that of Weaver and Dorsey (1965), who reared three species of Eulophidae
(Hymenoptera) from larvae of an unidentifed agromyzid leafminer on R.
pseudoacacia in West Virginia. A conversation with J. E. Weaver, West
Virginia University, leaves little doubt that the parasitized larvae he col-
lected were those of L. robiniae.

Liriomyza robiniae Valley, NEw SPECIES
Figs. 1, 2

Male.—Head: Front yellow, sometimes light orange anteriorly, darkened
at base of each fronto-orbital bristle, except foremost | or 2; ocellar area
black: 3 lower fronto-orbital bristles turned mesad; 2 upper fronto-orbitals
reclinate, anterior one somewhat the larger; front narrowly raised above
eye, slightly more than twice width of one eye at level of anterior ocellus;
both vertical bristles in black area, inner one at margin of dark color; par-
afrontal setulae lacking; gena (jowl) yellow, approximately 0.6 eye height;
face yellow; palpi light orange; antenna with 2 basal segments yellow, 3rd
segment round, orange, sometimes with light infuscation on outer basal 2;

782 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

a West
SS - . = ae
© a ee :
Fig. 1. Mine of L. robiniae in leaflet of black locust.

arista brown, pubescent, slightly longer than antenna, evenly tapering from
rather thick base.

Thorax: Scutum black, densely gray tomentose with humerus and noto-
pleuron largely yellow, black spot on anterior face of humerus; pleura yel-
low and black as follows: propleuron yellow; mesopleuron with black largely
confined to lower 2; pteropleuron largely yellow; hypopleuron largely black;
sternopleuron yellow along dorsal border, broadly around sternopleural
bristle, narowly anterior thereof; scutellum black, gray tomentose, yellow
mesally, especially between apical bristles, sometimes only faintly yellowish
basally; scutum with 4 irregular rows of acrostichal setulae anteriorly, grad-
ing into more regular rows behind transverse sulcus; wing 1.6—1.8 mm long,
hyaline, costa extending to M (4th vein), anterior crossvein approximately
opposite costal break; last section of Cu (Sth vein) 1.4-1.5 x as long as discal
cell; halter yellow; squamal cord and fringe dark brown; legs yellow, coxae
and femora partly blackish.

Abdomen: Dorsum black, with narrow yellow posterior margin of seg-
ments I—6 and yellow anterior corners of segment 1; postabdomen as in Fig.
2, epandrium yellowish, with yellowish bristles; ventral lobe (surstylus)
rounded with a few minute pale triangular scaly bristles; cerci not evident;
aedeagus slightly infuscated basally, otherwise colorless, details of disti-
phallus difficult to distinguish, but pair of narrow approximated processes
at apex (Fig. 2B); sperm pump (Fig. 2A) unusually small, on short duct.

Female.—External characters similar to male but with 3rd antennal seg-
ment sometimes more heavily infuscated and legs darker. Wing length 1.8-
2.0 mm.

Type-Material.—All specimens have the following locality and host in-

|

VOLUME 84, NUMBER 4 783

Fig. 2. Postabdomen of L. robiniae male, ventrolateral view. Drawn from macerated prep-
aration in water. A, Sperm pump, from microslide preparation. B, Outline of tip of aedeagus
as seen in direction of arrow above in dry specimen; not to scale.

formation: PENNSYLVANIA: Dauphin Co., Harrisburg, 2301 N. Cameron
St., taken on Robinia pseudoacacia. Holotype d: 22 April 1981, A. G.
Wheeler, Jr. (USNM). Allotype 2: 30 April 1979, AGW (USNM). Para-
types: 1¢, 12, 25 April 1979, AGW (CU): 29, 24 April 1980, T. J. Henry,
12, 23 April 1980, KRV, and 1 6 25 April 1979, AGW (PDA); 1d, 24
April 1979, KRV, and 1 2, 25 April 1979, KRV (USNM).

Specimens deposited in the Cornell University Insect Collection, Ithaca,
New York (CU); Pennsylvania Department of Agriculture, Harrisburg
(PDA); and the National Museum of Natural History, Washington, D.C.
(USNM).

Relationships.—Liriomyza robiniae is anomalous in the genus Liriomyza
because of its largely black scutellum, densely tomentose scutum, and post-
abdominal structure, especially the reduced cerci, colorless distiphallus that
is difficult to examine, and the very small, peculiarly shaped sperm pump.
However, a microscope preparation confirms its assignment to Liriomyza
by the presence of fine, abundant stridulatory spicules on the lateral abdom-
inal membrane of the male. Its only close relative, at least in North America,
seems to be a species from an unknown host in California, now in press in
K. A. Spencer’s comprehensive work on the Agromyzidae of California.
That species has a wholly black scutellum and has been given a name re-
ferring thereto; it lacks parafrontal setulae and the postabdomen is appar-
ently fairly similar to that of L. robiniae. It also differs from L. robiniae in
having but one upper fronto-orbital bristle, a subshining scutum, and black
pleura except for the narrow yellow upper margin of mesopleuron.

Comments.—Liriomyza robiniae runs to couplet 38 in Spencer's (1969)
key, although specimens do not agree with either choice, L. kenti Spencer
or L. nordica Spencer. In Frick’s (1959) key, specimens of L. robiniae trace
to L. eupatorii (Kaltenbach), a leafminer on Aster and Eupatorium in North
America.

784 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Larvae of L. robiniae were collected in black locust leaflets at the follow-
ing localities: NEW YORK: Delaware Co., Rt. 97, 2.6 mi. S. jct. with old
Rt. 17 in Hancock, 16 May 1979 and 27 May 1981, KRV; Cadosia, 16 May
1980, KRV; Sullivan Co., Long Eddy, along Rt 97, 16 May 1980 and 27
May 1981, KRV; Tompkins Co., Ithaca, Cornell University, 17 May 1980,
A. G. Wheeler, Jr.; Suffolk Co., Long Island, Yaphank, 30 May 1981, AGW.
PENNSYLVANIA: Centre Co., Rt. 45 nr. jct. with Rt. 26, betw. Pine
Grove Mills & Shingletown, 19 May 1980, AGW & KRV; Dauphin Co.,
Linglestown, 9 May 1979 and | May 1980, AGW & KRV; Clark’s Valley,
Rt. 325, 18 May 1979, KRV; Wayne Co., Damascus Twp., on road to Cal-
licoon, N.Y., nr. Curtis Nursery, 17 May 1979, KRV. VIRGINIA: Augusta
Co., I-81 S., 5.4 mi. N. Rt. 606, Raphine-Steeles Tavern Exit, 10 May 1979,
AGW: Montgomery Co., I-81, 2 mi. S. Roanoke Co. line, 26 April 1981,
AGW; Lancaster Co., Windmill Pt., 8 May 1981, AGW & KRV; James Co.,
Williamsburg, Wm. & Mary Campus, 9 May 1981, AGW & KRV. WEST
VIRGINIA: Berkeley Co., I-81 Rest Area W. of Potomac River, 10 May
1979, AGW.

Detection of L. robiniae has been hindered by the early and ephemeral
appearance of the adults and the short larval feeding period. The type-
material was collected shortly after leaf flush of black locust in the Harris-
burg area, when leaves were less than 4 cm long. In 1979, abandoned mines
were found as early as 9 May, and by 21 May no larvae were found in any
of the leaflets sampled. Small, heavily mined leaflets dropped from the trees;
larger leaflets with mines remained on the trees, but by 29 May the mined
areas had begun to dry and drop from the leaflets, thus preventing easy
detection of the larval feeding activities. By mid-June little evidence re-
mained to document the mining activities of the larvae.

The collecting records show this species is widespread, and I suspect that
additional early-season efforts will yield new distribution records from
throughout the native range of black locust. The tree is not native to states
and provinces north of Pennsylvania, where it has been naturalized north
to Nova Scotia, Quebec, and Ontario (Fernald, 1950). Thus L. robiniae
probably has been introduced into at least New York State through man’s
commerce or has followed the tree’s movement along highway banks, waste
land, and other areas disturbed by man.

ACKNOWLEDGMENTS

I thank George C. Steyskal, Systematic Entomology Laboratory, USDA,
Washington, D.C., for reviewing the manuscript and especially for his as-
sistance with the description, comments on relationships, and for Fig. 2. I
also thank my Harrisburg colleague, A. G. Wheeler, Jr., for reviewing the
paper and making suggestions for its improvement, and who, along with

VOLUME 84, NUMBER 4 785

Thomas J. Henry, Systematic Entomology Laboratory, USDA, collected
specimens of L. robiniae. J. E. Weaver, West Virginia University, Mor-
gantown, shared with me his observations on L. robiniae.

LITERATURE CITED

Fernald, M. L. 1950. Gray’s manual of botany. American Book Co., New York. 1632 pp.

Frick, K. E. 1959. Synopsis of the species of agromyzid leaf miners described from North
America (Diptera). Proc. U.S. Natl. Mus. 108: 347-465.

Spencer, K. A. 1969. The Agromyzidae of Canada and Alaska. Mem. Entomol. Soc. Can.
64, 311 pp.

Weaver, J. E. and C. K. Dorsey. 1965. Parasites and predators associated with five species
of leaf-mining insects in black locust. Ann. Entomol. Soc. Am. 58: 933-934.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 786-790

A NEW ALLOCAPNIA FROM WEST VIRGINIA
(PLECOPTERA: CAPNIIDAE)

RALPH F. KIRCHNER!

Department of the Army, Huntington District Corps of Engineers, Water
Quality Section, P.O. Box 2127, Huntington, West Virginia 25721.

Abstract.—Allocapnia frumi, n. sp., is described from specimens collect-
ed in West Virginia. It is included in the forbesi Group and is closely related
to A. curiosa Frison. A holotype male, allotype, and 37 paratypes are des-
ignated.

During an investigation of the winter stoneflies from Fayette, Greenbrier,
and Pocahontas counties, West Virginia, an undescribed species of Allo-
capnia was discovered. The following description and morphological terms
generally follow those of Ross and Ricker (1971).

Allocapnia frumi Kirchner, NEw SPEcIES

Male (Figs. 1-4).—Dark brown, almost black. Wings variable in length,
reaching from 2nd to Sth segment of abdomen. Length of body 5-7 mm.
Anterior 2 of 7th tergum produced into a conical process. Dorsal process
of 8th tergum with apex angled slightly forward with well separated lobes
forming a deep trough, anterior portion with a distinct white oval membra-
neous area. Apical segment of upper limb of epiproct about % length of
entire process and diamond shaped in dorsal aspect. Lower limb of epiproct
with apical segment moderately deep, its tip narrowed into a thin tongue.

Female (Figs. 5—7).—Dark brown, almost black. Wings variable in length,
reaching from 3rd to 9th segment of abdomen. Length of body 6-8 mm.
Eighth tergum entirely sclerotized to nearly so, or a mesal membranous area |
extending its whole length. Seventh and 8th sterna heavily sclerotized. Sev-
enth with posterior margin projecting well over anterior margin of 8th ster-
num. Eighth sternum with conspicuous lateral lobes, mesal area wide and
flat, its apex produced into a truncate lobe.

' The views of the author do not purport to reflect the position of the Department of the
Army or the Department of Defense. |

VOLUME 84, NUMBER 4 787

tetra

Figs. 1-4. Allocapnia frumi (male), terminalia. 1, Dorsal. 2, Lateral. 3, Dorsal process of

) 8th tergum, posterior aspect. 4, Dorsal process of 7th tergum, posterior aspect.

Type material—Holotype ¢ and Allotype, West Virginia, Greenbrier Co.,
Monongahela National Forest, Coats Run of North Fork Cherry River, 8.0
km east of Richwood, Rt. 39, 14 March 1981, R. F. Kirchner. Paratypes:
WEST VIRGINIA: Greenbrier Co., Coats Run, 3d, 17 March 1979, R. F.

788 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 5-7. Allocapnia frumi (female), terminalia. 5, Ventral. 6, Dorsal. 7, Lateral.

Kirchner and R. M. Meyer; 2¢, 12, 23 February 1980, R. F. Kirchner; 2
3,32, 16 March 1980, R. F. Kirchner, R. M. Meyer, and V. J. Marchese;
1d, 28 March 1981, R. F. Kirchner and V. J. Marchese. Pocahontas Co.,
Monongahela National Forest, Hills Creek, 24.1 km east of Richwood, Rt.

VOLUME 84, NUMBER 4 789

39, 1d, 16 March 1980, R. F. Kirchner, R. M. Meyer and V. J. Marchese.
Pocahontas Co., Monongahela National Forest, Sugar Creek of Williams
River, Forest Service Rt. 76, 52, 8 May 1980, R. F. Kirchner; 3¢, 9°,
3 May 1981, R. F. Kirchner and V. J. Marchese. Fayette Co., Big Hollow
of Paint Creek, 3.2 km northwest of Kingston, W. Va. turnpike, 3 ¢, 24
January 1980, R. F. Kirchner and L. K. Evans; 1d, 39, 27 February 1981,
R. F. Kirchner.

The holotype, allotype, and two paratypes will be deposited in the Na-
tional Museum of Natural History, Washington, D.C. (type No. 100222).
Other paratypes will be deposited in the collections of the Illinois Natural
History Survey, Champaign; Virginia Polytechnic Institute and State Uni-
versity, Blacksburg; C. H. Nelson, University of Tennessee at Chattanooga;
P. P. Harper, University of Montreal; and my personal collection.

Diagnosis.—Allocapnia frumi is a member of the forbesi group of Ross
and Ricker (1971), which includes A. curiosa Frison, A. forbesi Frison, A.
maria Hanson, A. minima (Newport), A. ozarkana Ross, and A. pechumani
Ross and Ricker. Within this group, A. frumi is most closely related to A.
curiosa in having the male terga highly ornamented and the female 7th
sternum well produced over the 8th sternum. The male of this new species
can be separated from others in the group by the prominent oval membra-
nous area of the 8th tergum (Fig. 1) (one male had an oval membranous
area on both the 7th and 8th terga), which is lacking in males of the other
species. The female of A. frumi may be distinguished by a wide truncate
mesal lobe of the 8th sternum (Fig. 5). In A. curiosa, the mesal area is
rounded posteriorly and without a mesal lobe (Ross and Ricker, 1971, Fig. 69).

Etymology.—This species is named in honor of Prof. W. Gene Frum,
Department of Biological Sciences, Marshall University. *

Remarks.—Allocapnia frumi was collected from first order mountain
streams ranging in altitude from 427 m in Fayette County to 1220 m in
Pocahontas County. The dominant riparian vegetation varied from cove
hardwoods to nearly pure stands of red spurce. Other Capniidae associated
with this species included A. curiosa Frison, A. frisoni Ross and Ricker,
A. harperi Kirchner, A. nivicola (Fitch), and Paracapnia angulata Hanson.
Allocapnia harperi was previously known only from the type-locality in
Virginia (Kirchner, 1980). It was collected from Greenbrier and Pocahontas
counties during this study.

ACKNOWLEDGMENTS

I am indebted to B. C. Kondratieff and J. R. Voshell, Jr., Virginia Poly-
technic Institute and State University, Blacksburg, for reviewing the manu-
script and to Penelope F. Kondratieff for preparing the illustrations. I also
thank P. P. Harper, University of Montreal, Quebec, for loan of specimens
of Allocapnia pechumani.

790 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

LITERATURE CITED

Kirchner, R. F. 1980. A new Allocapnia from Virginia (Plecoptera: Capniidae). Entomol.

News. 91: 19-21.
Ross. H. H. and W. E. Ricker. 1971. The classification, evolution, and dispersal of the winter

stonefly genus Allocapnia. Ill. Biol. Monogr. 45: 1-166.

—$— Le

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 791-796

CHROMOSOME AND ISOZYME STUDIES IN TRICHOGRAMMA
(HYMENOPTERA: TRICHOGRAMMATIDAE)!

AKEY C. F. HUNG

College of Agriculture and Life Sciences, University of Guam, Mangilao,
Guam 96913; present address: Beneficial Insect Introduction Laboratory,
IIBUI, Agricultural Research Service, Building 417, BARC-East, Beltsville,
Maryland 2070S.

Abstract.—Karyotype and isozyme analyses were carried out in Tricho-
gramma chilonis Ishii, T. evanescens Westwood, T. nubilale Ertle and Da-
vis, and T. pretiosum Riley. No significant differences were detected among
karyotypes of these four species as they all have n = 2SM + 2T + 1A.
However, these species can be readily distinguished on zymograms of su-
peroxide dismutase and esterase.

Trichogramma have been widely used in biological control projects in
various parts of the world. However, because of the inadequacy of classical,
morphological studies of Trichogramma, the precise identities of the species
and strains used are in most cases unknown. This problem has been some-
what alleviated by the use of male genitalia as a diagnostic morphological
character and crossbreeding experiments as a genetic approach in biosys-
tematics of this group of parasitic insects (see review article by Nagarkatti
and Nagaraja, 1977). As pointed out by Harland and Atteck (1933), the basic
solution to the problems of systematics in Trichogramma is the study of
biological characters, crossing relationships, cytology, and genetics of var-
ious races and species.

Due to the minute size of Trichogramma, cytological and isozyme studies
in these insects are rather difficult and have not been adequately carried
out. The only cytological study of Trichogramma was that of Fukada and
Takemura (1943). However, no detailed karyotype analysis was given in
their report. Voegele and Berge (1976) published the first paper on electro-
phoresis of Trichogramma. Further isozyme studies have been carried out
by Voegele and his associates (Jardak et al., 1979: Pintureau et al., 1980;

1 Approved as TA 21 by the Director of the Guam Agricultural Experiment Station.

792 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pintureau and Babault, 1981). Their results look promising for use in species
differentiation of these difficult parasitic Hymenoptera.

In 1979-80, karyotype and isozyme analyses were carried out at the Uni-
versity of Guam in order to determine the value of these two techniques in
the biosystematics of Trichogramma. The results of these studies are pre-
sented here.

MATERIALS AND METHODS

The following four species of Trichogramma were used in these studies:
T. chilonis Ishii (reared from sphinx moth eggs collected from wild taro
leaves on Guam), 7. evanescens Westwood (from a laboratory culture of
W. J. Lewis, USDA, Tifton, Georgia; positive identification of this sample
has yet to be determined), 7. nubilale Ertle and Davis (from a laboratory
culture of P. P. Burbutis, University of Delaware, Newark), and 7. pre-
tiosum Riley (from a laboratory culture of R. K. Morrison, USDA, College
Station, Texas). With the exception of 7. evanescens, species identifications
were confirmed by the author. Voucher specimens of these four species are
deposited at the Beneficial Insect Introduction Laboratory, II[BIII, USDA,
Beltsville, Maryland. :

Field collected eggs of a sphinx moth from wild taro were used as host
eggs. Freshly laid host eggs were collected in the morning, removed from
taro leaves with a camel hair brush, and stored at —10°C in a sealed vial
for three weeks in order to kill any egg parasites that might have already
parasitized the eggs. After three weeks, the host eggs were removed from |
the freezer and thawed at room temperature. They were glued in a cluster
of ten with Elmer’s glue® to a strip of filter paper and used for rearing the |
Trichogramma species according to the method of Morrison (1970).

Chromosome preparation.—Due to the minute size of Trichogramma, the |
technique of Hung et al. (1972) was modified. Four to five days after the |
host eggs were stung, Trichogramma pupae with light pink eyes or prepupae |
were removed from the host eggs and placed on a drop of colchicine-hy- |
potonic solution on a microslide. Testes and ovaries of pupae or brains of |
prepupae were dissected out and pushed aside to a corner. The remainder |
of the tissue was then wiped off the slide. The organs to be used were kept |
on the slide with another drop of colchicine-hypotonic solution for ten min- |
utes. To prevent the organs from drying out, the droplet was covered with |
a depression slide. After ten minutes, the solution was carefully removed
with a piece of filter paper. Special care was used to avoid contact of the |
filter paper with the organs and at the same time not to dry out the organs. },
Before the organs dried out, a wax pencil mark was made on the other side | |
of the slide to indicate the position of the organs. A drop of aceto-orcein |
was then placed on the organs and covered with a cover glass. The materials |
were squashed between filter paper with the thumb. One drop of immersion |

VOLUME 84, NUMBER 4 793

oil was placed on the slide at the position opposite to the wax pencil mark.
The pencil mark was then wiped off, and the slide was finally placed under
the microscope for examination. Chromosome photographs were taken with
Wild M20-EB Phase Microscope® using Fuji Minicopy film®.
Electrophoresis.—The vertical dual slab cell (Bio-Rad Model 220) with
0.75 x 140 mm spacer and Buchler 3-1500° constant power supply were
used. The Laemmli system (Laemmli 1970) of reagents and acrylamid gels
for SDS electrophoresis was followed in gel preparation, except that in all
cases SDS was omitted and sample buffer was replaced with 10% sucrose.
At least five individuals of each species were used for each sample well.
The wasps were homogenized in 0.01 ml of 10% sucrose on a 12-cavity
white porcelain plate with a glass rod. The homogenate in each cavity was
obsorbed onto a very narrow strip of Whatman #1 filter paper (ca. 5 x 0.1
mm). These strips were inserted into the wells already filled with electrode
buffer (tris base 6.0 g, glycine 28.8 g q.s. to | liter with deionized water).
The gel was initially run with constant current of 15 mA until the tracking
dye reached the separating gel which took about 3 hours. It was then run
for another 2 hours at constant power of 10 watts at which time the gel front
would have reached the bottom of the separating gel. During the run, the
gel was cooled with circulating ice water. After electrophoresis, the gel was
- stained for either esterase or superoxide dismutase. The esterase stain con-
sists of 100 ml sodium phosphate buffer, 0.12 M (pH 6.0); 1 ml 1% beta-
napthyl acetate (w/v) in 50% acetone-water (v/v); and 40 mg Fast Garnet
GBC salt. The stain for superoxide dismutase was composed of 15 mg NADP,
| 15 mg NBT, 5 mg PMS, 50 mg MgCl, and 75 ml 0.2 M Tris-HCl buffer (pH
8.0). All four species were used in esterase assay; however, the culture of
T. evanescens died later and was thus not included in superoxide dismutase
studies.

RESULTS AND DISCUSSION

As shown in Figs. 1-6, these four species of Trichogramma each have
n = 5 which is the same as that reported by Fukada and Takemura (1943)
for seven ‘‘strains’’ of Trichogramma sp. and is the predominant haploid
number in chalcidoids (Crozier, 1977). Further analyses revealed that they
not only have the same haploid number, but the morphology of these five
chromosomes in each species are the same, i.e. n = 2SM + 2T + IA as
indicated in the idiogram in Fig. 7. This striking homogeneity in the karyo-
types of these four species representing two different species groups (Na-
| garkatti and Nagaraja, 1977) is significant in the application of karyotype
analysis in the biosystematics of Trichogramma. It indicates that more ad-
vanced cytological techniques such as chromosome banding may have to
be developed for these minute insects.
Although no significant differences have been detected among karyotypes

794 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON |

ay Pe

-.

VOLUME 84, NUMBER 4 795

30

==) = =) =|
a= PR
30 3) Ba
NU
40- Ge
20 eee
‘f
17 10}- — |)
t=]
a it
Al ao =
1®) BB eet

50> == -—o
7 RE AS lal
Figs. 7, 8. 7, Idiogram based on Figs. 1-6; TCL = total chromosome length. 8, Diagram-

matic illustrations of superoxide dismutase (top) and esterase (bottom) banding patterns in
Trichogramma. CH = chilonis; EV = evanescens; NU = nubilale; PR = pretiosum.

efoml Gal

of these four Trichogramma species, they can be readily distinguished on
zymograms of superoxide dismutase and esterase (Fig. 8). Formal progeny
analyses to establish the number of loci have not been carried out because
at least five wasps have to be used in each well in order to obtain detectable
enzyme activity due to the level of sensitivity of slab gel. Nevertheless, the
banding patterns of the two enzymes are unique in each of these four species.
The esterase zymogram of T. evanescens reported here is different from
that of Pintureau and Babault (1981). Perhaps my sample is not evanescens
as I have questioned. Furthermore, whether the three-band phenotype of
esterase found in 7. evanescens represents a heterozygote at this locus has
yet to be determined.

A pooled sample of male progeny of a single virgin female has been used
to circumvent the difficulty of assaying a single wasp (Pintureau et al., 1980;

—

Figs. 1-6. Haploid and diploid karyotypes. 1, 2, Trichogramma chilonis, male and female.
3, 4, T. evanescens, male and female. 5, 7. nubilale, female. 6, T. pretiosum, female. Scale =
10 p.

796 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Pintureau and Babault, 1981). However, a technique that uses only one wasp
per sample well is still highly desirable. This technique would not only
facilitate the direct analysis of allele frequency of each population, but also
would enable us to use isozyme analysis in hybridization experiments (Hung
and Vinson, 1977). Nevertheless, unless chromosome banding pattern turns
out to be species specific, isozyme analysis will still be the most useful tool
in Trichogramma biosystematics.

ACKNOWLEDGMENTS

I am grateful to Dean W. P. Leon Guerrero, Associate Dean R. Muniap-
pan, J. L. Demeterio, Marilou T. Baccay, Sandra L. Dela Garza, and Ros-
alee Kikuchi for helping me to carry out this study in Guam. I thank P. P.
Burbutis, W. J. Lewis, and R. K. Morrison for sending me live cultures and
E. E. Grissell and D. R. Smith, Systematic Entomology Laboratory, USDA,
for reviewing the manuscript. This study was supported in part by USDA
Regional Project W-84.

LITERATURE CITED

Crozier, R. H. 1977. Evolutionary genetics of the Hymenoptera. Annu. Rev. Entomol. 22:
263-288.

Fukada, H. and M. Takemura. 1943. Genetical studies of Trichogramma. (In Japanese).
Jpn. J. Genet. 19: 275-281.

Harland, S. C. and O. M. Atteck. 1933. Breeding experiments with biological races of Trich-
ogramma minutum. Z. Indukt. Abstammungs. Vererbungsl. 64: 54-76.

Hung, A. C. F., H. T. Imai, and M. Kubota. 1972. The chromosomes of nine ant species
(Hymenoptera: Formicidae) from Taiwan, Republic of China. Ann. Entomol. Soc. Am.
65: 1023-1025.

Hung, A. C. F. and S. B. Vinson. 1977. Interspecific hybridization and caste specificity of
protein in fire ant. Science (Wash., D.C.) 196: 1458-1460.

Jardak, T., B. Pintureau, and J. Voegele. 1979. Mise en evidence d’une nouvelle espece de
Trichogramma (Hym., Trichogrammatidae). Phenomene d’intersexualite; etude enzy-
matique. Ann. Soc. Entomol. Fr. (N.S.) 15: 635-642.

Laemmli, U. K. 1970. Cleaveage of structural proteins during the assembly of the head of
bacteriophage T4. Nature 227: 680-685.

Morrison, R. K. 1970. A simple cage for maintaining parasites. Ann. Entomol. Soc. Am.
63: 625-626.

Nagarkatti, S. and H. Nagaraja. 1977. Biosystematics of Trichogramma and Trichogram- |
matoidea species. Annu. Rev. Entomol. 22: 157-176. |

Pintureau, B. and M. Babault. 1981. Caractérisation enzymatique de Trichogramma eva- |
nescens et de T. maidis (Hym.:Trichogrammatidae): Etude des hybrides. Entomophaga
26: 11-22.

Pintureau, B., R. Goujet, D. Martouret, and J. Voegele. 1980. Etude des esterases chez |
Trichogramma embryophagum Hartig (Hym.:Trichogrammatidae). Choix d’une souche |
pour lutter contre Laspeyresia pomonella L. (Lep.: Tortricidae) dans la région pari-
sienne. Bull. Soc. Entomol. Mulhouse, pp. 17-24.

Voegele, J. and J. B. Berge. 1976. Les Trichogrammes (Insectes Hymenop. Chalcidiens, ,
Trichogrammatidae), caractéristiques isoesterasiques de deux espeéces:Trichogramma

evanescens Westw. et T. achaeae Nagaraja et Nagarakatti. C. R. Acad. Sci. Paris 283: }
1501-1503.

|

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 797-821

SEXUAL BEHAVIOR, BIONOMICS, AND FIRST-INSTAR LARVAE
OF THE LAUTA AND DIVERSICORNIS GROUPS OF EPICAUTA
(COLEOPTERA: MELOIDAE)

RICHARD B. SELANDER

Departments of Genetics and Development and of Entomology, Univer-
sity of Illinois, Urbana, Illinois 61801.

Abstract.—Epicauta lauta (Horn) is removed from the Diversicornis Group
and placed in a group by itself. Male courtship behavior in E. lauta and E.
polingi Werner is described in detail. In both species the orientation phase
is short and the mounted phase long. The mounted phase involves distinct
riding and displaying subphases. On the basis of limited observation, Epi-
cauta arizonica Werner is not distinguishable behaviorally from E. polingi.

Data on adult activity, adult feeding behavior, and oviposition are sum-
marized for the Lauta and Diversicornis groups. Adults of Epicauta lauta
feed on Larrea; those of the Diversicornis Group favor Leguminosae (es-
pecially Prosopis). Hatching time of eggs and number of eggs per mass are
recorded for E. lauta, E. polingi, and E. arizonica. A rearing of E. polingi
on the eggs of the acridid Melanoplus differentialis (Thomas) is described.

The Lauta, Diversicornis, and Fabricii groups are characterized and com-
pared on the basis of anatomical characters of the first-instar larva. The
larva of E. lauta possesses several characteristics not found in other species
of the subgenus Macrobasis and one characteristic (only three minor setae
on each side of the head dorsally) that is unique in Epicauta. Larvae of E.
polingi and E. arizonica share many characteristics with E. fabricii (Le-
Conte) and E. murina (LeConte), of the Fabricii Group.

This paper treats the sexual behavior, bionomics, and larval anatomy of
species of blister beetles currently assigned to the Diversicornis Group of
the subgenus Macrobasis LeConte of the genus Epicauta Dejean. This group
was established by Werner (1949) on the basis of anatomical characters of
the adult male. Twelve species were included, all strictly North American
in distribution except Epicauta flagellaria (Erichson), which occurs in Pan-
ama and northern South America. Epicauta luteola Dillon (1952), described
subsequently from Texas, apparently should be included also. Species treat-
ed in detail herein are Epicauta lauta (Horn), E. polingi Werner, and E.
arizonica Werner. In the adult stage, as noted by Werner et al. (1966), E.

798 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

lauta differs from other species of the group in exhibiting relatively slight
sexual dimorphism of the antenna. Associated with this trait is a distinctive,
elaborate pattern of male courtship display, described below. Further, in
anatomical characters of the first larval instar, E. /auta is much less similar
to E. arizonica and E. polingi than these species are to E. fabricii (LeConte)
and E. murina (LeConte), of the Fabricii Group of Macrobasis. Taken to-
gether, the evidence clearly indicates that E. /auta must be removed from
the Diversicornis Group and assigned to a separate group, which I will
hereafter refer to as the Lauta Group.

For comparative purposes, descriptions of the larvae of species of the
Lauta and Diversicornis groups given in this paper are supplemented with
details regarding anatomical characteristics of Epicauta fabricii and E. mu-
rina not included in MacSwain’s (1956) larval descriptions of these species.

MATERIALS AND METHODS

Studies of sexual behavior were carried out largely at field sites under
relatively primitive conditions. Behavioral data were obtained by observa-
tion and photography of field-caught adults confined to cages in groups of
12-25. Some observations were made in sunlight, others under a bank of
fluorescent lamps. Temperature during observation and filming periods ranged
from 27° to 31°C. Data were taken in the form of tape recorded descriptions
of activities, still photographs, and, in the case of E. lauta and E. polingi,
200 and 800 feet, respectively, of 16 mm motion picture film (24 fps). In
describing the courtship behavior of E. lauta I occasionally record mean
duration or rate of movement for an activity. These values, derived from
data obtained by counting motion picture frames, are generally based on
samples of no more than five bouts of behavior and, accordingly, have little
or no value for comparative purposes. They are presented simply to give the
reader a rough idea of the tempo of the behavior.

Epicauta lauta was studied behaviorally at two localities (inclusive dates
of study in parentheses): Van Horn, Culberson County, Texas (5-VIII-59),
and Balmorhea, Texas (30-VIII/1-VIII-63). All material was taken at light.
At the time of the studies no food plant had been recorded for the species.
Adults from Van Horn were confined without food; those from Balmorhea
were given lettuce, which they ate.

Epicauta polingi was studied at Apodaca, Nuevo Leon, Mexico (3/21-
VII-60); Quemado, Maverick County, Texas (27-VIII/2-IX-61); Balmor-
hea, Reeves County, Texas (26/29-VII-63); and Ft. Davis, Jeff Davis Coun-
ty, Texas (8-VII-63). The bulk of the adult material of this species was
collected at lights. Captive adults were given flowers of Prosopis juliflora
and Solanum elaeagnifolium as food.

The behavior of Epicauta arizonica was studied only superficially on the

VOLUME 84, NUMBER 4 799

basis of a sample of adults from Culiacan, Sinaloa, Mexico (26/27-IX-72).
Lettuce was provided as food.

In describing postembryonic ontogeny, I have adopted Selander and
Mathieu’s (1964) use of the symbols T, FG, C, SG, P, and A for the four
phases of the larval stage (triungulin, first grub, coarctate, second grub), the
pupal stage, and the adult stage, respectively, with numerical subscripts to
indicate instars. The technique used in rearing Epicauta polingi is described
in the account of the rearing.

First-instar larvae described in this work were progeny of adults used in
the behavioral studies or additional field-caught material. The geographic
source and number of larvae studied are specified in the section on larval
anatomy.

Egg masses, which were generally laid on plant material or cage floors,
were placed in individual cotton-stoppered 3-dram vials and incubated at
100% RH in darkness. Larvae were killed in 70% ethyl alcohol (in water) a
few days after eclosion. Those selected for detailed study were dehydrated
through an alcohol series, cleared in oil of wintergreen and toluene, and
mounted on slides in Harleco Synthetic Resin.

The following conventions were adopted in describing larvae: Length of
the head is the distance from the posterior margin of the head capsule to
the clypeolabral suture. Coloration is that in preserved, unmounted speci-
mens. Setae in a transverse row, whether on the labrum, frontal margin of
the head, or abdominal terga and sterna, are numbered consecutively on
each side of the body, beginning with the seta nearest the midline and pro-
ceeding laterad.

In the text, values accompanying means in parentheses are estimated
standard errors.

SEXUAL BEHAVIOR

Following Selander and Mathieu (1969), I divide the courtship stage of
sexual behavior in Macrobasis into (1) an orientation phase, in which the
male may (depending on the species) touch or press the antennae and max-
illary palpi on the female but does not contact her with the body or legs,
and (2) a mounted phase, in which he holds her with at least the fore- and
hindlegs. The mounted phase may be divided into (2a) an active stimulus
or ‘‘display’’ subphase, in which the male performs stereotyped acts evi-
dently intended to stimulate the female, (2b) a genital presentation sub-
phase, in which he attempts to insert his genitalia in her genital opening,
and (2c) a riding subphase, in which he neither displays nor presents the
genitalia. The second subphase, which constitutes the entire mounted phase
in the Albida Group, was termed ‘‘abortive mounting’’ by Selander and
Mathieu (1969).

800 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

The mounted position of the male is basically the same in all species of
Macrobasis (Selander and Mathieu, 1969; Selander, unpublished
data). Depending on the size of the male, his head is at the level of the
female’s or somewhat behind. The forelegs are brought to the sides of the
female’s prothorax, where principal contact is made by the first tarsal seg-
ment. In the Albida and Fabricii groups the male’s foretarsi are merely
pressed against the sides of the prothorax (foreleg clasp); in the Lauta and
Diversicornis groups the first segment of the foretarsus is inserted under the
hind margin of the female’s pronotum (foreleg Jock). The hindlegs grasp the
female on the lateral margins of the elytra or on the sides of the abdomen
just below the elytral margin. Here contact is made by the posterior side of
the apex of the hindtibia, which is provided with a hindtibial comb appar-
ently instrumental in maintaining the grasp (Selander and Mathieu, 1969).
In the Lauta and Diversicornis groups the midlegs are extended to the sides
as soon as the fore- and hindlegs are positioned properly and are not used
to grasp the female at all.

In all groups of Macrobasis studied behaviorally a fixed series of activities
occurs immediately before copulation to which the name precopulatory se-
quence has been applied (Selander and Mathieu, 1969). The sequence is
initiated by the female’s tipping sharply forward, opening the last segment
of the abdomen, and turning it slightly upward. In direct response, the male,
if not already mounted, mounts the female immediately, extrudes the gen-
italia downward and forward and inserts them in her genital opening. The
female remains still until the genitalia are coupled, at which point she lowers
the body. As a prelude to dismounting, the male usually raises the midlegs
or brings them to the substrate at the sides of the female. Finally, he releases
the foreleg grasp and either walks off her or falls over backward and then
rights himself. During copulation the sexes face in opposite directions, con-
nected only by the genital coupling.

Epicauta lauta (Horn)

Courtship.—The orientation phase is characteristically brief, lasting a few
seconds to a few minutes. The male appears to recognize a female only
when within a distance of a few centimeters. He then turns to face her and
approaches directly, with little indication of wariness and with no tendency
to hesitate or posture before her. The antennae are directed forward and
held immobile; the maxillary palpi are fully extended forward. There is no
attempt to antennate or palpate the female.

As the male mounts the female he brushes the tips of the maxillary palpi
lightly over her elytra. On reaching the dorsal position he immediately es-
tablishes the foreleg lock and hindleg grasp, raising his body well above the
female’s dorsum. The apex of the foretibia, as well as the first foretarsal

VOLUME 84, NUMBER 4 801

Fig. 1. Courtship in Epicauta lauta. a, Riding. b, Antennal embracing.

segment, may be inserted beneath the female’s pronotum. The posterior
surface of the foretibia rests lightly on the female, conforming by its cur-
vature to the shape of the humeral area of the elytra. The positioning of the
hindlegs is nearly as precise as that of the forelegs. The apex of the hindtibia
rests just on the lateral margin of the female’s elytron, near the level of the
hind margin of the third visible abdominal sternum. The hindtarsi do not
touch her.

The mounted phase of courtship is characteristically of long duration (sev-

802 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Midleg Mid- & hindleg

Genital Hindleg Palpation/ (<> Antennal
presentation ~ ~ rubbing ~ antennal lashing <~——1 embracing
! Berney
Y
Precopulatory
sequence Riding
|
1
Y
Copulation Orientation 2

Fig. 2. Sequence of male sexual behavior in Epicauta lauta. Relative frequencies of the
observed transitions are suggested by the thicknesses of the arrows. Transitions not actually
observed are represented by dashed arrows.

eral minutes to an hour or more). It is characterized by extended bouts of
display interspersed with brief bouts of riding.

In the riding subphase (Fig. la) the head is elevated, the maxillary palpi
are folded, and the antennae assume the position shown in the figure or are
directed even more strongly backward, parallel to the body. The femur of
the midleg projects horizontally to the side, with the tibia and tarsus sloping
diagonally downward; the tarsi sometimes rest on the substrate at each side
of the female. In no case that I observed were the midlegs fully extended
or the femur directed above the horizontal. During both the riding and dis-
play subphases the end of the male’s abdomen is open and the genitalia
partially extruded.

Riding seems to occur chiefly in response to sudden movement of the
female or an especially vigorous attempt to dislodge the male by brushing
over the back with the mid- or hindlegs.

The male courtship display of Epicauta lauta is unusually complex for a |
species of Macrobasis. It involves three principal activities: leg rubbing,
antennal embracing (AE), and palpation/antennal lashing (PAL). These
occur sequentially (Fig. 2), although the first two usually overlap at least |
briefly.

Leg rubbing is a compound activity consisting of hindleg rubbing (HLR)
and midleg rubbing (MLR), which are performed separately as a rule but |
on occasion occur simultaneously. In any case, the head is elevated and the |
antennae are directed backward, as in riding. In HLR, which is typically |
the initial activity in a bout of display, the hindlegs are straightened, directed |
backward, and rubbed rapidly from side to side over the female’s elytra. |
The two legs move synchronously and in phase. Frequently the tarsi curl |
over the end of the female’s abdomen, as in Fig. 1b. A bout of HLR lasts |

™ = — -_— — I
——————— ——

SS

VOLUME 84, NUMBER 4 803

about two s; at 30°C, complete leg strokes occurred at a mean rate of 6.3
per s.

MLR consists of curling the midtarsi and rapidly and rather delicately
dabbing or brushing them on the legs and sides of the female’s abdomen in
a poorly controlled, rather indiscriminate manner. Frequently the behavior
occurs in several brief bursts of activity following one another in rapid
succession. Mean duration of bouts of MLR was about 14 s: mean rate of
movement (at 30°C) was 7.4 strokes per s.

In the typical pattern of courtship, HLR alternates with repeated cycles
of AE and PAL. However, in what I interpret to be especially intense
behavior, AE alternates with simultaneous rubbing by both the mid- and
hindlegs. In this behavior the midlegs are straightened and directed back-
ward in the same manner as the hindlegs and move synchronously with
them, in opposite phase.

It is interesting that MLR may be performed as a substitute or HLR (or
vice versa). For example, in two filmed sequences in which the female is
tipped to the side and resting on one of the male’s hindlegs, there is simul-
taneous movement of the free hindleg and the opposite midleg.

AE (Fig. 1b) is a precise manipulation of the female’s antennae by the
male’s. To initiate it the male, while lowering the head, brings each antenna
forward just to the outside of the corresponding antenna of the female and,
by bending his antennae at the junction of segments I and II, crosses his
flagella in front of the female’s head, completely encircling her antennae in
a very loose embrace. If the female does not resist, each of her antennae
slides into the bend in the male’s antenna. The male then presses his an-
tennae inward and raises his head, bringing the female’s antennae straight
up and parallel in front of his head, where they are held momentarily before
release. In film timings (at 30°C) males spent a mean of about | s bringing
their antennae forward and a mean of 4 s embracing the female’s.

Frequently AE is followed immediately by HLR. However, when the
female is passive, a pattern of extended, rhythmic cycling between AE and
PAL develops. In each cycle there is a relatively extended period of PAL,
followed by a single performance of AE, during which palpation is not
interrupted but, rather, is directed to the top of the female’s head.

The palpation element of PAL is an animated, vigorous rubbing of the

_ maxillary palpi on the female’s pronotum produced by the male’s rapidly

and repeatedly nodding the head. At 30°C, mean duration of bouts of rubbing
was 1.2 s, mean rate of head movement 5.4 nods per s. Palpation is always
accompanied by antennal lashing, in which the male directs his antennae
forward and moves them rapidly from side to side over the female's. The
movement is poorly controlled and is apparently not directed at any specific
part of the female’s antennae. Occasionally, the male’s flagellum may be
thrown into a complete loop; and in one filmed sequence a loop momentarily

804 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

encloses one of the female’s antennae. There is no indication in the films
that the male attempts to grasp the female’s antennae or manipulate them
in a deliberate manner, but one cannot help but be impressed by the simi-
larity of this behavior to the much more highly developed and stereotyped
antennal whipping and curling of some species of the Albida Group.

Females did not attack males in the orientation phase of courtship and
were, in general, rather tolerant of courting males in the mounted phase.
The most common negative response of the female is to brush over the
back with the middle or hind legs. This often interrupts the male’s display
cycle but is apparently ineffectual in dislodging him. A second negative
response is to lower the head and antennae, making it difficult or impossible
for the male to perform AE. Ina third negative response the female elevates
her head and antennae as far as possible, pressing the latter against the front
of the male’s head. In both observed cases of this behavior the male lowered
the head onto the female’s pronotum and appeared to push against it with
the mandibles.

Genital presentation apparently occurs very infrequently during court-
ship. I did not record a single complete bout of this behavior. However, on
one occasion, following HLR, a male strongly extruded the genitalia, di-
rected them downward, and began lowering the end of the abdomen. It is
on this basis that I indicate in Fig. 2 a possible transition of HLR to genital
presentation.

Precopulatory sequence and copulation.—Not observed.

Epicauta polingi Werner

Courtship.—In this species, as in E. /auta, the orientation phase of court-
ship is brief, the mounted phase is generally lengthy, and there is a definite
display subphase of the mounted phase. |

The male of E. polingi seems rather wary when approaching a female;
otherwise, his behavior is essentially like that of E. /auta. A figure of an
orienting male of E. polingi is given by Selander and Mathieu (1969).

In nearly all observed bouts of courtship the male mounted from directly | '

or diagonally behind the female. As he mounts he slides the maxillary palpi

along the female’s elytra. On reaching her dorsum he quickly inserts the | ,
foretarsi under the female’s pronotum and brings the ends of the hindtibiae };

to the lateral margins of her elytra, exactly as in E. lauta.

The male in one filmed sequence, having just mounted, lashed the anten-
nae over those of the female for several seconds, at times throwing a com-
plete loop in one of the flagella and loosely encircling one of the female’s
flagella, in the fashion described for E. /auta. In another case, a male ini-
tiating a bout of courtship that led shortly to copulation performed essen-
tially this same behavior while standing in front of the female, just before

VOLUME 84, NUMBER 4 805

mounting. No comparable behavior was noted in any other bouts of court-
ship.

All but a small fraction of the male’s time in the mounted phase is devoted
to riding. In this behavior (Fig. 3a) the head is moderately elevated and the
midlegs are fully extended and directed to the sides. In low-level riding the
flagellum is directed nearly straight forward, the midfemur is nearly hori-
zontal, and the rest of the midleg droops slightly. In high-intensity riding,
performed when the female moves suddenly or brushes over her back, an-
tennal segment I is directed upward and the midlegs are elevated well above
the horizontal, with the tarsi curving upward, as the male clearly tries to
keep these appendages as far as possible from the female. The male’s gen-
italia are extruded throughout the mounted phase. The degree of extrusion
is consistently greater than in E. lauta.

Periodically, especially when the female is still, the male slowly lowers
the antennae and tries to perform antennal rubbing (AR) (Fig. 3b). In this
act antennal segment I is directed diagonally forward and the flagella curve
downward, closely paralleling each other distally. By moving antennal seg-
ment I, the male then rubs the flagella rapidly and vigorously forward and
backward several times on the top and front of the female’s head. The
antennae move synchronously and in phase. Two or more brief bouts of
AR may be performed in succession. In many females that I have examined
the clothing setae on the midline of the head are abraided, presumably as a
result of AR.

There is a second form of antennation, antennal touching (AT), observed
less frequently than AR, in which the male inclines the head to one side and
delicately touches the end of his opposite antenna on the front of the fe-
male’s head. In one case AT was followed immediately by AR; in another,
two bouts of AT occurred in succession. In three filmed bouts of the be-
havior males touched the female’s head 6, 12, and 2 times, respectively.

The only other active component of display in the mounted phase is a
delicate, casual dabbing of the maxillary palpi on the top of the female’s
head. In my observations it was performed sporadically and then only brief-
ly.

Genital presentation subphase.—Males rarely attempted to insert the gen-
italia except in the course of the precopulatory sequence. In a few instances
a male presented the genitalia as soon as he mounted, a form of behavior
suggesting the ‘‘abortive mounting”’ of the Albida Group.

Precopulatory sequence.—As a rule, tipping by the female occurs while
the male is mounted, usually as an immediate response to AR. When the
female tips, the male usually dabs the maxillary palpi lightly on her head or
pronotum. Males that were large enough to insert the genitalia without mov-
ing backward continued to perform AR during genital presentation. After

806 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

3

Fig. 3. Courtship in Epicauta polingi. a, Riding (after Selander and Mathieu, 1969; copy-
right, Board of Trustees, University of Illinois, used with permission). b, Antennal rubbing.

VOLUME 84, NUMBER 4 807

the genitalia are coupled the male lowers the midlegs to the substrate, if he
has not already done so. In two separate timings, the precopulatory se-
quence, from presentation of the genitalia until ‘“‘turnoff’’ of the male, lasted
100 and 130 s, respectively.

If a female is ready to copulate when approached by a male, she may
solicit copulation by tipping during the orientation phase of courtship. This
behavior was observed in two cases in which reared females about a month
old were exposed to males for the first time.

Copulation.—I observed several bouts of copulation in E. polingi but did
not record the duration of any.

Epicauta arizonica Werner

In my very limited observation of this species I was unable to detect any
significant difference in its courtship behavior and that of E. polingi. As in
that species, the orientation phase is brief and the mounted phase involves
long periods of riding interspersed with short bouts of display. The riding
behavior appears to be identical. The display consists of male antennal
manipulation of the female’s head at least superficially like antennal rubbing
in E. polingi.

DISCUSSION

The Lauta and Diversicornis groups both differ markedly from the Albida
Group with respect to the general pattern of male courtship behavior. In
the Albida Group, as described by Selander and Mathieu (1969), the ori-
entation phase dominates courtship temporally and incorporates stereo-
typed visual and tactile stimulatory acts by the male, whereas the mounted
phase entails only genital presentation, typically performed forceably and
briefly. An intermediate pattern is found in the Fabricii Group, where the
mounted phase incorporates a distinct riding subphase but, as in the Albida
Group, lacks a display subphase (Selander and Mathieu, unpublished data).

There is, among species of Macrobasis, a strong correlation between the
length of the mounted phase of male courtship and the strength of modifi-
cation of the male’s fore- and hindlegs. In the Lauta and Diversicornis groups,
where, as we have seen, the male spends nearly all of the courtship period
mounted on the female, the forelegs are strongly modified to facilitate the
foreleg lock (Fig. 4) and the hindtibial comb is consistently present and well
developed. In contrast, modification of the male forelegs in the Albida and
Fabricii groups consists largely of loss of one of the tibial spurs, and the
hindtibial comb is vestigial or absent in most species.

Antennal segment I is somewhat longer and thicker in the male of Epi-
cauta lauta than in the female, but the sexual dimorphism is minor compared
to that in the Diversicornis Group. The adaptive significance of the great

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

d

808

SS

25>
7 ee

2
aa

>>
Z

——=F

KC

\\

S25
a

1

sy
——_

\\

AWN

<F

SS
x

oe
AO

WSS
SSS

_Z

BF

Ze

AZ

=

Tibia and tarsus of male foreleg. a, Epicauta lauta, anterior view. b, Same, posterior

Fig. 4.
view. c, E. polingi, anterior view. d, Same, ventral view.

enlargement of the first two antennal segments of the male in E. polingi, E.
arizonica, and many other species of Macrobasis is difficult to interpret

solely in mechanical terms. Despite the lengthening of these segments, the
male’s antennae are not much longer than the female’s, so the dimorphism

VOLUME 84, NUMBER 4 809

presumably reflects more than a means of increasing the effective distance
at which they may be employed in tactile stimulus. That the modifications
are important mechanically in the curling and other contortions of the fla-
gellum seems unlikely since males of many other species of Meloidae lacking
marked sexual dimorphism of the antennae perform equal elaborate flagellar
movements in courtship. Almost certainly, the antennal dimorphism in Mac-
robasis involves visual stimulation of the female by the male. In the Albida
Group, males possessing enlarged basal antennal segments characteristically
posture in front of females, literally displaying the antennae, whereas in
males without the enlargement this behavior is much less developed or
absent (Selander and Mathieu, 1969). Moreover, in some species of Macro-
basis the enlarged antennal segments contrast strongly with the remainder
of the antenna and with the body in being yellow in color, as though designed
to attract attention.

BIONOMICS

Adult activity —Adults of Epicauta lauta and the species of the Diver-
sicornis Group are attracted to lights at night, often in huge numbers. They
are not strictly nocturnal, however, since large populations of some of the
more common species have been found feeding in the daytime. Adults of at
least one species of the Diversicornis Group migrate in large swarms. On
the morning of 21 July 1955 at Lo de Lomeda, in the mountains of Nayarit,
I found a migrating population of thousands of E. beckeri (Duges) adults
crossing the highway that connects Tepic and Mazatlan. The beetles moved
in slow, steady flight two to 15 feet above the ground, coming up in waves
a few minutes apart from a low area on one side of the highway and dropping
down into a canyon on the other. Some individuals alighted and then took
off again. None walked an appreciable distance and none was found feeding
on the vegetation nearby. I observed the migration for 75 minutes; it was
still in progress when I left the site.

In Arizona, the seasonal range of adults of Epicauta lauta is June to
November (Werner et al., 1966). In Apodaca, Nuevo Leon, Hernandez
(1960) took adults of the species in a light trap from May to November; the
percentage distribution by month of the catch (N = 7663) was: May, 2.8;
June, 13.1; July, 6.5; August, 50.7; September, 25.8; October, 1.0; and No-
vember, 0.2. Epicauta polingi has been taken from April to September, E.
arizonica from July to September, and E. liebecki Werner from May to July
(Werner, 1945; Werner et al., 1966).

Feeding behavior.—Although Epicauta lauta is one of the more com-
monly collected species of meloids in the southwestern United States, no
food plant was known before Hurd and Linsley (1975) reported finding adults
on Larrea tridentata at several localities in southern Arizona.

The Diversicornis Group resembles the Fabricii Group in apparently fa-

810 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table |. Hatching time of eggs and number of eggs per mass.
Hatching Time (Days) Eggs per Mass
N Temp. N
Species Mean SE Range Masses (°C) Mean SE Range Masses
E. lauta 18.0 2.00 16-20 2 2 355 0.0 —- I
E. arizonica 16.0 0.00 a 3 27 229 14.2 201-248 3
E. polingi 15.1 0.61 12-21 18 223 171 13.8 128-220 7

® Two masses (Balmorhea, Texas) hatched in 17 and 19 days, respectively, at 27°C. Tem-
perature was not controlled for the other masses.

voring plants of the family Leguminosae. Werner et al. (1966) reported
finding Epicauta liebecki on Prosopis several times (‘‘once in great num-
bers’’) and E. arizonica on Prosopis foliage at night. I have found E. polingi
feeding on Prosopis juliflora flowers at night in Texas and have taken an
adult here and there on Solanum elaeagnifolium in the day. Hernandez
(1960) collected 28 adults from S. elaeagnifolium in Nuevo Leon. Captive
adults used in my behavioral studies ate the flowers of Solanum elaeagni-
folium but clearly preferred those of Prosopis juliflora. 1 occasionally saw
them chewing on the leaves of Prosopis. Reared adults of E. polingi lived
40-87 days [mean 59.5 (7.77)] on a diet of Medicago sativa and Solanum
tuberosum (four cases) or Medicago alone (one case). Solanum dulcamara
was refused by these adults, and a local, spiny Solanum and a local Amar-
anthus were refused by adults of E. arizonica collected at light in Culiacan,
Sinaloa.

I have taken Epicauta diversicornis in Nayarit and Michoacan on un-
identified legumes and in Morelos on Prosopis. In addition, in Guerrero I
once found the species feeding in great abundance on the leaves of a viny
shrub (without flowers) that was identified by the botanist Faustino Miranda
as a species of Malpighiaceae.

Epicauta isthmica Werner was taken in large numbers by W. H. Young
at Cotaxtla, Veracruz, in July 1958 on Crotalaria.

Eggs.—Available data on hatching time of eggs and number of eggs per
mass are summarized in Table 1. Hatching time is comparable to that re-
corded for E. murina (Gilbertson and Horsfall, 1940) and species of the
Albida Group but much shorter than that of E. fabricii (Horsfall, 1943).
Epicauta polingi is one of several species whose eggs I have tested for
ability to produce blisters by taping eggs to my forearm. The test was pos-
itive.

Postembryonic development.—Epicauta polingi was reared on diapaus-
ing, year-old eggs of the acridid grasshopper Melanoplus differentialis
(Thomas) in 1963 by J. M. Mathieu and myself. Twenty-five 7-day-old T
larvae from an egg mass laid by a female from Balmorhea, Texas, were

VOLUME 84, NUMBER 4 811

Table 2. Cumulative time (days) for larvae of Epicauta polingi to reach developmental
periods.

Period Mean SE N
FG, 4.5 38 228
FG, Set 43 19
FG, 7.6 68 14
FG, 10.2 .66 14
FGspr° 17.4 70 i
FG, 14.0 — I
FGepe? 21.0 = |
Cz 21.0 — I
C; 26.0 — !
PE 26.9 1.22 9
A; 36.6 .87 5

4 Data for one larva, greatly delayed in reaching FG, (14 days), omitted entirely.
> Postfeeding period in FG (after transfer from food to sand).

confined individually with Melanoplus eggs in glass tubes and placed at
27°C, 100% RH, in darkness in a homemade incubator. Twenty-three larvae
(92.0%) fed and reached FG,. As the rearing progressed, 11 larvae and one
pupa were killed as specimens; one larva died after several months in the
C phase. FG; or FG, larvae were given food until they stopped eating and
then transferred to 3-dram vials packed with moist sand.

Developmental timing, summarized in Table 2, is well within the norm
for rearings of species of Epicauta. Eleven larvae had four instars in the
FG phase; one had five. As reported previously (Selander and Weddle,
1969), nine (75.0%) of 12 larvae followed the pattern T-FG-P-A; the rest
followed the pattern T-FG-C. One of the two C, larvae and the single C,
larva were chilled at 5°C from day 86 to day 255 of the rearing and then
returned to 27°C. The former died 32 days after chilling. The latter ecdysed
in 61 days to form a SG, that was structurally abnormal in retaining some
coarctate characteristics.

The incidence of the pattern T-FG-P-A is unusually high, especially at
27°C. Taken together with the lengthy seasonal range of adult activity, it
almost certainly indicates that E. polingi has more than one generation a
year.

THE FIRST-INSTAR LARVA

MacSwain (1956) distinguished the first-instar larvae of Epicauta fabricii
and £. murina from those of other species of Epicauta as follows: (1) ab-
dominal sternite VII well developed, complete; maxillary palpal segment III
of a unique form (short, with sensory area subcircular, dorsal, extending
less than % length of segment, set with numerous long papillae); and (3)

812 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

labial palpal segment II with two dorsobasal setae, each extending beyond
the apex of the segment. Epicauta arizonica and E. polingi agree with this
diagnosis except that sternite VII is divided on the midline of the body.
However, the significant point, it seems to me, is that the sternite is nearly
as fully developed in these species as in those of the Fabricii Group, whereas
no other species of the genus is comparable in this respect. Another note-
worthy similarity between larvae of the Diversicornis and Fabricii groups
is the presence of two dorsobasal setae on segment III of the maxillary
palpus. All other species of Macrobasis have one. As recorded in the key
and descriptions that follow, the two groups closely resemble each other in
several other anatomical characters, including chaetotaxy of the frontal area
of the head, labrum, and antennae and the form of the labrum, mandibles,
gula, and cardo.

Epicauta lauta shares none of the special similarities linking the Diver-
sicornis and Fabricii groups. Abdominal sternum VII is no more strongly
or extensively sclerotized than usual for the genus. The maxillary cardo is
better developed. Maxillary palpal segment III is slender and elongate; its
sensory area is positioned dorsolaterally and is sparsely set with short,
peglike papillae; and there is only a single dorsobasal seta. Labial palpal
segment II has only a single dorsobasal seta also. As a matter of fact, the
larva of E. lauta is one of the most distinctive types yet described in Ma-
crobasis. It differs from all other larvae of the subgenus in having the labrum
(and clypeus) much enlarged, the mandibles greatly elongated and bearing
an unusually large number of teeth, the lanceolate setae of femur I more
numerous, and the reticulations of the abdominal sclerites devoid of pos-
terior projections. In addition, it is the only species of the genus Epicauta
possessing three, rather than four, basal setae on each side of the dorsum
of the head.

The form of the mandibles and maxillary palpi in E. /auta suggests the |
condition in E. atrata (Fabricius), a species of the nominate subgenus of |
Epicauta for which I recently obtained evidence indicating larval predation |
on the eggs of blister beetles rather than those of grasshoppers, the usual
prey of larvae of Epicauta (Selander, 1981). The modifications of the mouth-
parts are not as extreme in E. /auta. Further, spiniform cuticular evagina- |
tions are relatively well developed, whereas in E. atrata and in other species _
that I believe share its larval prey-type, the evaginations are poorly devel-
oped or absent. Still, the mouthparts of E. /auta differ sufficiently from the |
norm of the genus as to suggest a special study of larval feeding behavior |
of the species at the earliest opportunity.

Formal diagnoses of the Lauta, Diversicornis, and Fabricii groups in the |
larval stage are given in the following key, which further serves to empha- |
size the distinctiveness of the Lauta Group with respect to the others.

VOLUME 84, NUMBER 4 813

DIAGNOSTIC KEY TO THE LAUTA, DIVERSICORNIS, AND
FABRICII GROUPS BASED ON THE FIRST-INSTAR LARVA

1. Median pair of setae on anterior margin of frontal area of head set
close to margin; gula flared anteriorly, well separated from labium;
stemma 2 as large as mesothoracic spiracle; apical setae of anten-
nal segments II and III short; labrum more than 7/5 as wide as head;
mandible slender, elongate, with 15-18 teeth, these small, truncate;
maxilla with cardo large; maxillary palpal segment III 2 as long as
wide, with a single dorsobasal seta and with sensory area dorsolat-
eral, bearing short, peglike papillae; labial palpal segment II with a
single dorsobasal seta; reticulations of abdominal sclerites lacking
posterior projections; sternum VII weakly sclerotized, none of the
posterior marginal setae sharing a common sclerite; caudal seta
14x as long as segments VH-IX combined; femur I with 8—9 (rarely
7) lanceolate setae; tarsungulus distinctly curved ....... Lauta Group

— Median pair of setae on anterior margin of frontal area of head set
well back from margin; gula gradually wider anteriorly, touching
labium; apical setae of antennal segments II and HI long; labrum 3
as wide as head; mandible robust, with 13 or fewer teeth; cardo
vestigial; maxillary palpal segment III at most 17/s x as long as wide,
with two dorsobasal setae and with sensory area dorsal, bearing
long, spiniform papillae; labial palpal segment II with 2 dorsobasal
setae; reticulations of at least abdominal tergites and pleurites with
distinct posterior spiniform projections; sternum VII strongly scler-
otized, with a distinct sternite (divided or not); caudal seta at most
12/sx as long as segments VII-IX combined; femur I with 7 (rarely
6) lanceolate setae; tarsungulus weakly curved .................... 4

2. Gular setae not attaining anterior margin of gula; stemma 12x as
large as mesothoracic spiracle; apical setae of antennal segment II
reaching or surpassing apex of III; mandible with 5-12 teeth, these
truncate or not; metanotum with line of dehiscence complete; retic-
ulations of abdominal sternites with distinct posterior projections;
seta | of posterior row of tergite V at most %4 as long as tergite; seta
1 of median transverse row of tergite V '/s as long as tergite; sternum
VII with sternite divided on midline; caudal seta at least 1¥%3x as
long as segments VII-IX combined ............. Diversicornis Group

— Gular setae surpassing anterior margin of gula; stemma no larger
than mesothoracic spiracle; apical setae of antennal segment II not
attaining apex of III; mandible with 11-13 teeth, these truncate;
metanotum with line of dehiscence incomplete; reticulations of ab-
dominal sternites lacking posterior projections; seta | of posterior
marginal row of tergite V nearly as long as tergite; seta | of median

814 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

transverse row of tergite V % length of tergite; sternum VII with a
complete, undivided sternite; caudal seta 1'/10 as long as segments
VH-IXicombined 44:crcise BE See ee Fabricii Group

Description of the Species

Epicauta lauta (Horn)
Figs. 5, 7a, d

Color of head, pro- and mesonotum, and abdominal tergites I-V or I-VI
light brown; metanotum and tergites VI-IX or VII-IX dark brown. Body
surface weakly reticulate; reticulations without posterior projections.

Head 34—*/s as long as pro- and mesonotum combined, widest near middle,
l'/10X as wide as long, strongly constricted behind middle, basal width 7/10
maximum width; dorsal surface with 18 major and 3 minor setae on each side;
stem of epicranial suture 2/s as long as head; distance from bifurcation of
suture to level of first pair of setae on frontal area no more than 3/s length
of stem. Gula flared, much widened anteriorly, more than 7x as wide at
apex as at base, well separated from labium; length 4/s maximum width of
head; distance from base to gular setae 4/s—?/10 length of gula; setae not
reaching anterior margin. Stemma large. Antenna elongate; segment II 2!/2—
27/10 as long as III, anteroventral apical seta minute, much shorter than
other two, which reach about middle of III, sensory organ at least as long
as III; segment III with apical setae about 2 as long as segment, terminal
seta 1!/10—17/10< as long as II. Labrum with 6 setae in median row, seta 3 of
row well separated from lateral margin, seta | much shorter than rest; lateral
margin with 3 short setae. Mandible 3/s—*/5 as long as head, '/3 to nearly 2/s
as wide as long, bearing 15—18 teeth (14-16 visible in outline); distal seta
longer than basal one. Maxilla with cardo bearing a long, conspicuous seta;
palpal segment III with sensory area extending */s length of segment, not
extending onto ventral surface at apex, papillae sparse, length of 2-seg-
mented sensory appendix 7/10 width of segment II of labial palpus. Labium
with premental plate bearing 4 setae; palpal segment II more than 2x as
long as wide, dorsobasal seta extending beyond apex of segment, 2-seg-
mented sensory appendix prominent, about 3/s as long as that of maxillary
palpus. Thorax with line of dehiscence well developed on all segments.
Abdomen with 10 setae in posterior marginal row on tergites I-VIII; setae
not divided apically; spiniform evaginations at bases of posterior marginal
setae moderately developed on tergites I-VI, vestigial on VII; tergite V 27/10—
3x as wide as long, seta | of posterior marginal row 7/10—3/4 as long as
tergum, ratio of lengths of setae 1, 2, and 3 of posterior marginal row 1:1.2-
1.3:0.7-0.8, seta | of median transverse row thin, %4 as long as tergite,
pleurites ventral, that of V °/10 as wide as long, spiracle near center of dorsal
¥4; spiracle I % or less as large as mesothoracic spiracle, at least 144X as
large as HII-VIII, which are subequal in size; sterna I-VI weakly sclerotized,

VOLUME 84, NUMBER 4

7) Mi
SS ORS o

o
"Mj

Zs
—/
Ly

Fig. 5. Head of first-instar larva of Epicauta lauta. a, Dorsal view. b, Ventral view.

815

816 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

VIII-IX strongly so. Leg I with lanceolate setae on femur heavy; leg III
with femur about 12x as long as coxa, tibia 1’2x as long as femur, tar-
sungulus 3/19 as long as tibia, longer seta 7/10 as long as tarsungulus and
about |'/sx as long as shorter seta. Length of body 2.7 mm. Length of
caudal seta 0.8 mm.

Material studied.—18 larvae, adults from Van Horn, Culberson County,
Texas, 6-VIII-59, R. B. Selander and J. C. Schaffner; 27 larvae, adults from
highway 40, 29 mi. SW Ciudad Lerdo, Durango, Mexico, 31-VII-60, R. B.
Selander and J. M. Mathieu.

Remarks.—This species is distinguished at a glance from all other species
of Macrobasis by the form of the mandible and maxillary palpus.

There are typically eight lanceolate setae on femur I. About a third of the
specimens examined have eight on one femur and nine on the other; one
specimen has seven and eight.

Epicauta arizonica Werner
Figs. 6, 7b, e

Color of head and pronotum dark brown; meso- and metanotum and ab-
dominal terga light brown. Body surface strongly reticulate; reticulations of |
abdominal sclerites, including sternites, with distinct spiniform posterior —
projections.

Head subequal in length to pro- and mesonotum combined, widest near
middle, 1'/10x as wide as long, strongly constricted behind middle, basal
width 7/19—3/4 maximum width; dorsal surface with 18 major and 4 minor
setae on each side; stem of epicranial suture 2/s as long as head; distance
from bifurcation of suture to level of first pair of setae on frontal area no
more than 2/s length of stem. Gula 4x as wide at apex as at base, projecting
anteriad; length 4/; maximum width of head; distance from base to gular |
setae 4/s length of gula. Stemma small. Antenna moderately long; segment
II less than 22x as long as III, anteroventral apical seta much shorter than
others, longer than III and reaching its apex, other 2 setae surpassing apex |
of III, sensory organ longer than III; segment III with apical setae much |
longer than segment, terminal seta 1?/3—-14/5x as long as II. Labrum with 6
setae in median row, seta 3 of row near lateral margin, seta | about as long |
as rest; lateral margin with I—2 short setae. Mandible 3/s as long as head, |
¥Y2 as wide as long, bearing 5-6 teeth (4 visible in outline); teeth large, |
rounded; distal and basal setae subequal in length. Maxilla with cardo small,
very thin, bearing a short seta; palpal segment II short, barrel-shaped, at
most 17/5 as long as wide, bearing 2 dorsobasal setae, sensory area extending |
*/s length of segment, not extending onto ventral surface at apex, papillae
dense, length of 2-segmented sensory appendix nearly equal to that of seg-
ment II of labial palpus. Labium with premental plate bearing only 2 setae;
palpal segment II nearly 2 as long as wide, dorsobasal setae extending |
beyond apex of segment, 2-segmented sensory appendix prominent, nearly |

817

VOLUME 84, NUMBER 4

I Va

Y

Fig. 6. Head of first-instar larva of Epicauta arizonica. a, Dorsal view. b, Ventral view.

818 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

4/5 as long as that of maxillary palpus. Thorax with line of dehiscence well
developed on all segments. Abdomen with 10 setae in posterior marginal
row on tergites I-VIII; setae not divided apically; spiniform evaginations at
bases of posterior marginal setae moderately developed on tergites I-VI,
vestigial on VII; tergite V 3!/s—33/sx as wide as long, seta | of posterior
marginal row 7/10-3/4 as long as tergite, ratio of lengths of setae 1, 2, and 3
of posterior marginal row 1:1.1:0.8-1.1, seta | of median transverse row
thin; pleurites ventral, that of V °/10 as wide as long, spiracle near center of
dorsal '/3; spiracle I 7/10—4/s as large as mesothoracic spiracle, at least 11/5 x
as large as II-VIII, which are subequal in size; sterna I-VI weakly sclero-
tized, VII moderately so, VII-IX strongly so, VII with sternite divided on
midline into 2 plates each of which contains posterior marginal setae | and
2: caudal seta 13/10-17/s x as long as segments VII-IX combined. Leg I with
lanceolate setae on femur heavy; leg III with femur about 17/10 as long as
coxa, tibia 1'/4-13/10 x as long as femur, tarsungulus weakly curved, about '/s
as long as tibia, longer seta about 7/10 as long as tarsungulus and about 17/s
length of shorter seta. Length of body 2.0 mm. Length of caudal seta 0.9
mm.

Material studied.—23 larvae, adults from Culiacan, Sinaloa, Mexico, 26/
27-IX-72, R. B. Selander and A. Berrios-Ortiz (Notes 601-72 RBS).

Remarks.—One specimen has seven lanceolate setae on femur I on one
side of the body and six on the other.

Epicauta polingi Werner
Fig. 7c

Color of head, pro- and metanotum, and (usually) abdominal tergites VU
IX brown; mesonotum and tergites I-VI light brown.

Mandible bearing 10-12 teeth (7-11 visible in outline); teeth medium in |
size, truncate.

Material studied.—15 larvae, adults from Balmorhea, Reeves County, |
Texas, 26-VII-63, R. B. Selander and J. M. Mathieu (Notes 32-63 RBS); 15
larvae, adults from Ft. Davis, Jeff Davis County, Texas, 8-VIII-63, R. B.
Selander and J. M. Mathieu (Notes 22-63 RBS); 8 larvae, adults from Que-
mado, Maverick County, Texas, 27-VIII-61, R. B. and J. M. Selander (Notes |
32-63 RBS); 14 larvae, adults from Quemado, Texas, 29-VIII-61, R. B. and >
J. M. Selander (Notes 35-61 RBS); 33 larvae, adults from Apodaca, Nuevo |
Leon, Mexico, 3/28-VII-60, R. B. Selander and J. M. Mathieu; 10 larvae, |
adults from Ciudad Victoria, Tamaulipas, Mexico, 9-VIII-62, R. B. Selander
and R. H. Storch (Notes 13-62 RBS).

Remarks.—The only differences between this species and Epicauta ari-
zonica of use in identifying larval specimens are those of coloration and the
form of the mandible, as described above.

VOLUME 84, NUMBER 4 819

7

Fig. 7. a-c, Right mandible of first-intar larva, ventral view. a, Epicauta lauta. b, E. ari-
zonica. c, E. polingi. d—e, Abdominal segments VII-IX of first-instar larva, ventral view. d,
Epicauta lauta. e, E. arizonica.

820 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Despite the extensive geographic range from which samples of larvae of
E. polingi have been obtained, I have been unable to detect significant
geographic variation in either color or structural characters. All specimens
examined have seven lanceolate setae on femur I.

Epicauta fabricii (LeConte)

Reference.—MacSwain. 1956: 53, pls. 8-9.

Color light brown; pronotum sometimes suffused with brown. Reticula-
tions of abdominal tergites and pleurites with distinct spiniform projections;
these not quite so strongly developed as in E. arizonica or E. polingi, not
evident on sternites.

Head with basal width 4/s—?/10 maximum width. Length of gula at least !/2
maximum width of head. Antenna with anteroventral seta of segment II
much shorter than others, no longer than III, not reaching its middle, other
2 apical setae surpassing middle of III, not attaining its apex. Labrum 3 as
wide as head, with seta | of median row distinctly shorter than setae 2 and
3. Mandible bearing 11-13 teeth (9-10 visible in outline); teeth medium in
size; distal seta longer than basal one. Metanotum with line of dehiscence
incomplete, not attaining margins. Abdomen with spiniform evaginations at
bases of posterior marginal setae moderately developed on tergites I-IV,
smaller on V, vestigial on VI-VII; tergite V with posterior marginal setae
1, 2, and 3 subequal in length, 4/s—?/i0 length of tergite, seta 1 of median
transverse row thin; sterna I-VI weakly sclerotized, VII-IX strongly so. |
Length of caudal seta 0.4 mm.

Material studied.—10 larvae, adults from West Jordan, Salt Lake County,
Utah, 11-VII-49, R. B. Selander; 15 larvae, adults from Urbana, Champaign |
County, Illinois, 31-V-63, R. B. Selander and J. M. Mathieu (Notes 4-63
M); 19 larvae, adults from Urbana, Illinois, 4-VII-61, R. B. Selander and J. |
K. Bouseman (Notes 4—61 RBS); 51 larvae, adults from Arkansas, received
from W. R. Horsfall. |

Remarks.—The above description is limited to characters in which E.
fabricii differs significantly from E. arizonica and E. polingi. A more com- |
prehensive description is given by MacSwain (1956). I have found no ap-

preciable differences among samples of E. fabricii from Utah, Arkansas, and §}-

Illinois.

Epicauta murina (LeConte)

Reference.—MacSwain, 1956: 54, pl. 9.

Remarks.—As far as I can determine, from MacSwain’s (1956) description
and a single larval specimen that I have examined, E. murina differs from
E. fabricii only in having a somewhat less robust mandible.

VOLUME 84, NUMBER 4 821

ACKNOWLEDGMENTS

For their help in collecting adults and obtaining larvae I am indebted to
A. Berrios-Ortiz, J. K. Bouseman, J. C. Schaffner, J. M. Selander, and, in
particular, J. M. Mathieu. Dr. Mathieu also took an active part in the rearing
of Epicauta polingi. A special note of appreciation is extended to W. R.
Horsfall for a gift of larvae of Epicauta fabricii and other species of Meloi-
dae studied by him in Arkansas. The drawings of adult behavior and male
forelegs were done by Alice Prickett, who also inked the drawings of larval
anatomy. Field work was supported in part by grants from the National
Science Foundation.

LITERATURE CITED

Dillon, L. S. 1952. The Meloidae (Coleoptera) of Texas. Am. Midl. Nat. 48: 330-420.

Gilbertson, G. I. and W. R. Horsfall. 1940. Blister beetles and their control. S.D. State Coll.
Agric. Exp. Stn. Bull. 340, 23 pp.

Hernandez, J. D. 1960. Estudio taxonoémico y datos biologicos de la familia Meloidae (Co-
leoptera) en los alrededores de Monterrey, N.L. Unpubl. thesis, Instituto Tecnologico
y de Estudios Superiores de Monterrey, Escuela de Agricultura, 163 pp.

Horsfall, W. R. 1943. Biology and control of common blister beetles in Arkansas. Univ.
Arkansas Agric. Exp. Stn. Bull. 436, 55 pp.

Hurd, P. D., Jr. and E. G. Linsley. 1975. Some insects other than bees associated with
Larrea tridentata in the southwestern United States. Proc. Entomol. Soc. Wash. 77:
100-120.

MacSwain, J. W. 1956. A classification of the first instar larvae of the Meloidae (Coleop-
tera). Univ. Calif. Publ. Entomol. 12, 182 pp.

Selander, R. B. 1981. Evidence for a third type of larval prey in blister beetles (Coleoptera,
Meloidae). J. Kans. Entomol. Soc. 54: 755-783.

Selander, R. B. and J. M. Mathieu. 1964. The ontogeny of blister beetles (Coleoptera, Me-

loidae). I. A study of three species of the genus Pyrota. Ann. Entomol. Soc. Am. 57:

711-732.

1969. Ecology, behavior, and adult anatomy of the Albida Group of the genus Epi-
cauta (Coleoptera, Meloidae). Ill. Biol. Monogr. 42, 168 pp.

Selander, R. B. and R. C. Weddle. 1969. The ontogeny of blister beetles (Coleoptera, Me-
loidae). Il. The effects of age of triungulin larvae at feeding and temperature on devel-
opment in Epicauta segmenta. Ann. Entomol. Soc. Am. 62: 27-39.

Werner, F. G. 1945. A revision of the genus Epicauta in America north of Mexico (Coleop-
tera, Meloidae). Bull. Mus. Comp. Zool. 95: 421-517, 7 pls.

——.. 1949. Epicauta diversicornis and its allies in the Neotropical Region. Psyche (Camb.,
Mass.) 56: 74-80.

Werner, F. G., W. R. Enns, and F. H. Parker. 1966. The Meloidae of Arizona. Univ. Ariz.
Agric. Exp. Stn. Tech. Bull. 175, 96 pp.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 822-827

NEW SPECIES AND NEW COLLECTION RECORDS OF
CERATOPOGONIDAE (DIPTERA) FROM SRI LANKA

FRANCIS E. GILES AND WILLIS W. WIRTH

(FEG) Biology Department, Loyola College, Baltimore, Maryland 21210;
(WWW) Systematic Entomology Laboratory, IIBIII, Agricultural Research
Service, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Two new species of Ceratopogonidae from Sri Lanka are de-
scribed and illustrated: Parabezzia orientalis, the first of the genus to be |
found in the Orient, and Atrichopogon schizonyx. New collection records
from Sri Lanka are presented for seven species of Alluaudomyia.

This paper is one in a series reporting on the biting midges of the family |
Ceratopogonidae collected by members of the Smithsonian Ceylonese Insect |
Project. We thank Karl V. Krombein, director of the project, for the op- |
portunity to study this material. We are grateful to Molly K. Ryan for mak-
ing the illustrations.

For explanation of methods of measurement and of ratios see Giles et al. |
(1981). The first values presented are those of the holotype followed by the
range of variation of the paratypes in parentheses.

Genus Alluadomyia Kieffer

The genus Alluaudomyia is a cosmopolitan group of approximately 114 §
species. A revision of the Oriental species with a key was published by §
Wirth and Delfinado (1964). Delfinado and Hardy (1973) listed three species jj
from the island, marginalis Wirth and Delfinado, spinosipes Tokunaga, and 9
xanthocoma (Kieffer). Four additional species are represented in the Cey- |
lonese Project collections.

Alluaudomyia bifurcata Wirth and Delfinado
New Record.—SRI LANKA: Col. Dist., Beyagama, sea level, 20-
2evini. 19737 Ga Ekis. 1 ¢-
Alluaudomyia formosana Okada

New Records.—SRI LANKA: Keg. Dist., Kitulgala Resthouse, 3-@j,
5.11.1979, UVL trap, K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, Ss. |

VOLUME 84, NUMBER 4 823

Siriwardane, and T. Gunawardane, 109; Rat. Dist., Ratnapura, 1—3.11.1979,
UVL trap, K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, S. Siriwar-
dane, and T. Gunawardane, 2°.

Alluaudomyia fuscipes Wirth and Delfinado

New Record.—SRI LANKA: Uggalkaltota, 5.11.1970, D. Davis and W.
Rowe, light trap, 1°.

Alluaudomyia maculosipennis Tokunaga

New Record.—SRI LANKA: Matale Dist., Bandarapola, 12.v.1974, UVL
trap, Gans and Prasanna, | °.

Alluaudomyia marginalis Wirth and Delfinado

New Record.—SRI LANKA: Uggalkaltota, 5.ii.1970, D. Davis and W.
Rowe, light trap, 1°.

Alluaudomyia spinosipes Tokunaga

New Record.—SRI LANKA: Uggalkaltota, 5.11.1970, D. Davis and W.
Rowe, light trap, 2°.

Alluaudomyia xanthocoma (Kieffer)

New Record.—SRI LANKA: Keg. Dist., Kitulgala Resthouse, 3—5.11. 1979,
UVL trap, K. V. Krombein, P.B. Karunaratne, T. Wijesinhe, S. Siriwar-
dane, and T. Gunawardane, Id.

Genus Parabezzia Malloch

The genus Parabezzia has been considered as mainly a New World genus
with 18 known species from the Americas. There are four known Afrotrop-
ical species, falcipennis Clastrier (1960) from Zaire, insolita Vattier and
Adam (1966) from the Congo Republic, and obscura de Meillon and Wirth
(1981b) and stagni de Meillon and Wirth (198la) from South Africa. The

| discovery of a new species from Sri Lanka in the Oriental Region indicates
| that the genus is still more widespread than formerly suspected.

Parabezzia orientalis Giles and Wirth, New Species
Fig. |

Female holotype.—Wing length 0.90 (0.89-1.12, n = 5) mm; breadth 0.40
{0.39-0.48, n = 5) mm.

Head: Brown. Eyes well separated. Antennal pedicel light brown, flagel-
lum darker; verticils well developed on all segments; length of flagellar seg-
ments (Fig. la) in proportion of 42-36-30-3 1-32-3 1-32-30-38-43-44-44-61, an-
tennal ratio 0.87. Palpus (Fig. 1b) light brown, 4-segmented; lengths of
segments in proportion of 10-20-34-21; 3rd segment with 2 (1-2, n = 5) cap-

824 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mee Sele Gas Clie Cas WAS ea a ee
a

<< —

Fig. 1. Parabezzia orientalis, female. a, Antenna. b, Palpus. c, Wing. d, Spermathecae. e,
Fifth tarsomere and claws of foreleg. f, Same of midleg. g, Same of hindleg.

itate sensilla; palpal ratio 3.29 (3.14-3.86, n = 5). Proboscis dark brown,
short, P/H ratio 0.43 (0.43-0.53, nm = 3); mandible with 12 (10-12, nm = 5)
large teeth, the distal ones slightly longer.

Thorax: Dark brown, prescutellar median area and large humeral areas |
paler brown. Legs dark brown from coxae to tibiae; hindtibial comb with 7 |
(7-8, n = 4) spines, the 3rd from the spur longest. Tarsi pale, hindbasitarsus |

and Sth tarsomeres light brown; claws (Fig. le—g) slightly unequal, long and
slender, the longer claw on each leg with proportional lengths of 29, 29, 28.

Wing (Fig. Ic): Hyaline; costa, base of subcosta, radial veins, medial |

stem, basal arculus, and base of Ist A slightly infuscated, all other veins
faint; costal sections I-II-III with lengths in proportion of 96-75-18, Rs 52,

Rs extending 0.95 (0.89-0.97, n = 5) of wing length; costa with slight pale |

basal swelling covered with fine microtrichia, also bearing 2 (2-4, n = 5)
large setae; fringe of costal setae longer than the width of costa, extending

to the wing tip, the fringe consisting of a series arising from the anterior §

edge of the costa. Halter pale.

Abdomen: Brown with segments 8, 9, and cerci darker. Spermathecae |
(Fig. 1d) 2, ovoid with short necks; unequal, measuring 0.071 by 0.054 mm |

and 0.061 by 0.053 mm; a tubelike rudimentary spermatheca present.

Male.—Unknown.

Distribution.—Sri Lanka.

Types.—All on slides in phenol balsam. Holotype 2, Sri Lanka, Mon-
aragala District, Mau Ara, 10 mi. E Edawalawa, 100 m, UV light, 24-26
Sept. 1977, coll. K.V. Krombein, P.B. Karunaratne, T. Wijesinhe, and M. |

|

VOLUME 84, NUMBER 4 825

Jayawiera (Type no. 76119, USNM). Paratypes, 42 , same data as holotype.
Holotype and 2 paratypes deposited in the National Museum of Natural
History, Washington, D.C. (USNM); | paratype will be deposited in the
Colombo National Museum, Colombo, Sri Lanka, and another in the Na-
tional Museum at the University of Sri Lanka at Peradeniya.

Discussion.—Parabezzia orientalis is the first species of Parabezzia to be
recorded from the Oriental Region.

Parabezzia falcipennis (male) and insolita (female) differ from orientalis
by the possession of a conspicuous beadlike swelling at the base of the
costa. In P. stagni the costa extends markedly past the end of vein R4 +
5, nearly attaining vein MI and the wing tip, the antennal ratio is 1.0, and
the spermathecae lack sclerotized necks. Parabezzia obscura is known only
from the male; it is marked very similar to P. stagni, but has a 3-segmented
palpus.

Genus Atrichopogon Kieffer

The genus Atrichopogon is difficult taxonomically and the Oriental species
are poorly known, although they are abundantly represented in nearly all
ceratopogonid collections. We have been unable to make specific determi-
nations of the Smithsonian Sri Lanka material except for the following species
which possesses remarkable tarsal claws.

Atrichopogon schizonyx Giles and Wirth, NEw SPECIES
Fig. 2

_ Female Holotype.—Wing Length 0.87 mm; breadth 0.43 mm.
_ Head: Brown. Eyes finely pubescent above, bare below; contiguous for
a distance equal to 3 facets. Antenna (Fig. 2a) light brown with well-devel-
| oped verticils on all segments; segment 15 with terminal papilla; flagellar
segments with lengths in proportion of 26-14-17-20-20-20-20-22-74-72-76-74-
100, antennal ratio 2.49; segments 3-5 disciform and closely appressed.
'Palpus (Fig. 2c) light brown; segments in proportion of 21-25-38-26-22; 3rd
“Segment swollen, spindle-shaped, with sensory pit moderately large and
shallow; palpal ratio 2.1. Proboscis brown, moderately long, sections A-B-
C (see Wirth, 1980) with lengths as 50-24-26; mandible (Fig. 2f) with 17 large
teeth becoming smaller proximad.

Thorax: Brown, pleural regions lighter. Legs with coxae brown, trochan-
ters light brown; femora and tibiae light brown with faint basal and apical
light bands; hindtibial comb with 8 spines; tarsi light brown, hindtarsal ratio
3.15; paired claws (Fig. 2g—i) of each leg with lateral claw bifid with small
tooth, mesal claw trifid, the middle tooth longest.

Wing (Fig. 2e): Light brown becoming paler caudad, veins darker. Ma-
crotrichia few on anterior veins. Costal ratio 0.73; 2nd radial cell 4x length
of Ist. Halter slightly infuscated.

826 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

See Se | SECESO(\)

(BX o S, GS | 0.@.

Fig. 2. Atrichopogon schizonyx. a, c, e-j, Female. b, d, k, Male. a, b, Antennae. c, d, |
Palpi. e, Wing. f, Mandible. g, Fifth tarsomere and claws of foreleg. h, Same of midleg. 1, |
Same of hindleg. j, Spermatheca. k, Male genitalia.

Abdomen: Light brown darkening toward tip. Spermatheca (Fig. 2j) 1,
partially collapsed, measuring 0.106 by 0.074 mm, ovoid with short slender |
neck.

Male Allotype.—Similar to female except for usual sexual differences.
Genitalia (Fig. 2k) with 9th sternum short, caudal margin convex with ap- |
proximately 17 setae; 9th tergum about 2 length of 9th sternum. Basistyle
stout, about as wide as long, extending slightly past tip of 9th tergum; basal
apodeme with slender base, swelling at midportion, then extending cephalad |
and tapering to a long point; dististyle stout basally, tapering quickly and
smoothly from base in a moderate curve and ending in a sharp point. Ae-
deagus with basal arms stout, arch about 2x as broad as long; main body |
with rounded shoulders, slightly broader than long, with a short caudal
process terminating in a cap.

Distribution.—Sri Lanka.

Types.—On slides in phenol balsam. Holotype 2, Sri Lanka, Ratnapura —
District, Ratnapura, 1-3 Feb. 1979, UVL trap, coll. K. V. Krombein, P. B.
Karunaratne, T. Wijesinhe, S. Sirtwardane, and T. Gunarwardane. Allotype
3, Sri Lanka, Uggalkaltota, 5 Feb. 1970, coll. D. Davis and W. Rowe, light
trap. Both deposited in USNM (Type no. 76120).

Discussion.—The name A. schizonyx refers to the unusually cleft claws, |

VOLUME 84, NUMBER 4 827

by which it can readily be distinguished from related species. Although
nothing is known of the life history of this species, the large mandibular
teeth and cleft claws suggest that it is an insect parasite.

Atrichopogon schizonyx appears to be an ordinary member of the genus
without any noteworthy features except for the unusual condition of the
tarsal claws. The usual condition for the tarsal claws in Atrichopogon is for
the female claw to be simple on the end, but a considerable number of
species have the claws slightly bifid at the very tip or each bearing a slender
spur partway down on the side. The male claws are usually both slightly
cleft at the tip. Tokunaga, however, has described three Atrichopogon species
departing from this pattern. In A. gressitti Tokunaga, (in Tokunaga and
Murachi, 1959), known only from the male from the Caroline Islands, the
claws are ‘‘stout, with tips bifid and each with a stout median tooth on the
lateral side.”’ In A. unguis Tokunaga (1962) from the Yaeyama Islands, the
female claws are ‘‘cleft into 3 parts, lateral | small, other 2 equal,’’ and
male ‘‘claws bifid at ends.”’ In A. xanthopygus Tokunaga (1962) from the
Yaeyama Islands, the male claws are all **‘equally bifid and each with a small
tooth on lateral side.”’

LITERATURE CITED

Clastrier, J. 1960. Notes sur les Cératopogonidés. X. Cératopogonidés de la République du
Congo (2). Arch. Inst. Pasteur Alger. 38: 258-298.

Delfinado, M. D. and D. E. Hardy. 1973. A catalog of the Diptera of the Oriental Region.
Vol. 1 Suborder Nematocera. The University Press of Hawaii, Honolulu. 618 pp.

Giles, F. E., W. W. Wirth, and D. H. Messersmith. 1981. Two new species of biting midges
and a check list of the genus Culicoides (Diptera: Ceratopogonidae) from Sri Lanka.
Proc. Entomol. Soc. Wash. 83: 537-543.

Meillon, B. de and W. W. Wirth. 198la. Subsaharan Ceratopogonidae (Diptera) VI. New
species and records of South African biting midges collected by A. L. Dyce. Ann. Natal
Mus. 24: 525-561.

—. 1981b. Subsaharan Ceratopogonidae (Diptera). VII. The biting midges of the Kruger
National Park, South Africa, exclusive of the genus Culicoides. Ann. Natal Mus. 24:
563-601.

Tokunaga, M. 1962. Biting midges of the Ryukyu Islands (Diptera: Ceratopogonidae). Pac.
Insects 4: 153-217.

Tokunaga, M. and E. K. Murachi. 1959. Insects of Micronesia Diptera: Ceratopogonidae.
Insects Micronesia 12(3): 100-434.

Vattier, G. and J. P. Adam. 1966. Les Ceratopogonidae (Diptera) des Grottes de la Répub-
lique du Congo (Brazzaville). Ann. Speleol. 21: 711-773.

Wirth, W. W. 1980. A new species and corrections in the Africhopogon midges of the
subgenus Meloehelea attacking blister beetles (Diptera: Ceratopogonidae). Proc. Ento-
mol. Soc. Wash. 82: 124-139.

Wirth, W. W. and M. D. Delfinado. 1964. Revision of the Oriental species of Alluaudomyia
Kieffer (Diptera, Ceratopogonidae). Pac. Insects 6: 599-648.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 828-844

BIOLOGY AND IMMATURE STAGES OF SETACERA ATROVIRENS,
A GRAZER OF FLOATING ALGAL MATS
(DIPTERA: EPHYDRIDAE)'

B. A. FOOTE

Department of Biological Sciences, Kent State University, Kent, Ohio
44242.

Abstract.—Information is presented on the life history and larval feeding
habits of Setacera atrovirens (Loew), a common and widely distributed
shore fly species in lentic freshwater habitats. Adults and larvae are asso-
ciated with floating algal mats where the larvae feed rather non-selectively
on algal cells, including those of various blue-green genera. The egg, mature
larva, and puparium of S. atrovirens are described and illustrated.

Miscellaneous observations on the habitat distribution, life cycle, and
larval feeding habits are given also for S. durani (Cresson) and S. pacifica
(Cresson).

The family Ephydridae, shore flies, consists of at least 1200 species in the
world (Rohdendorf, 1974), with over 400 species in 68 genera being recorded
from America north of Mexico (Deonier, 1979). This vast fauna is reflective
of the large amount of adaptive radiation that has occurred within this family
of wetland-inhabiting flies. Among the radiations into different habitats has
been the invasion of floating algal mats by at least three Nearctic genera of |
the tribe Ephydrini of the subfamily Ephydrinae. Adults of species of Cir- |
rula, Ephydra, and Setacera typically are found on the surface of the shal-
low water of lake margins, ponds, and marshes. Larvae of several of these |
species have been collected repeatedly from floating or partially submerged |
mats of filamentous algae, although they may be actually feeding micropha- |
gously on the interstitial unicellular algae growing within the filamentous |
matrix (Brock et al., 1969; Collins, 1975; Simpson, 1973).

The biology, life history, and larval feeding preferences of certain of the |
algal mat-inhabiting species have been elucidated previously. Aldrich (1912), |
Beyer (1939), Brock et al. (1969), Collins (1975, 1977, 1980a, 1980b), Hennig |

' Research supported by NSF grant DEB-7912242.

VOLUME 84, NUMBER 4 829

(1943), Johannsen (1935), Nemenz (1960a, 1960b), Ping (1921), Simpson (1973,
1976, 1979), Tragardh (1903), Wirth (1971, 1975), and Zavatarri (1921) have
presented information on species of Ephydra. Apparently the only refer-
ences to the immature stages of Cirrula are those of Mathis and Simpson
(1981) and Simpson (1973) dealing with the life history of C. gigantea Cres-
son, a species inhabiting algal mats in salt marshes along the northeastern
coast of North America. The only author who has published biological in-
formation on Nearctic species of Setacera is Johannsen (1935), who illus-
trated the immature stages of S. atrovirens (Loew) and S. needhami Johann-
sen. Beyer (1939) discussed in considerable detail the ecology, life cycle,
and larval morphology of an European species, S. micans (Haliday) (as
Ephydra).

The genus Setacera currently contains eight species in the Nearctic Re-
gion (Mathis, 1982), one in the Neotropics (Wirth, 1968), one in the
Afrotropical Region (Cogan, 1980), and five in the Palaearctic Region (Wirth,
1975). Setacera atrovirens is widely distributed in North America, occurring
between 55° and 105° west longitude and 38° and 49° north latitude (Mathis,
in press). In contrast, S. durani (Cresson) has been recorded primarily in
the Southwest, and S. pacifica (Cresson) apparently is known only from the
western states and provinces (Wirth, 1965).

This publication is the fourth in a series of papers devoted to ephydrid
species whose larvae can utilize blue-green algae (Foote, 1977, 1981la, 1981b).
The present paper gives life history data and discusses the larval feeding
habits of S. atrovirens, a common and widely distributed species in fresh-
water lakes, ponds, and marshes. Additionally, the egg, mature larva, and
puparium of that species are described and illustrated. Fragmentary biolog-
ical observations are also given for S. durani and S. pacifica.

MATERIALS AND METHODS

Most of the field observations on S. atrovirens were obtained near Kent,
Ohio, in Portage County. Supporting observations for this species and for
S. pacifica were obtained along the south shore of Flathead Lake east of
Polson, Montana, in Lake County. Most of the field work dealing with the
latter species was conducted at a highly alkaline pond located 5.0 miles
south of Ronan, Montana, on the Ninepipes Wildlife Refuge.

The laboratory rearings were carried out in an environmental chamber
programmed to give a photoperiod of 15L:9D and a temperature of 22°C
(+1°C). Monocultures of most of the algae used in the larval feeding tests
were obtained from the University of Texas Culture Collection of Algae
(Starr, 1978). Each algal monoculture was established on a nutrient agar
substrate in sterile petri plates, and feeding tests were performed as given
in Zack and Foote (1978). Monocultures of algae utilized in the tests, along
with their UTEX strain numbers, are listed below. Species lacking UTEX

830 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

numbers were obtained from the phycology laboratory at Kent State Uni-
versity.

Cyanophyceae

Anabaena variabilis (B-377)
Anabaena sp.
Cylindrospermum sp. (LB-942)
Gloeocapsa sp. (795)
Lyngbya sp.

Nostoc commune (584)
Oscillatoria chalybea (B-386)
Oscillatoria tenuis (B-428)
Phormidium sp. (1540)
Synechococcus leopoliensis (625)

Chlorophyceae

Chlamydomonas sp.
Chlorella vulgaris (29)

Bacillariophyceae
Navicula pelliculosa (668)

Chrysophyceae
Botrydiopsis alpina (295)

LIFE HISTORY OF SETACERA ATROVIRENS

Adults of Setacera atrovirens, like those of most other species of the tribe
Ephydrini, are typically found on the surface of shallow water of small
ponds, marshes, and lakes. They have elongated tarsi and straight claws,
adaptations which permit adults to move over the surface film in a manner
resembling that of the water striders of the family Gerridae (Hemiptera).
The mouthparts are rather broad and fleshy and apparently allow adults to
lap up microorganisms from the water surface. Deonier (1965) recorded
adults as common in the floating algal-mat habitat and rare on muddy shores
in lowa, while Scheiring and Foote (1973) collected specimens only in the
mud-shore habitat in Ohio. In contrast, I have found adults primarily on
floating algal mats and only rarely on muddy shorelines.

Deonier (1972) in a study of the gut contents of adult Ephydridae reported
that diatoms and other algal cells were about equally represented in guts of
S. atrovirens. | have observed adults of §. atrovirens applying their mouth-
parts to the surface of algal mats and subsequently found cells of numerous
algal genera in their alimentary canals. Adults fed readily on a variety of
algal monocultures in the laboratory but survived longest on a diet of Na-

i eee
a a

VOLUME 84, NUMBER 4 831

vicula pelliculosa, although good survival was obtained also on cultures of
Anabaena variabilis and Chlamydomonas sp. Adults died within 2 or 3 days
in pure cultures of Chlorella vulgaris.

The premating period varied between 2 and 4 days (n = 4). No overt
courtship behavior was observed, and males seemingly attempted to cop-
ulate with any suitably sized individual, including other males. Males were
rebuffed by all adults of the wrong species, by males of S. atrovirens, and
frequently even by females of that species. Mating behavior seemed to be
of the assault type described by Spieth (1974). Copulation lasted anywhere
from a few minutes to well over an hour. During mating the pair remained
relatively inactive, although females frequently fed intermittently. During
copulation the male assumed a position dorsal to the female and facing in
the same direction. The preoviposition period in 4 laboratory-reared females
ranged between 4 and 7 days and averaged 6 days.

An indication of the potential fecundity of S. atrovirens was obtained by
determining the number of developing eggs per ovariole and the total num-
ber of ovarioles in each of 8 females that were collected in northeastern
Ohio during August, 1979. The number of ovarioles per female varied be-
tween 28 and 36 and averaged 31.3. The number of detectable oocytes per
ovariole averaged 7.8 and ranged between 6 and 11. Assuming that each
ovariole is capable of producing 8 eggs during the life of a female and that
each female possesses 31 ovarioles, the average fecundity is around 250
eggs. The range would be between 168 (28 x 6) and 396 (36 x 11). Actual
egg counts obtained from field-collected and laboratory-reared females were
considerably less than the values given above. Two females collected in
nature deposited an average of only 40 eggs each in the breeding chambers.

In the laboratory rearings, females oviposited into monocultures of An-
abaena variabilis, A. sp., Cylindrospermum sp., Oscillatoria spp., and Na-
vicula pelliculosa. In contrast, no eggs were deposited in cultures of Chlor-
ella vulgaris. Eggs were widely scattered over the algal substrate and no
clumping was noted. They were generally oriented horizontally and occa-
sionally were completely imbedded in the alga mat. The eggs were elongate-
ovoid in shape, lacked terminal filaments, and possessed a distinct pinkish
cast. The incubation period varied between | and 2 days and averaged 1.8
days (nm = 45) in the laboratory rearings.

Eggs in nature were usually found in floating mats dominated by blue-
green algae. In Ohio, they were found in a small mat of Oscillatoria sp.
growing in a shallow, mud-bottomed rain pool having a surface area of less
than 2 m?. They were also taken abundantly in a mixed mat of Anabaena
and Spirogyra that was floating on the surface of a small marsh. In Montana,
eggs were discovered in a floating mat composed largely of species of Nos-
toc and Cylindrospermum located in shallow water along the south shore
of Flathead Lake. A few filamentous green algae were present in the mat

832 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

also, and unicellular interstitial algae, particularly diatoms, were abundant.
Also included in the mats were floating fragments of macrophytes, partic-
ularly debris derived from Typha and Potamogeton. A series of 3 petri dish
samples of the mat was taken along a transect extending from the sand
shore to the far edge of the algal growth. Very few Setacera eggs were
found within algae occurring on the moist shoreline sand, whereas they were
equally abundant in samples taken at the water’s edge and 2 m away from
the shore. Eggs were particularly common in growths of Nostoc but rela-
tively uncommon in colonies of Cylindrospermum.

Newly hatched larvae quickly began ingesting algae and seemingly pre-
ferred small unicells such as those of various species of diatoms. Older
larvae apparently could utilize a broader range of algal species and fre-
quently ingested trichomes of blue-green algae. A gut sample of a nearly
mature larva collected in an Oscillatoria mat in Ohio contained numerous
trichomes of that genus but also included considerable quantities of detritus.
The gut of a third-instar larva taken from the Nostoc-Cylindrospermum mat
at Flathead Lake, Mont. contained many trichomes of Nostoc, several frag-
ments of Cylindrospermum, numerous cells of Cosmarium, and a few spec-
imens of Oocystis, Scenedesmus, Navicula, and Pediastrum. Whether all
these algal taxa were being digested and assimilated was not determined.
To determine whether larvae preferred Cylindrospermum or Nostoc, 5 sec-
ond instars were placed in the center of a petri dish which contained 2
samples of each of those 2 genera. The observations began at 3 PM and
ended 6 hours later. By the end of the first hour 4 larvae were on Nostoc
and only one was on Cylindrospermum, a situation that did not change
during the subsequent 5 hours. A second test involved Anabaena sp., the
larval food of S. pacifica at a highly alkaline (pH 9.4) pond located south
of Ronan, Mont. Ten second- and third-instar larvae of S. atrovirens were
placed in a dish containing Anabaena, and 10 others were placed in Nostoc
collected at the Flathead Lake site. Larvae seemingly fed equally well in
both algal genera, suggesting that larvae of these 2 Setacera species are not
trophically segregated.

Examination of floating balls of Nostoc collected along the south shores
of Flathead Lake revealed that the algal colonies were noticeably damaged
due to larvae burrowing through the gelatinous matrix. The colonies became
riddled with holes and soon assumed a shredded, frayed appearance. Shortly
thereafter, they lost any semblance of cohesiveness and disappeared.

Larvae frequently fed while completely submerged but seemingly had to
return to the water surface periodically to place the posterior spiracles in
contact with air. This generally presented no problem, as algal mats in
nature typically floated at or near the surface. Similar behavior was noted
in S. micans in Europe by Beyer (1939).

Table | summarizes data obtained in laboratory feeding tests utilizing

VOLUME 84, NUMBER 4 833

Table |. Results of larval feeding tests for S. atrovirens using different algal monocultures.

Percent Reaching Different Life Stages

Alga n 21 3L Pp A
Cyanophyceae
Anabaena variabilis 30 100 100 97 97
Anabaena sp. 10 100 80 80 80
Cylindrospermum sp. 30 100 70 50 40
Gloeocapsa sp. 30 97 83 40 0
Lyngbya sp. 30 97 90 46 27
Nostoc commune 30 100 33 20 20
Oscillatoria chalybea 30 43 7 0 0
Oscillatoria tenuis 30 100 100 97 47
Phormidium sp. 30 100 97 40) 7
Synechococcus leopoliensis 30 100 93 10 0
Chlorophyceae
Chlamydomonas sp. 20 100 85 35 20
Chlorella vulgaris 30 0 0 0 0
Bacillariophyceae
Navicula pelliculosa 30 100 100 77 50
Xanthophyceae
Botrydiopsis alpina 10 100 10 10 0

unialgal cultures. Larvae completed development and eventually produced
adults in monocultures of certain blue-green species as well as in cultures
of Chlamydomonas sp. and Navicula pelliculosa. In contrast, no larval
development occurred in cultures of Chlorella vulgaris. Interestingly, not
all species of blue-greens permitted development, as no adults were obtained
in cultures of Synechococcus leopoliensis, Gloeocapsa sp., and O. chaly-
bea. The Oscillatoria tests were patricularly intriguing in that O. tenuis
seemed quite satisfactory as a larval food, whereas O. chalybea gave very
poor growth.

Table 2 presents data on the duration of the larval and pupal periods in
those monocultures that permitted complete development. Developmental
times were shortest in A. variabilis (14 days); somewhat longer in Anabaena
sp., Cylindrospermum sp., Chlamydomonas sp., Lyngbya sp., Phormidium
sp., and N. pelliculosa (17-21 days), and longest in Nostoc commune (24.9
days).

Shortly before forming puparia, larvae moved away from the algal colo-
nies and sought out floating or slightly submerged stems or narrow leaves
of such aquatic macrophytes as Potamogeton and Myriophyllum. Occa-
sionally, larvae attached themselves to filaments of green algae. The last
proleg was used to attach the mature larva to the support. Interestingly,

834 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Table 2. Developmental times in days for S. atrovirens using different algal monocultures.

Larval Period Pupal Period Combined Period
Algal x Ss x s x Ss

Anabaena variabilis (n = 29) 6.9 0.37 Ted 0.75 14.1 0.92
Anabaena sp. (n = 8) 12.8 0.46 7.5 0.93 20.3 1.39
Cylindrospermum sp. (n = 12) 1322 1.60 6.7 eit 19:9 0.90
Nostoc commune (n = 7) 17.4 0.98 eS 0.79 24.9 0.90
Lyngbya sp. (n = 1) 10.0 — 7.0 —_ 17.0 —
Phormidium sp. (n = 1) 10.0 — 7.0 — 17.0 —
Chlamydomonas sp. (n = 1) 14.0 — 7.0 — 21.0 —
Navicula pelliculosa (n = 10) 9.6 0.84 7.4 0.84 17.0 leds

many puparia were formed below the water surface and thus out of contact
with atmospheric air. Submergence had no apparent effect on pupal devel-
opment, as numerous adults were obtained from puparia that were as much
as 10 cm below the surface. Under laboratory conditions, the pupal period
lasted about 7 days (Table 3).

Several successful attacks on Setacera larvae by larvae of water scav-
enger beetles (Hydrophilidae) were noted in field-collected samples of algal
mats. Hydrophilid larvae were quite abundant in nearly all of the mats
examined and probably served as the primary predator of the larval stages
of the fly, although nymphs of damselflies may also have affected larval
populations. Several adults of a species of chalcidoid wasp were reared from
a few puparia that had been collected in nature.

With a preoviposition period of 6 days, an incubation period of 2 days, a
larval period of 10 days, and a pupal period of 7 days, the life cycle can be
completed in approximately 25 days. This suggests that at least 4 generations
are produced in the northern states during a warm season lasting from late
May to late September. Overwintering probably occurs as adults in a state
of reproductive diapause (Beyer, 1939). Examination of 8 females collected
in nature on October 10, 1980, revealed undeveloped ovaries and numerous
fat deposits within the abdominal cavity. The latest record for a female in
northern Ohio was obtained on November 7, but no effort to discover over-
wintering adults in nature has been made. The first seasonal record for an
adult was obtained on April 5.

|

OBSERVATIONS ON OTHER SPECIES OF SETACERA
Setacera durani (Cresson)

Adults of this species were swept from the surface of an algal mat that
had developed in Sonoita Creek at Patagonia, Ariz. The stream was receiv-
ing effluent from a sewage lagoon with the result that blue-green algae and

VOLUME 84, NUMBER 4 835

Table 3. Life cycle data for S. atrovirens in northeastern Ohio. Rearings maintained at
22°C, with Anabaena variabilis serving as adult and larval food.

Flight Period Early April—early November (?)
Premating Period 2-4 days

Preoviposition Period 47 days

Incubation Period 2 days

Larval Period 5-7 days

Pupal Period 7-8 days

Length of Life Cycle 18-24 days

Fecundity 168-396 eggs/female
Number of generations/year 4+

other algal taxa had developed extensive growths in and along the water-
course. Several larvae and puparia were collected from the mat.

Setacera pacifica (Cresson)

Large populations of this western species were found during the summer
of 1973 and 1980 at highly alkaline ponds located at the Ninepipes Wildlife
Refuge near Ronan, Mont. The pH of the study pond averaged above 9
during both summers, and the water contained large quantities of carbonate
and bicarbonate ions. Algae were abundant, particularly species of the blue-
green genera Anabaena and Oscillatoria, and formed extensive floating mats
in the shallow nearshore water. Other algal genera found in the mats were

| Navicula, Pandorina, Euglena, Cosmarium, Staurastrum, Scenedesmus,
_ and Tetraedron. The shores became increasingly exposed as the pond dried
during the months of July and August, with the result that a wide band of
highly alkaline mud developed around the open water. Aquatic plants were
abundant, particularly hornwort (Ceratophyllum demersum L.), water-mil-
foil (Myriophyllum spicatum L.), and pondweed (Potamogeton pectinatus
L., P. zosteriformis Fernald).
_ Other species of Ephydridae repeatedly collected at the alkaline ponds
| were Paracoenia bisetosa (Coquillett), Lamproscatella muria Mathis, Sca-
| tella paludum (Meigen), Scatophila despecta (Haliday), Discocerina obscu-
rella (Fallén), and Hydrellia spp. Occasionally, Psilopa olga Cresson and
Philotelma alaskense Cresson also appeared. The commonest species was
P. bisetosa, which occurred by the thousands on the muddy shorelines.
Except for a species of Hydrellia, whose larvae mined the leaves of pond-
_ weed, all of the associated species were far more abundant on shoreline
muds than on the floating algal mats. To determine the microspatial distri-
| bution of adults of P. bisetosa and S. pacifica, 5 pan traps containing water
and a detergent (Grigarick, 1959) were placed along a transect perpendicular
| to the shoreline. One pan was placed on the mud shore, one was at the

836 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

shoreline, and 3 were on the floating algal mat 0.5, 1.0, and 2.0 m away
from the shoreline, respectively. They were positioned at the pond after
sunset and were collected 24 hours later. The results are given below:

Pan I (on shore mud): 29 Setacera, 67 Paracoenia

Pan II (at shoreline): 169 Setacera, 134 Paracoenia

Pan III (0.5 m from shoreline): 111 Setacera, 24 Paracoenia
Pan IV (1.0 m from shoreline): 48 Setacera, 12 Paracoenia

Pan V (2.0 m from shoreline): 102 Setacera, 45 Paracoenia

Although the 2 species showed considerable spatial overlap, S. pacifica was
most abundant on the floating mat while P. bisetosa reached its greatest
abundance on the organic-rich mud of the shoreline. Interestingly, very few
larvae of Paracoenia were found in the offshore algal mats. Conversely, no
Setacera larvae were present in samples of the shore mud, even though
they were fairly common in pieces of the algal mat that had been stranded
on the mud by dropping water levels.

Eggs and larvae of S. pacifica were abundant in the algal mats, particu-
larly in areas where Anabaena formed conspicuous gelatinous growths. The
eggs possessed a pinkish color and frequently were imbedded in the mat.
As in §. atrovirens, no clumping of eggs was noted. The incubation period
of 8 eggs lasted an average of 2.3 days in a laboratory rearing.

Larvae at the alkaline pool were not restricted to Anabaena colonies, as
numerous individuals were found within a mat composed largely of a species
of the filamentous green alga Rhizoclonium. Also present in this mat were
abundant cells of the diatom genera Navicula and Synedra and the desmid
genera Closterium, Cosmarium, and Staurastrum. Examination of the gut
contents of 2 third-instar larvae revealed a nearly pure assemblage of Cos-
marium Cells. Only a few cells of the other genera of desmids were present,
and there were no fragments of the alga genus Rhizoclonium. The gut of
another third-instar larva collected elsewhere in the mat contained numer-
ous cells of Cosmarium, plus representatives of Cymbella (diatom), Scene-
desmus (green alga) and the blue-green genera Chroococcus and Meris-
mopedia. Laboratory observations of a larva collected in the Rhizoclonium
mat showed that it moved along the filaments of that alga while scraping off
epiphytic algal cells with its mouthparts. |

Larvae collected from mats dominated by Anabaena definitely were in-
gesting trichomes of that alga as determined by examination of the contents
of 2 nearly mature larvae. However, other algal genera were also present
in the guts, such as Chroococcus, Cosmarium, and Staurastrum. Labora- |
tory-reared larvae also freely ingested trichomes of a species of Nostoc that
was serving as the primary food of S. atrovirens along the shores of Flathead
Lake.

As those of S. atrovirens, larvae occasionally fed below the water surface

EE aaa

VOLUME 84, NUMBER 4 837

but returned periodically to the surface to renew their air supply. However,
most larvae retained contact with atmospheric air via their posterior spira-
cles while feeding within the algal mats. The larval period of 6 laboratory-
reared larvae that had fed on Anabaena averaged 10 days.

Puparia in nature were affixed by their last abdominal proleg to thread-
like pieces of aquatic macrophytes or to filaments of algae. Two 0.3 m°
samples of potential substrate for puparia were taken in the pond whose
depth at the point of sampling was 0.2 meters. Sample I consisted largely
of algal filaments, whereas sample II contained both algal filaments and
several leaves and stems of Ceratophyllum, Myriophyllum, and Potamoge-
ton. Sample I contained 11 larvae and 3 puparia, whereas sample II pro-
duced 39 larvae and 28 puparia. These results plus general field observations
suggest that mature larvae rarely attach themselves to algal strands but
rather become affixed to narrow-leaved macrophytes. In sample II, all 28
puparia were attached to either Ceratophyllum or Myriophyllum, even though
Potamogeton was equally abundant. Similar results were obtained else-
where in the study site.

Puparia frequently were formed below the water surface and thus out of
contact with atmospheric air. The greatest depth for attachment was ap-
proximately 10 cm below the surface of water that was 30 cm deep. Adults
successfully escaped from these puparia. In the laboratory rearings, the
pupal period lasted 6-7 days for males and 7-8 days for females (n = 10).

Enemies of S. pacifica included parasitic nematodes that were in the
abdominal cavities of adults, hydrophilid larvae that preyed on the larval
stages, and small chalcidoid wasps that emerged from the puparia.

As in S. atrovirens, there were several generations a year, with consid-
erable overlapping of generations. The total life cycle could be completed
in 25—30 days.

DESCRIPTIONS OF IMMATURE STAGES OF SETACERA ATROVIRENS
Egg (Fig. 13).—Length: 0.68-0.75 mm, * = 0.70, width: 0.23-0.26 mm

¥ = 0.25 (n = 10). Elongate, ellipsoidal to cylindrical, both ends rounded;

chorion appearing minutely papillose micropyle terminal on small papilla,
micropylar end bluntly rounded, opposite end somewhat more tapering;
living embryo with faint pinkish color.

Mature third-instar Larva (Fig. 1).—Length: 10.5-14.0 mm, + = 12.0;
width: 1.7-2.0 mm, * = 1.8 (n = 5). Elongate, nearly cylindrical, both ends
tapering, posterior end telescoping and forming apically branched breathing
tube. Integument nearly concolorous and transparent. without distinct pat-
tern; surface densely covered by somewhat darkened, elongate spinules, no
spinules broadened (Fig. 6); integument of each segment bearing 2 kinds of
sensilla, each rayed sensillum with elongate base and 3—4 apical branches

838 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

BT A)

Figs. 1-10. Setacera atrovirens, mature larva. 1, Lateral habitus. 2, Lateral view of anterior
end. 3a, Rayed sensillum. 3b, Rosette sensillum. 4, Posterior spiracle. 5, Facial mask. 6,
Dorsum of segment 8. 7, Venter of segment 8. 8, Lateral view of cephalopharyngeal skeleton.
9, Dorsal view of parastomal bars and dorsal bridge. 10, Ventral view of hypostomal and
associated sclerites. Abbreviations: A = antenna; AHB = anterior hypostomal bridge; AP =
anal proleg; ASp = anterior spiracle; BT = breathing tube; CSP = circular sensory plate; DB =
dorsal bridge; DC = dorsal cornu; DS = dentate sclerite; HS = hypostomal sclerite; ILR = in-

VOLUME 84, NUMBER 4 839

(Fig. 3a), each rosette sensillum very short and bearing several narrow
branches apically (Fig. 3b). Prolegs present on venter of segments 5-12,
anal proleg largest, all prolegs bearing heavy, recurved crochets apically
(Fig. 7). Segment | (pseudocephalic) (Fig. 2, 5) frequently invaginated, bear-
ing antennae apicodorsally, circular sensory plate apicoventrally, and facial
mask ventrally; antennae elongate, appearing 2-segmented; circular sensory
plates with rims unbroken, each plate bearing few peg-like structures; facial
mask (Fig. 5) with several rows of comb-like structures. Segment 2 (pro-
thoracic) spinulose, bearing anterior spiracles posterolaterally; each spiracle
(Fig. 2) somewhat elongate and bearing 3-4 finger-like papillae apically.
Segments 3 and 4 very similar, densely spinulose, and bearing prolegs ven-
trally; each segment with 4 rayed sensilla (Fig. 1), one rosette sensillum
next to each dorsal, dorsolateral, and ventrolateral rayed sensillum. Segment
12 (caudal) bearing breathing tube posteriorly and anal proleg and perianal
pad ventrally (Fig. 15); breathing tube branched apically, with branches
capped by spiracular plates, each spiracular plate (Fig. 4) bearing 4 semi-
oval spiracular openings, circular scar, and 4 highly branched spiracular
processes; perianal pad at apex of anal proleg posterior to crochets, pad
nearly circular (Fig. 16); anal proleg large and somewhat curved forward
(Fig. 15).

Cephalopharyngeal skeleton (Fig. 8).—Length: 0.60-0.65 mm, * = 0.62

| (n = 5). Mouthooks paired, not connected dorsally; hook part slender in

lateral view, broader in dorsal view and bordered by several accessory
teeth; basal part narrow, with indication of elongate window. Dentate scler-

_ ite below basal part of each mouthhook narrowly triangular. Ligulate sclerite

narrow and elongate. Hypostomal sclerite (Fig. 10) composed of 2 lateral
rods seemingly connected by 2 narrow and strap-like hypostomal bridges,
anterior bridge somewhat broader than posterior bridge. Epistomal sclerite
broad, poorly pigmented, and fenestrate anteriorly. Parastomal bars (Fig.
9), arising near but not connected to anterior border of pharyngeal sclerite,
running forward between and dorsal to hypostomal sclerite, bars not con-

' nected to each other or to epistomal sclerite anteriorly. Inner longitudinal

rods of hypostomal sclerite converging anteriorly (Fig. 10). Pharyngeal sclerite
(Fig. 8) rather deeply pigmented; dorsal cornua slender, connected ante-
riorly by dorsal bridge; bridge narrow, with few small windows (Fig. 9);

oS

ner longitudinal rod; ISF = intrasegmental fold; LPP = lateral pharyngeal process; MH = mouth-
hook; OLR = outer longitudinal rod; P = proleg; PB = parastomal bar; PS = pharyngeal
sclerite; PHB = posterior hypostomal bridge; PSp = posterior spiracle; RaS = rayed sensil-
lum; RoS = rosette sensillum; SpO = spiracular opening; SpS = spiracular scar; VC = ven-
tral cornu; W = window.

840 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ventral cornua broader, each with large window posteroapically; subcircular
lateral process arising from anterior margin of pharyngeal sclerite above
parastomal bars; floor of pharyngeal sclerite with 9 ridges (Fig. 14), lateral
2 ridges incomplete, remaining 7 ridges each with well-developed lateral
lamellae apically that form filtering surface.

Puparium (Fig. 11, 12).—Length: 7.5-8.0 mm, (* = 7.8; width: 1.4-1.8
mm, < = 1.6 (n = 5). Somewhat swollen at midlength, with both ends ta-
pering and upturned; anterior end invaginated, bearing anterior spiracles
laterally; posterior end elongated into breathing tube. Prolegs on segments
5—11 somewhat reduced compared to larva but bearing distinct crochets
apically; anal proleg largest, curved forward and frequently attaching pu-
parium to narrow-leaved macrophyte or algal filament. Integument nearly
concolorous brown, without dorsal pattern, densely spinulose, no spinules
broadened. Perianal pad somewhat invaginated, subcircular.

DISCUSSION

Species of Setacera apparently are ecologically unified by their preference
for the floating algal-mat habitat. However, little information is available
concerning ecological isolation among the eight species occurring in North
America. There are indications that some species prefer lentic habitats that
have particular water chemistries. For example, S. atrovirens has been found
most frequently in freshwater habitats with low alkalinity, whereas S. pa- |
cifica apparently prefers waters of high alkalinity. Thus, these two species |
may be segregated by habitat. Certainly they do not seem to be segregated
trophically, as the larvae of both species readily fed upon and completed
development on a wide variety of algal foods. Furthermore, in laboratory
tests, larvae of S. atrovirens consumed Anabaena, the blue-green algal genus
heavily grazed in nature by S. pacifica. Similarly, S. pacifica developed
readily on cultures of Nostoc, the preferred larval food of S. atrovirens at
Flathead Lake. Equally obvious is that these two species are not separated |
temporally, as both occur throughout the warm season and are multivoltine.

Results of the larval food tests involving S$. atrovirens and 14 species of |
algae belonging to four classes (Table 1) suggest that larvae of this species
are rather generalized trophically. Nine species of 8 genera of 3 algal |
classes permitted complete larval growth. These data suggested that S. atro-
virens may be slightly less generalized than Scatella stagnalis (Fallén), as
that species was able to attain the adult stage on 13 algal species (Zack and
Foote, 1978). Moreover, the latter species was able to utilize certain algal
taxa that are seemingly unsuitable as larval foods for S. atrovirens (e.g.,
Gloeocapsa, Chlorella, Botrydiopsis). However, larvae of S. stagnalis de-
veloped very poorly on cultures of Cylindrospermum and Nostoc, genera
of blue-green algae which gave excellent growth in S. atrovirens.

VOLUME 84, NUMBER 4 841

13 14

Figs. 11-14. Setacera atrovirens, immature stages. 11, Dorsal view of puparium. 12, Lateral
view of puparium. 13, Egg. 14, Cross section of pharyngeal ridges. Abbreviations: M = mi-

| cropylar end; PR = pharyngeal ridge.

An interesting question concerns the ecological relationship of species of
Setacera to those of the two other genera of Nearctic Ephydrini associated
with floating algal mats. Species of Cirrula do not present much of a problem
as they are restricted to coastal salt marshes, habitats rarely (if ever) utilized
by North American species of Setacera. However, there are numerous ex-
amples of geographic and perhaps habitat overlap among species of Ephydra
and Setacera. Thus, S. atrovirens is sympatric with E. riparia Fallén in the

842 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Figs. 15-16. Setacera atrovirens, mature larva. 15, Lateral view of caudal segment. 16,
Perianal pad. Abbreviations: ASI = anal slit; other abbreviations as in Figs. 1-10.

northeastern states. There may be some habitat segregation in these two
species as E. riparia is most abundant in saline waters having relatively high
chloride ion concentrations, whereas S. atrovirens seemingly prefers fresh-
water habitats having low concentrations of salts. Other examples of geo-
graphic co-existence of species in these two genera occur in the western
states. Thus, S$. durani and E. packardi Wirth co-exist in southern Arizona,

and S$. needhami occurs with E. hians and E. packardi in Washington (Zack, |
personal communication). Once again, however, it is possible that each of |
these sympatric species is segregated into shallow-water habitats having |

different water quality. It is significant that neither W. N. Mathis (personal

communication), nor R. S. Zack (personal communication), nor I have col- |

lected species of both genera regularly in the same habitat.

Setacera larvae undoubtedly play several ecological roles in lentic eco- |

systems. The mechanical damage caused by the burrowing larvae probably
accelerates the disruption and eventual decomposition of floating algal mats.

Thus, the feeding activities of the larvae must be important in nutrient cy- |

cling. The larvae, particularly when abundant, are certainly involved in food

chains, as they are utilized by predacious and parasitoid insects and may |
be consumed by fishes and birds. More practically, the utilization of such |
heterocystous blue-green algae as Anabaena, Cylindrospermum, and Nos- |

VOLUME 84, NUMBER 4 843

toc by Setacera larvae and adults probably have unsuspected effects on the
nitrogen economy of shallow water habitats, as these algae are important
nitrogen-fixers (Peters, 1978). Finally, it should be mentioned that the bac-
terium responsible for Legionnaires disease has been found in floating mats
of blue-green algae, where it apparently utilizes algal extracellular products
as carbon and energy sources (Tison et al., 1980). This suggests that adults
of Setacera and other genera of alga mat-visiting Ephydridae could serve
as dispersal agents for this pathogenic bacterium.

ACKNOWLEDGMENTS

Appreciation is expressed to W. N. Mathis, Department of Entomology,
Smithsonian Institution, for his taxonomic aid and advice. All figures were
executed by Tana L. Smith.

LITERATURE CITED

Aldrich, J. M. 1912. The biology of some western species of the dipterous genus Ephydra.
J. N.Y. Entomol. Soc. 20: 77-90.

Beyer, A. 1939. Morphologische, okologische und physiologische Studien an den Larven der
Fliegen: Ephydra riparia Fallén, E. micans Haliday und Cdnia fumosa Stenhammar.
Kiel. Meeresforsch. 3: 265-320.

Brock, M. L., R. G. Wiegert, and T. D. Brock. 1969. Feeding by Paracoenia and Ephydra
(Diptera: Ephydridae) on the microorganisms of hot springs. Ecology 50: 192-200.

Cogan, B. H. 1980. 71. Family Ephydridae, pp. 655-669. Jn Crosskey, R. W., ed., Cataloque
of the Diptera of the Afrotropical Region. Brit. Mus. Nat. Hist., London.

Collins, N. 1975. Population biology of a brine fly (Diptera: Ephydridae) in the presence of
abundant algal food. Ecology 56: 1139-1148.

—. 1977. Mechanisms determining the relative abundance of brine flies (Diptera: Ephy-
dridae) in Yellowstone thermal spring eflluents. Can. Entomol. 109: 415-422.

—. 1980a. Population ecology of Ephydra cinerea Jones (Diptera: Ephydridae), the only
benthic metazoan of the Great Salt Lake (USA). Hydrobiologia 68: 99-112.

—. 1980b. Developmental responses to food limitation as indicators of environmental
conditions for Ephydra cinerea Jones (Diptera). Ecology 61: 650-661.

Deonier, D. L. 1965. Ecological observations on Iowa shore flies (Diptera, Ephydridae).
Proc. lowa Acad. Sci. 71: 496-510.

—. 1972. Observations on mating, oviposition, and food habits of certain shore flies

(Diptera: Ephydridae). Ohio J. Sci. 72: 22-29.

1979. Introduction—A prospectus of research in Ephydridae, pp. 1-19. Jn Deonier,
D. L., ed., First Symposium on the Systematics and Ecology of Ephydridae (Diptera).
North Am. Benthol. Soc.

Foote, B. A. 1977. Utilization of blue-green algae by larvae of shore flies. Environ. Entomol.
6: 812-814.

—. 198la. Biology and immature stages of Lytogaster excavata, a grazer of blue-green

algae (Diptera: Ephydridae). Proc. Entomol. Soc. Wash. 83: 304-315.

1981b. Biology and immature stages of Pelina truncatula, a consumer of blue-green
algae (Diptera: Ephydridae). Proc. Entomol. Soc. Wash. 83: 607-619.

Grigarick, A. A. 1959. Bionomics of the rice leaf miner, Hydrellia griseola (Fallén) in Cali-
fornia (Diptera: Ephydridae). Hilgardia 29: 1-80.
Hennig, W. 1943. Ubersicht tiber bisher bekannten Metamorphosestadien der Ephydriden,

844 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

mit Neubeschreibungen nach Material der Deutschen Limnologischen Sunda-Expedi-
tion. Arb. Morph. Taxon Entomol. Berlin-Dahlem 10: 105-138.

Johannsen, O. A. 1935. Aquatic Diptera. Part II. Orthorrhapha-Brachycera and Cyclorrha-
pha. N.Y. Agric. Exp. Stn. Mem. 177: 1-62.

Mathis, W. N. 1982. Studies of Ephydrinae (Diptera: Ephydridae), VII: Revision of the
genus Setacera Cresson. Smithson. Contrib. Zool. 350, 57 pp.

Mathis, W. N. and K. W. Simpson. 1981. Studies of Ephydrinae (Diptera: Ephydridae), V:
The genera Cirrula Cresson and Dimecoenia Cresson in North America. Smithson.
Contrib. Zool. 329: 1-S1.

Nemenz, H. 1960a. Beitrage zur Kenntnis der biologie von Ephydra cinerea Jones 1906
(Dipt.). Zool. Anz. 165: 218-226.

—. 1960b. On the osmotic regulation of the larvae of Ephydra cinerea. J. Insect Physiol.
4: 38-44.

Ping, O. 1921. The biology of Ephydra subopaca Loew. Mem. Cornell Univ. Agric. Exp.
Stn. 49: 557-616.

Peters, G. A. 1978. Blue-green algae and algal associations. Bioscience 28: 580-585.

Rohdendorf, B. 1974. The Historical Development of Diptera. Univ. Alberta Press. xv +
360 pp.

Scheiring, J. F. and B. A. Foote. 1973. Habitat distribution of the shore flies of northeastern
Ohio. Ohio J. Sci. 73: 152-166.

Simpson, K. W. 1973. Biology and immature stages of certain Ephydridae (Diptera) occurring
in North America. Ph.D. Diss., Cornell Univ.

—. 1976. The mature larvae and puparia of Ephydra (Halephydra) cinerea Jones and
Ephydra (Hydropyrus) hians Say (Diptera: Ephydridae). Proc. Entomol. Soc. Wash.
78: 263-269.

——. 1979. Evolution of life histories in the Ephydrini, pp. 99-109. In Deonier, D. L.,
ed., First Symposium on the Systematics and Ecology of Ephydridae (Diptera). North
Am. Benthol. Soc.

Spieth, H. T. 1974. Courtship behavior in Drosophila. Annu. Rev. Entomol. 19: 385-405.

Starr, R. C. 1978. The Culture Collection of Algae at the University of Texas at Austin. J.
Phycol. 14(Suppl.): 47-100.

Tragardh, I. 1903. Zur Anatomie und Entwicklungsgeschichte der Larve von Ephydra riparia
Fallén. Arkiv. Zool. I; 1-42.

Tison, D. L., D. H. Pope, W. B. Cherry, and C. B. Fliermans. 1980. Growth of Legionella
pneumophila in association with blue-green algae (Cyanobacteria). Appl. Environ. Mi-
crobiol. 39: 456-459.

Wirth, W. W. 1965. Family Ephydridae, pp. 734-759. In Stone, A., et al., eds., A Catalog
of the Diptera of America North of Mexico. U.S. Dep. Agric., Agric. Handb. 276.

—. 1968. Family Ephyridae, pp. I-43. In Papavero, N., ed., A Catalogue of the Diptera
of the Americas South of the United States. Dep. Zool. Sec. Agric. Sao Paulo, Brazil.

——. 1971. The brine flies of the genus Ephydra in North America (Diptera: Ephydridae).
Ann. Entomol. Soc. Am. 64: 357-377.

———. 1975. A revision of the brine flies of the genus Ephydra of the Old World (Diptera:
Ephydridae). Entomol. Scand. 6: 11-44.

Zack. R. S. and B. A. Foote. 1978. Utilization of algal monocultures by larvae of Scatella
stagnalis. Environ. Entomol. 7: 509-511.

Zavattari, E. 1921. Ricerche morfologiche ed etologiche sul dittero alofilo Ephydra bivittata
Loew. R. Comit. Talassogr. Ital. Venezia Mem. 83, 58 pp.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 845-848

CONICOBRUCHUS ALBOPUBENS (PIC) (COLEOPTERA: BRUCHIDAE)
AND ITS HOST CYAMOPSIS TETRAGONOLOBA (L.)
(LEGUMINOSAE), WITH THE DESIGNATION
OF A LECTOTYPE

JOHN M. KINGSOLVER

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C.
20560.

Abstract.—Conicobruchus albopubens attacks seeds of Guar (Cyamopsis
tetragonoloba) in India and Pakistan but has not yet been found in the guar-
growing areas of south central United States. To facilitate identification,
illustrations of salient characters are provided. The lectotype of albopubens
is selected.

Guar (Cyamopsis tetragonoloba (L.), also known as C. psoraloides DC.)
is widely grown in northern India and Pakistan. The legume fruits are used
as a vegetable, the foliage is fed to cattle, and fruits and foliage have me-
dicinal value. Growing plants provide shade for ginger plants, and foliage
is used as green manure and as a cover crop. In recent years, this plant has
been successfully grown in southwestern Oklahoma and north central Texas
but in fairly small acreages (50,000 acres in 1978). The seed produced from
U.S. cultivation and from more than 40,000 metric tons imported into the
U.S. from India and Pakistan is used in paper manufacturing, as a thickener
and binder in various processed foods and in cosmetics, and in a number
of other applications (Whistler and Hymowitz, 1979). Small acreages of guar
are also grown in Australia, Brazil, and South Africa.

On the Indian subcontinent, seeds of guar are attacked by the bruchid
Conicobruchus albopubens (Pic). Arora (1977) listed this plant (as Cy-
amopsis psoraloides) as the only host of C. albopubens, and most of the
specimens in the National Museum of Natural History (USNM) are labeled
with this host. One series of specimens from Coimbatore, S. India, however,
was reared from indigo seeds (Indigofera prob. tinctoria L.). Indigofera is
closely related to Cyamopsis.

S. R. Wadhi, National Bureau of Plant Genetic Resources, New Delhi,
recently sent to me for confirmation specimens of C. albopubens that his

846 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

i ut
AU)

Figs. 1-5. Conicobruchus albopubens. 1, Habitus, dorsal. 2, 6 genitalia, median lobe. 3,
3 genitalia, lateral lobes. 4, Pygidium. 5, Antenna, <.

VOLUME 84, NUMBER 4 847

laboratory had identified using Arora’s key to species (1977). They are iden-
tical to specimens sent to the USNM by Arora. Because this bruchid was
described from the Sudan, I requested the loan of type-material from the
British Museum, and R. D. Pope kindly sent two cotypes. In all details
including those in the male genitalia, the specimens from India and the type-
specimens are identical.

Although the geographical origin of C. tetragonoloba is obscure, Whistler
and Hymowitz (1979) suggest that the cultivar may have developed from
fodder supplies of C. senegalensis Guill. and Perrin brought into India by
Arabs for their horses during the eighth century. This theory, if correct,
could explain the Sudan-India distribution of the beetle, assuming that it
remained constant throughout the changes in the host plant from C. senega-
lensis to its present cultivar C. tetragonoloba. Only one species of Cy-
amopsis (C. senegalensis) is known from Sudan and Senegal (Brown and
Massey, 1929), but there is no evidence that the type-specimens of Coni-
cobruchus albopubens were associated with this plant in the Sudan.

The following short redescription of C. albopubens is given for conve-
nience although Pic’s description (1931) is unusually informative and Aro-
ra’s redescription (1977) is definitive despite the poorly reproduced habitus
drawings (Figs. IXA, IXB).

Conicobruchus albopubens (Pic)
Figs. 1—5

Bruchus albopubens Pic, 1931: 26.
Conicobruchus albopubens: Arora, 1977: 34.

Description.—Body length, 2.0—-2.5 mm; body width, 1.0-1.3 mm. Integ-
ument black except pro- and mesolegs and 3 proximal antennal segments
red to reddish yellow. Vestiture of densely placed, pure white slender setae
more or less evenly distributed over body except narrow white stripe in
middle of pronotum and middle of pygidium (Figs. | and 3); antenna slender,
serrate, longer in male than in female (Fig. 5); body shape as in Fig. 1;
metafemur without subapical ventral armature; male genitalia (Fig. 3) with
median lobe short, ventral valve acutely triangular, armature of internal
sac as shown (Fig. 2). (See also Arora, 1977, figs. 17 and 18.)

Type.—Lectotype 3d, with labels R. F. Medani, H. W. Bedford 173-
25, Cotton, Sudan Govt., Ent. Coll. C8065, Bruchus albopubens n. sp. (in
Pic’s handwriting), red circular type label. This specimen is hereby desig-
nated and labeled as Lectotype. In British Museum (Natural History),
London.

Discussion.—Although Conicobruchus albopubens is not known from the
United States, it remains a potential threat to the guar industry in this coun-

848 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

try, especially if seed stocks for breeding purposes were to be imported
from the Indian subcontinent. Imported industrial guar is in the form of
broken or split seeds, and this rules out almost completely the importation
of live immature or adult beetles.

Conicobruchus albopubens seems out of place among the other much
larger species of Indian Conicobruchus but appears to be more closely re-
lated to unidentified African species in the USNM collection. A thorough
taxonomic study of this Old World tropical genus would be necessary to
more clearly determine its relationships. Vats (1979) described the larva of
C. albopubens.

ACKNOWLEDGMENTS

My thanks to S. R. Wadhi for sending the specimens that launched this
small study, and to R. D. Pope for loan of type-material from the British
Museum collections. Mary Lou Cooley drew the habitus and pygidium.

LITERATURE CITED

Arora, G. L. 1977. Taxonomy of the Bruchidae (Coleoptera) of northwest India. Part 1.
Adults. Orient. Insects Suppl. No. 7, 32 pp.

Brown, A. F. and R. E. Massey. 1929. Flora of the Sudan. London. 502 pp.

Pic, M. 1931. Nouveautes diverses. Mel. Exot.-Entomol. 57: 1-36.

Vats, L. K. 1979. On the larvae of Conicobruchus Decelle (Coleoptera: Bruchidae). Res.
Bull. Panjab Univ. Sci. 27(1/2): 111-113.

Whistler, R. L. and T. Hymowitz. 1979. Guar: Agronomy, production, industrial use, and
nutrition. Purdue Univ. Press, W. Lafayette, Indiana. 124 pp.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 849-854

TWO NEW SPECIES OF HETEROSPILUS (HYMENOPTERA:
BRACONIDAE) FROM MEXICO BEING INTRODUCED
AGAINST THE COTTON BOLL WEEVIL,
ANTHONOMUS GRANDIS
(COLEOPTERA: CURCULIONIDAE)

PAUL M. MARSH

Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser-
vice, USDA, Beltsville Agricultural Research Center, Beltsville, Maryland
20705.

Abstract.—Descriptions are provided for two new species of the braconid
genus Heterospilus: annulatus, n. sp., from Corregidora, Tabasco, Mexico;
and megalopus, n. sp., from Cardenas, Tabasco, Mexico. Both species have
been reared from the cotton boll weevil, Anthonomus grandis Boheman,
and possibly from its close relative A. hunteri Burke and Cate, and are
being cultured and released in Texas.

The genus Heterospilus Haliday belongs in the braconid subfamily Do-
ryctinae, most species of which parasitize wood-boring beetle larvae. How-
ever, species of Heterospilus exhibit an unusually wide range of hosts cov-
ering three insect orders: Coleoptera (Anobiidae, Bostrichidae, Bruchidae,
Buprestidae, Cerambycidae, Curculionidae, Languriidae, Mordellidae,
Scolytidae); Lepidoptera (Gelechiidae, Incurvariidae, Olethreutidae, Pyr-
alidae); and Hymenoptera (Cephidae, Tenthredinidae). In nearly all of these
records, the hosts have cryptic boring habits in stems. In addition there are
two unusual records: one species is parasitic on larvae of the genus Mi-
crostigmus (Hymenoptera: Sphecidae); the other is recorded from galls of
Eurosta solidaginis (Fitch) (Diptera: Tephritidae) on ragweed. In spite of
this diverse host range, the genus is morphologically homogeneous. The
single character that distinguishes Heterospilus from all but one of the other
genera in the Doryctinae is the absence or weakness of the first intercubital
vein of the forewing, thus making the first and second cubital cells confluent
(Fig. 1). The genus has not been studied in the Western Hemisphere, but
apparently there are several species groups based on a variety of other
characters. None of these groups seems to be correlated with a particular
host group from the above list.

850 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

@

Figs 1, 2. Wings of Heterospilus. 1, H. megalopus. 2, H. annulatus.

The two species described here are being studied by James R. Cate, B.
J. Porter, and associates at Texas A&M University, College Station. The
species were collected at localities in the state of Tabasco, Mexico, where
they were definitely reared from larvae of Anthonomus grandis Boheman
in Hampea nutrica Fryxell. In addition to the specimens examined by me,
both species were collected at other localities in the state of Campeche
where they were reared from what was assumed to be A. hunteri Burke and
Cate (J. R. Cate, personal communication). However, since there is some
difficulty in distinguishing larval remains of A. grandis from those of A.
hunteri, which is restricted to Hampea trilobata Standley, I therefore as-
sume that the host of specimens of the Heterospilus species reared from
Anthonomus on Hampea trilobata in Campeche is A. hunteri.

Both species of Heterospilus have been released in areas around College
Station, Brownsville, Elsa, and Rosebud, Texas.

Heterospilus annulatus Marsh, New SPEcIEs
Bigs. 2.50: 75 310,14

Female.—Length of body, 2.5—3.5 mm; ovipositor, 0.50-0.75 mm. Color:
Head dark brown, face and mandibles often light brown to honey yellow,
palpi white; scape and pedicel yellow with longitudinal brown stripe later-
ally, flagellum brown except flagellomeres 16-19 which are white, numbers
16 and 19 sometimes partially to completely brown; thorax dark brown,
mesonotum, scutellum, propodeum, and anterior part of mesopleuron fre-
quently light brown; abdomen dark brown, terga | and 2 yellow medially,
terga 3-7 sometimes light brown, entire venter yellow; legs light yellow,
coxae and trochanters nearly white, apical tarsomeres brown; tegula yellow,
wing base white.

Head: Coriaceous,! vertex sometimes imbricate behind ocelli (Fig. 5),
face often rugulose; eyes large, malar space 4 eye height; ocellocular dis-

' All terms for sculpturing are based on Harris, 1979, Calif. Dep. Food Agric., Entomol.
Occas. Pap. No. 28, 31 pp.

VOLUME 84, NUMBER 4 851

eee
Gwe,
wewesne eS

7
ee,

Figs. 3-5. Heads of Heterospilus. 3, H. megalopus, dorsal view. 4, H. megalopus, anterior
view. 5, H. annulatus, dorsal view.

tance 2.5x diameter of lateral ocellus; antenna with 24-27 flagellomeres.
Thorax: Mesonotal lobes (Fig. 6) coriaceous, notauli scrobiculate and meet-
ing posteriorly in a wide porcate area, scutellar disc coriaceous but some-
times nearly smooth in small specimens, scutellar furrow with 4 or 5 cross
carinae; mesopleuron (Figs. 10, 11) coriaceous, subalar furrow scrobiculate;
propodem areolate-rugose with 2 small triangular coriaceous areas at base.
Abdomen (Fig. 7): First tergum short and broad, 1.5 as wide at apex as
long, costate-rugose, strongly arched with anterior elevation smooth; terga
2+3 with 2 transverse scrobiculate furrows, costate-rugose on basal /% be-

fore Ist transverse furrow, remainder coriaceous; remainder of terga cori-

|

-——$_—

aceous, terga 4 and 5 with transverse scrobiculate furrow at base (usually
hidden under apex of preceeding tergum); ovipositor about as long as terga

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

terospilus, dorsal views. 6, 7, H. annulatus. 8, 9,

abdomens of He

Figs. 6-9. Thoraces and
-galopus.

H. me

————— a

VOLUME 84, NUMBER 4 853

Figs. 10-13. Thoraces of Heterospilus, lateral views. 10, 11, H. annulatus. 12, 13, H.
megalopus.

1-3 or about % as long as forewing. Wings (Fig. 2): First segment of radius
% length of 2nd segment, 2nd segment of radius % length of 2nd intercu-
bitus, Ist intercubitus indicated only by light infuscation near radius; ner-
vulus slightly postfurcal.

Male.—Essentially similar to female; length of body 2-3 mm; antenna
with 22-26 flagellomeres; stigma in hindwing about as long as its distance
from wing base, broad, greater than 2 width of wing at stigma location.

Holotype.— , Corregidora, Tab., Mexico, August 1979, J. R. Cate col-
lector, ex. Anthonomus grandis. Deposited in USNM (National Museum of
Natural History, Washington, D.C.).

Paratypes.—24 2,25 ¢, same data as type. Deposited in USNM and Texas
A&M University.

Remarks.—This species is distinguished from all other described U.S.
species by the white annulus on the antenna in both sexes. It is similar to
annulicornis Muesebeck from Brazil, which also has a white annulus on the

854 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

female antenna, but annulatus is distinguished from annulicornis by the
yellow and brown markings on the abdomen and thorax, the brown stripe
on the scape and pedicel, and by the fact the male of annulicornis does not
have the white annulus on the antenna.

This species also is apparently similar to ashmeadi Shenefelt but the
unique holotype is missing both antennal flagella and some characters are
obscured by glue. However, ashmeadi appears to have a smooth meso-
notum, smooth notauli, and is entirely dark brown or black.

Heterospilus megalopus Marsh, New SPEcIEs
Figs 13. 458.9223

Female.—Length of body, 3.5 mm; ovipositor, 1.25 mm. Color: Head
honey yellow, antenna yellow, apical flagellomeres light brown, palpi white;
thorax brown, black markings around scutellum, prothorax sometimes light-
er brown; abdomen brown, often with dark brown or black on abdominal
terga 2-4, venter honey yellow; tegula and wing base yellow; legs light
yellow, nearly white.

Head: Vertex and frons strigate (Fig. 3), face rugulose, temples smooth;
eyes large (Fig. 4), malar space about !/12 eye height, temples about !/4 eye
width, face narrow, about 34 eye width; ocellocular distance equal to di-
ameter of lateral ocellus; antenna with 26 flagellomeres. Thorax: Mesonotal
lobes (Fig. 8) coriaceous, notauli scrobiculate, meeting posteriorly in a wide
rugose area; scutellar disc smooth, scutellar furrow broad and with one
median cross carina; mesopleural disc (Figs. 12, 13) smooth, subalar furrow
scrobiculate; propodeum rugose with 2 large triangular smooth areas at base.
Abdomen (Fig. 9): First tergum slightly longer than apical width, costate;
terga 2+3 costate on basal 34, smooth on apical 4, transverse sutures at
most indicated by weak lines laterally, remainder of abdominal terga smooth;
Ovipositor about % as long as abdomen or slightly less than 2 of forewing
length. Wings (Fig. 1): First segment of radius equal to 2nd, 2nd segment
of radius slightly longer than Ist intercubitus; nervulus postfurcal by about
its own length.

Male.—Essentially as in female; length of body, 2.5—3.5 mm; antenna with
22-24 flagellomeres; stigma in hindwing as long as its distance from wing
base, broad, nearly as broad as wing base near stigma.

Holotype.—? , Cardenas, Tab., Mexico, July 1979, J. R. Cate collector,
ex Anthonomus grandis. Deposited in USNM.

Paratypes.—3 ¢, 16 3, same data as holotype. Deposited in USNM and
Texas A&M University.

Remarks.—This species is similar to fasciatus Ashmead from St. Vincent
and Grenada because of the single carina in the scutellar furrow, but me-
galopus is distinctive by its large eyes, the heights of which are nearly 12
times as long as the malar space.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 855-857

NOTE
Insect Label Production Using a Personal Computer

A personal computer and an “‘inexpensive’’ dot matrix printer make it
possible to prepare small runs of insect and other labels in a very efficient
manner. The system I use is aCROMEMCO Z2D® computer with a Texas
Instruments 820® printer.

The TI820® compressed print option fits 9 characters on the width of a
0.5 inch wide label; three lines of print occupy 0.5 inch. I have found these
labels to be of the proper dimension for use of point-mounted Coleoptera.
The system is fast since the printer runs at 150 characters per second.

The print quality of the label is about that of a good typewriter except
that each letter is formed from a series of 7 vertical dots instead of being
fully formed as with a typewriter. The listing of the program will indicate
the general legibility of the print. The TI820® uses dual tractors with pin
feed so that only computer forms can be printed. I use a continuous com-
puter form which is card stock containing 3 x 5 inch cards such as is used
by libraries in cataloging their holdings.

The computer program presented here uses a BASIC interpreter and will
run on most personal computers (with BASIC) except that the procedure
to fill a string is unique to the CROMEMCO 16K BASIC® which was used
to develop and run the program.

The program assumes that input from the user comes from the keyboard
of the printer. A CRT terminal keyboard could be used for input but then
statements 340 through 360 of the program must be changed so that the
output is written to the printer instead of to the CRT.

The program produces eight (8) 10-spaced labels as a group with a blank
space between each label. The number of lines per label is here limited to
three but the number of lines and the width of the label can be changed by
the user upon modifying the program.

The user first specifies how many groups of 8 labels is wanted by supply-
ing a “REPLICATION FACTOR.”’ For instance, if he wants 80 labels he
would enter *‘10.’’ Next, the user supplies information for each line of the
label following the prompt from the program. A small label of one or two
lines can be prepared by entering a “**’’ as indicated by the program. After
producing the labels wanted the program loops back to the beginning and
asks for the next REPLICATION FACTOR. Any negative number, i.e. —1,
will stop the program.

A listing of the program and an example of its use is shown in Fig. 1.

856

SSSSSZEBSSq

260

SER SSS8SS838R

328

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Rem PROGRAM TO PRODUCE INSECT LABELS
Rem AUTHOR: 0D. G. Kissinger
Rem
Rem PRODUCES 8 LABELS ON A LINE EACH LABEL 9 CHARACTERS WIDE AND
Rem SEPARATED FROM ADJACENT LABELS BY A SPACE.
Rem
Rem SET UP TO PRODUCE LABELS WITH AT MOST 3 LINES
Rem
Dim A$(79),B$(79),C$(79),N$(9),Z7$(9) = Rem DIMENSION STRINGS
7${-1)=" "+Z$(-1) : Rem SET 7$ TO ALL BLANKS
Rem BEGIN MAIN LOOP OF PROGRAM - STATEMENTS 120 THROUGH 400
Input"WHAT REPLICATION FACTOR (EXIT WITH -1) ".G
If GO Then Stop
Input"DATA FOR LINE 1 ",N$(0,8) : Rem ENTER UP TO 9 CHARACTERS
N$(Len(N$))=Z$ = Rem SET UNUSED PORTION OF LINE TO BLANKS
A$(-1)=NS+A$(-1) = Rem PRODUCE 8 COPIES OF FIRST LINE
L=1 : Rem THERE IS AT LEAST ONE LINE IN THE LABEL
N$="""= Rem EMPTY THIS STRING
Input"DATA FOR LINE 2 (END LABEL WITH #) °,N$(0,8)
If N${0,0)="#" Then 310
L=2 : Rem THERE ARE TWO LINES IN THE LABEL
N$(Len(N$) )=Z$
BS (-1)=N$+B$(-1)
N$="" = Rem EMPTY THIS STRING
Input"DATA FOR LINE 3 (END LABEL WITH *) °,N$(0,8)
If N$(0,0)="#" Then 310
L=3 = Rem LABEL CONTAINS 3 LINES
NS(Len(N$) )=Z$
C$(-1)=N$+C$(-1)
Rem LOOPS TO PRINT G GROUPS OF LABELS EACH WITH L LINES
Print
For 1 To G
For K=1 To L
If K=1 Then Print A$
If K=2 Then Print BS
If K=3 Then Print C$
Next K
Next J
Print = Rem END GROUP OF LABELS WITH A BLANK LINE
Goto 120

————<——__

VOLUME 84, NUMBER 4 857

>run

WHAT REPLICATION FACTOR (EXIT WITH -1) 2
DATA FOR LINE 1 RIVERSIDE

DATA FOR LINE 2 (END LABEL WITH #) Bt)
DATA FOR LINE 3 (END LABEL WITH #) #

RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE

C0 CO CO C0 CO C0 CO BL
RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE RIVERSIDE
CO CO CO CO CO C0 PI) CO

WHAT REPLICATION FACTOR (EXIT WITH -1) -1
#44130 StopHtt
»>

Fig. |. Listing of the program and an example of its use. Labels shown are actual size.

David G. Kissinger, Department of Biostatistics and Epidemiology, School
of Health, Loma Linda University, Loma Linda, California 92350.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 857-858

NOTE

Ferrisia lobdellae, a New Name for Ferrisia setosa
(Lobdell, 1930) (Homoptera: Pseudococcidae)

Ferrisia setosa (Lobdell) is presently without a valid name. This mealybug
was originally discovered on the roots of Liguidambar styraciflua at Durant,
Mississippi, by Lobdell (1930. Ann. Entomol. Soc. Am. 23: 209-236) and
named Trionymus setosus. Ferris (1950. Atlas of the Scale Insects of North
America, Series V: The Pseudococcidae, Pt. I, pp. 85-95) treated it as a
valid species and transferred it to the genus Ferrisiana. But the generic
name Ferrisiana is now considered as an unjustified change for Ferrisia
Fullaway, 1923 (vide, Morrison and Morrison. 1966. U.S. Dep. Agric. Misc.
Publ. 1015: 1-206). Therefore, this species presently is called Ferrisia setosa

(Lobdell).
| However prior to Lobdell’s 1930 paper, Hempel (1900. Rev. Mus. Paul-
| ista, Sao Paulo 4: 386) described a mealybug on Ficus in Brazil as Dacty-
| lopius setosus, and he mentioned it again in a second publication (Hempel.
1900. Ann. Mag. Nat. Hist. (7) 6: 396). Hempel’s mealybug was referred to
as Pseudococcus setosus (Hempel) by Fernald (1903. Mass. Agric. Exp.
Stn. Spec. Bull. 88: 109) and by MacGillivray (1921. The Coccidae. Scarab
Co., Urbana, Illinois, p. 133), was later placed in the genus Ferrisia, and

858 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

presently has been sunk as a synonym of F. virgata (Cockerell) (vide, Mo-
hammed Ali. 1970, Indian Mus. Bull. 5(2): 108).

Thus the setosus of Lobdell (1930), which is a valid species, is a junior
secondary homonym of the setosus of Hempel (1900), and both are presently
included in the genus Ferrisia. There being no other available name for the
species discovered by Lobdell, Ferrisia lobdellae Varshney, NEw NAME, is
here proposed as a replacement name.

The holotype of Trionymus setosus Lobdell, 1930, is in the Collection of
the Entomological Museum, Mississippi State (Richard L. Brown, personal
communication). Four ‘‘type material’ slides are deposited in the Coccoi-
dea Collection, University of California, Davis; and one paratype slide (not
in good condition) is in the Collection of the National Museum of Natural
History, Beltsville, Maryland.

The replacement name is in honor of Ms. Gladys Hoke Lobdell, the dis-
coverer of the species.

Incidentally, it may also be pointed out here that Ferris (1950: 88) and
McKenzie (1967. Mealybugs of California with Taxonomy, Biology, and
Control of North American species (Homoptera: Coccoidea: Pseudococci-
dae). Univ. Calif. Press, Berkeley, p. 179) have made a minor unintentional
error by stating that Takahashi ‘‘in 1927°’ changed Ferrisia to Ferrisiana.
In fact, Takahashi proposed this change in 1929 (Trans. Formosa Nat. Hist.
Soc. 19(104): 425-431).

I am grateful to the Director, Zoological Survey of India, Calcutta, for
providing facilities and permission to publish this note. I also thank D. R.
Miller and M. B. Stoetzel, Systematic Entomology Laboratory, IIBIII, Agric.
Res. Serv.. USDA, Beltsville, Maryland, for making useful suggestions on
this manuscript.

Rajendra K. Varshney, Zoological Survey of India, 34, Chittaranjan Av-
enue, Calcutta, 700 012, India.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 858-859

NOTE

Distributional Records of Platypodidae (Coleoptera) in Maryland

In the recent Catalog of North American Platypodidae (S. L. Wood. 1979.
U.S. Dept. Agric., Agric. Handb. 529-141) no species are listed as occurring
in Maryland. Three species, Platypus compositus Say, P. flavicornis (Fa-

VOLUME 84, NUMBER 4 859

bricius), and P. quadridentatus (Olivier), are recorded from surrounding
states and would be expected to occur in Maryland.

From examining the collections of the Maryland Department of Agricul-
ture, University of Maryland, E. J. Ford, and C. L. Staines, the following
records were observed:

Platypus compositus: Prince Georges Co., Laurel, 24/VIII/1977; Wicom-
ico Co., 16/VIII/1976.

Platypus flavicornis: Wicomico Co., Koppers, 24/VIII/1976; Worcester
Co., 16/X/1980.

Platypus quadridentatus: Baltimore City, Leakin Park, 7/V1I/1976; Prince
Georges Co., College Park, 2/IV/1946; Somerset Co., Shelltown, 10/V/1969;
Worcester Co., Berlin, 9/XII/1977.

I thank C. Mitter (University of Maryland), and E. J. Ford for allowing
me to examine the collections under their care.

C. L. Staines, Jr., 3302 Decker Place, Edgewater, Maryland 21037.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 859-860

NOTE

An Oriental Fly Found in Trinidad (Diptera: Platystomatidae)

Among specimens of Richardiidae sent to me by Fred D. Bennett, Direc-
tor of the Commonwealth Institute of Biological Control, was found a female
specimen that I at first thought was a member of the widespread American
genus Senopterina. However, the specimen showed characters, one of which
was a well-developed mesopleural bristle, that forbade my placing it in Sen-
opterina. Further study soon revealed that it was Plagiostenopterina du-
biosa Malloch, 1931, Proc. U.S. Natl. Mus. 78 (art. 15): 13-15. The type
specimen from Singapore and others from that locality and from Colombo,
Sri Lanka; Mindanao, Philippines; Selangor Gombak Forest Reserve, Ma-
laya; Chiengmai, Thailand; and Sarawak are in the U.S. National Museum
and have been compared with the specimen Bennett took from his labora-
tory window in Curepe, Trinidad, in March 1982. Bennett had done some
very interesting rearing of Richardiidae in the laboratory, but inasmuch as
there was nothing in the laboratory from southern Asia that might have
yielded the platystomatid and as nothing is known about the life history of

860 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Plagiostenopterina species, we cannot even guess what brought the species
to Trinidad.

George C. Steyskal, Cooperating Scientist, Systematic Entomology Lab-
oratory, IIBITT, Agricultural Research Service, USDA, % National Museum
of Natural History, Washington, D.C. 20560.

ANNOUNCEMENT

Back issues of the Index-Catalogue of Medical and Veterinary Zoology
are available free of charge by requesting them from:

Index-Catalogue of Medical & Veterinary Zoology
Animal Parasitology Institute

Building 1180, BARC-East

Beltsville, Maryland 20705 USA

Users and libraries should complete their sets now as reprinting of back
issues is not planned.

A compendium of the world’s literature on human and animal para-
sitology, the catalogues are divided into seven parts: Part 1, authors;
Part 2, Protozoa; Part 3, Trematoda and Cestoda; Part 4, Nematoda and
Acanthocephala; Part 5, Arthropoda and Miscellaneous Phyla; Part 6,
Subject Headings and Treatment; Part 7, Hosts. Several Special Publica-
tions are also available: No. 1: Checklist of the Internal and External
Parasites of Deer; No. 2: Bibliography of Chagas Disease (1909-1969);
No. 3: Ticks and Tickborne Diseases; No. 4: Checklist of Types in the
U.S. National Parasite Collection; No. 5: List of Translations; No. 6:
Nematoda and Nematode Diseases (and its companion publication, a re-
printed edition of the Stiles and Hassall Roundworm Catalogue), and an un-
numbered Special Publication, Trematoda and Trematode Diseases.

|
|

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 861-862

Book REVIEW

The North American Grasshoppers. Volume I, Acrididae: Gomphoceri-
nae and Acridinae. By Daniel Otte. Harvard University Press, Cam-
bridge, Massachusetts. 275 pp., illus. 76 figs. 1981. Cost: $45.00, hard-
back.

In 1961 James A. G. Rehn and Harold J. Grant, Jr. published the first
systematic revision of the grasshoppers of North America. In that volume,
which appeared as Monograph No. 12 of the Academy of Natural Sciences
of Philadelphia, the authors focused only on the Tetrigidae, Eumastacidae,
Tanaoceridae, and Romaleinae, and dealt only with species occurring in
North America, north of Mexico. Daniel Otte succeeded Rehn and Grant
at the Academy of Natural Sciences and undertook the task of completing
the monograph. His approach has been drastically different from that of
Rehn and Grant and may well prove to be much more successful, not only
in scope and aesthetics, but also in the breadth of audience that will be
receptive to it.

This book is the first of a three-volume set on the Orthoptera of North
America. Volume II will cover the band-winged grasshoppers of the subfam-
ily Oedipodinae, and volume III will cover all the grasshopper groups not
covered in volumes I and II.

The format of this volume is well-organized, easy to read, and concise.
It treats all species of gomphocerine and acridine grasshoppers that have
been described from North America, including Canada, Mexico, and Central
America. Otte includes helpful identification keys for each genus and pro-
vides excellent figures and distribution maps as well. For each species, the
distribution, recognition characters, information on behavior, ecology, and
life history, and pertinent references are detailed. The phylogenetic rela-
tionships of genera are discussed as well.

Additional valuable information is compiled neatly within five appendices,
including taxonomic changes made in the book, a list of the genera and
species of Gomphocerinae and Acridinae and their synonymy, a comparison
of the various classification systems of the treated genera, and a pronunci-
ation guide for the genera. Also included is a glossary and fairly complete
list of references.

But the highlight of this text is the showy display of color habitus figures
of every species of gomphocerine and acridine grasshoppers in North Amer-
ica. These figures, rendered by the author in colored pencil and pastel,
permit nearly anyone to quickly identify a grasshopper simply by overview-
ing the plates, which are conveniently clustered in the center of the book.

862 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

There are, in fact, few detracting qualities about this book. Some of the
figures do not agree in number with the text (e.g., on p. 15, Fig. 19D should
read ‘*‘Figure 16D’’). Also, ‘‘carina’’ and ‘‘carinula’’ are incorrectly used
interchangeably in the text and on Fig. 26. These shortcomings are minimal,
however, compared with the tremendous utility this book will provide to all
persons interested in acridology.

David A. Nickle, Systematic Entomology Laboratory, ITBIII, Agricultur-
al Research Service, USDA, % National Museum of Natural History,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 862-863

Book Review

Faunal Affinities, Systematics and Bionomics of the Orthoptera of the
California Channel Islands. By D. C. F. Rentz and David B. Weissman.
University of California Publications, Entomology, University of Califor-
nia Press, Berkeley. Volume 94, xiv + 240 pp., 337 figs. 1982.

Because they provide a microcosmic view of the complexity of the nature
of environments, islands have a special appeal to zoologists and botanists
for studies in biogeography. The California Channel Islands are well suited
for such studies. The eight islands in the chain lie between 21 and 94 kilo-
meters off the coast of southern California. They are divided into two groups,
the northern four islands, which are close in proximity to one another and
were connected to the mainland about 400,000 years ago, and the southern
islands which are more distant from each other and lack early geophysical
affinities with the mainland.

David C. F. Rentz and David B. Weissman together have an extensive
background in studies of the Orthoptera of North America and are probably
the world’s most knowledgeable orthopterists of the California fauna. Their
joint interest in the Orthoptera began more than ten years ago, and in this
volume they have provided us with the first comprehensive study of the
insect fauna on this remarkable island chain. They have spent more than
six years collecting Orthoptera on the islands and the adjacent mainland
and, during the course of their collecting, have made detailed studies of
populations, behavior, and ecology of the insects.

Their research approach is diverse, employing several techniques to gain

a

VOLUME 84, NUMBER 4 863

information on the faunal affinities of the Orthoptera of the study area. For
example, testes were removed from adult male Orthoptera specimens in
each area to compare karyotypes of populations on the various islands.
Immature insects were reared to adulthood to establish their identity and to
obtain such information about the insect’s biology as the number of moults
during its development, the presence and duration of periods of diapause,
and the occurrence of parasites. Most of the sound-producing Orthoptera
were tape-recorded and their songs analyzed and described. Wind tunnel
tests of several Orthoptera were made to evaluate whether it were possible
for these species to have used flight to achieve their present distribution.
Finally, collecting sites on each island were described ecologically, and the
species collected at these sites were listed.

The authors then use their faunal survey to evaluate zoogeographic re-
lationships of the islands and the adjacent California mainland. They indi-
cate that 23% of the 52 Orthoptera of the Channel Islands are endemic and
that of the 39 species of nonendemic Island species, 34 also occur on the
adjacent mainland. They suggest that the endemism is autocthonous, that
species evolved anew on the islands and do not represent relic populations
of species that were once more widespread. What is important here is that
Rentz and Weissman offer a hypothesis or model based on their research
on Orthoptera that other researchers can employ or test.

The second half of this comprehensive volume is devoted to the system-
atics and bionomics of the island fauna of Orthoptera. An excellent key is
provided for the identification of all species on the islands and adjacent
mainland. For each species, recognition characters, geographic distribution,
karyotype, habitat (and often food preferences), seasonal occurrence, and
discussion are given. Twelve new species and ten new subspecies are de-
scribed for the first time. On the basis of calling songs, six species of crickets
of the genus Gryllus are identified, though they are not given names.

The excellent figures include photographs of collecting sites and karyo-
types and line drawings of island maps and morphological features of the
island species.

The work of Rentz and Weissman is a major contribution to the biogeog-
raphy of the Channel Islands. In scope and presentation of information
pertinent to the concepts of island biogeography, it is a model for other
researchers to follow.

David A. Nickle, Systematic Entomology Laboratory, IIBIII, Agricultur-
al Research Service, USDA, % National Museum of Natural History,
Washington, D.C. 20560.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 864-866

OBITUARY

LUELLA WALKLEY MUESEBECK
1905-198 |

Luella Walkley Muesebeck was born May 22, 1905, in McClure, Penn-
sylvania, as Luella May Walkley. Her father was a career Army man, and
the family moved frequently. Luella’s only brother, Charles, was born in
New Jersey and her only sister, Orlena, in Rhode Island. When Luella was
six years old, the family moved to the West Coast, her father having been
assigned to Fort Worden, in the Puget Sound area of Washington, and they
lived on the base. Here they remained until 1915 when they moved to New
York State where they lived until Luella’s father returned from Europe at
the end of the First World War and was reassigned to Fort Worden. The
family again lived on the base until 1923 when the father retired from the
Army and built a home for them at nearby Port Townsend.

Luella attended the high school in Port Townsend 1919-1923 and Western

VOLUME 84, NUMBER 4 865

Washington College of Education 1924-25. During the following several
years she taught in public schools in Burlington and Dabob, Washington,
and Crowheart, Wyoming. With the rest of the family still in Port Townsend,
she then attended the University of Washington in Seattle, graduating in
1933 with a major in zoology and a minor in journalism. Here she came
under the influence of Dr. Trevor Kincaid, head of the Department of Zo-
ology, and developed a keen interest in biology and natural history. She
participated in numerous field excursions involving study of water and shore
biology of the Puget Sound region and of the biology of the nearby Cascade
Mountains.

Soon after this she went to Washington, D.C., and for several years taught
in the public schools there. In 1942 she was appointed Naval Research
Analyst in the Office of Censorship, and in 1946 she transferred to the
Department of Agriculture, being appointed to a subprofessional position in
the Division of Insect Identification of the old Bureau of Entomology and
Plant Quarantine. Although originally assigned to the cataloging section of
the Division she was soon placed on the professional staff. Her first re-
sponsibility here was in bibliographical research, in which she excelled. In
this connection she coauthored with me certain sections of the Catalog of
the Hymenoptera of North America north of Mexico, published by The
Department of Agriculture in 1951 (U.S. Dep. Agric., Agric. Monogr. 2),
and also a work entitled “‘Type species of the genera and subgenera .. . of
the superfamily Proctotrupoidea,’’ published in 1956 (Proc. U.S. Natl. Mus.
105: 319-419). Her familiarity with the International Rules of Zoological
Nomenclature and the Opinions of the International Commission, together
with unusual aptitude in the interpretation and application of them, contrib-
uted significantly to the success of the latter project. Soon after this she was
appointed specialist in the hymenopterous family Ichneumonidae with re-
sponsibility for supplying needed identifications and conducting research in
the systematics of the group. Intermittently she also continued study of the
small coleopterous family Lathridiidae which she had begun while at the
University of Washington.

Luella retired in January 1969 and continued to make her home in the
outskirts of Silver Spring, Maryland, where she had more than two acres
of lawn, garden, and woodland. She was devoted and skillful in rose culture
and sometimes exhibited her roses in the annual exhibitions sponsored by
the Potomac Rose Society, winning a number of prizes. She knew and loved
the song birds and saw to it that those in her area had food in winter and
summer. Her knowledge of the local wildflowers also was exceptional; she
even cultivated some of these in a natural setting at the edge of her wood-
land.

For many years Luella had suffered from a respiratory ailment, which
became seriously aggravated in 1976. She attributed her worsening condi-

866 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

tion to the increased air pollution in her area, and eventually she decided to
return to the Puget Sound region of Washington where she had spent her
youth. Efforts to become readjusted and settled here weakened her further,
but she acquired a comfortable and nicely situated home on Camano Island.
She and I were married there in April 1980. Her health continued to worsen,
however, and on March 12, 1981 she suffered a stroke which was followed
by her death on March 21. Besides me, her husband, only a sister, Orlena
W. Braun, of Jensen Beach, Florida, survives her. Her passing left an es-
pecial void in the lives of both of us, but it also has grieved a large number
of friends for all who knew her loved her. Luella was a member of the
Entomological Society of Washington.

Carl F. W. Muesebeck, 7/5 North Ell Road, Camano Island, Washing-
ton 98292.

PROC. ENTOMOL. SOC. WASH.
84(4), 1982, pp. 867-872

SOCIETY MEETINGS

883rd Regular Meeting—January 7, 1982

The 883rd Regular Meeting of the Entomological Society of Washington
was called to order by President Margaret S. Collins at 8:00 PM on January
7, 1982 in the Naturalist Center, National Museum of Natural History.
Twenty-five members and three guests attended.

Minutes of the previous meeting were read and approved. No new mem-
bers were reported for the month.

President Collins introduced a motion to increase membership dues to
$13 and change Article 4, Section | of the Constitution to read ‘‘thirteen”’
instead of ‘‘eight,’’ as recommended by the treasurer at the previous meet-
ing. The motion was properly seconded and unanimously approved by voice
vote.

Membership Chairman Joyce Utmar produced the names of the persons
making up the membership committee. In addition to herself, they are Victor
Adler and Thomas Wallenmaier.

President-elect Manya Stoetzel noted that the annual banquet will be held
on June 3, 1982. Various alternate sites for the banquet were discussed.

The speaker for the evening was Jack E. Lipes of the Office of Interna-
tional Cooperation and Development, USDA, who presented a talk entitled
‘Selected Insects—Their Distribution, Economic Importance, and Biolog-
ical Control Potential.’’ Most of the talk centered on Central America and
included Mediterranean Fruit Fly, Citrus Blackfly, Citrus Whitefly, Coconut
Palm Weevil, and various Lepidoptera whose larval forms have urticating
setae. Slides of the Middle East were also shown.

There were no notes or exhibitions. Visitors were introduced and the
meeting was adjourned at 9:15 PM, after which refreshments were served.

Thomas E. Wallenmaier, Recording Secretary

884th Regular Meeting—February 4, 1982

The 884th Regular Meeting of the Entomological Society of Washington
was Called to order by President Margaret S. Collins at 8:00 PM on February
4, 1982 in the Naturalist Center, National Museum of Natural History. Sev-
enteen members and four guests attended.

Minutes of the previous meeting were read and approved.

The speaker for the evening was Dr. Luther P. Brown of the Department
of Biology, George Mason University. His talk was entitled ‘*Behavioral
Ecology and Life History of a Horned Beetle—Bolitotherus cornutus (Pan-

868 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

zer).’’ The behavior of this tenebrionid beetle was studied extensively by
Dr. Brown at the Mountain Lake Biological Station in Virginia, to test
whether resource defense polygamy was being practiced by members of the
population. Details were given on life history, including courtship behavior,
oviposition, and larval and pupal descriptions. This species was found to
feed on three species of bracket fungi, and the adult males are noted for
horns extending from the pronotum. An interesting question and answer
period followed the talk.

NOTES AND EXHIBITIONS

President Collins announced that a field research station has been estab-
lished at Kartabo, Guyana, and inquiries from interested scientists are wel-
come. T. J. Spilman announced the death of Marion R. Smith at age 87 on
December 29, 1981. Dr. Smith received degrees from Clemson College, Ohio
State University, and the University of Illinois, and from 1921 to 1937 taught
at Mississippi State College. He was a specialist on the systematics of ants
and from 1937 to 1964 worked for the U.S. Department of Agriculture, being
best known as the author of ‘“‘House Infesting Ants of the Eastern United
States.”’

Visitors were introduced and the meeting was then adjourned at 9:30 PM,
after which refreshments were served.

Thomas E. Wallenmaier, Recording Secretary

885th Regular Meeting—March 4, 1982

The 885th Regular Meeting of the Entomological Society of Washington
was called to order by President Margaret S. Collins at 8:00 PM on March
4, 1982 in the Naturalist Center, National Museum of Natural History.
Twenty-six members and nine guests attended.

Minutes of the previous meeting were read and approved. T. J. Spilman
read the names of the following applicants for membership:

Robert W. Kelley, Department of Entomology, Fisheries and Wildlife,
Clemson University, Clemson, S.C. 29631.

John D. Pinto, Department of Entomology, University of California, Riv-
erside, Calif. 92521.

President Collins read a letter from the Smithsonian Archives asking
whether the Society would want to place its inactive records in their custody.
The question will be brought up at the next executive committee meeting
and then voted on at a regular meeting subsequent to the former. President
Collins gratefully acknowledged a gift of $1000 from Bennet A. Porter. Pres-
ident Collins also noted that she is now the delegate from the Society to the

a ie
$e ——

a

VOLUME 84, NUMBER 4 869

Washington Academy of Sciences. The Hospitality Committee was given
an acknowledgment of appreciation for its dedicated service throughout the
year.

The speaker for the evening was Dr. Howard G. Sengbusch, Professor
Emeritus of Biology, State University College of Buffalo. His talk was en-
titled ‘‘Starting from Scratch, or Some Ectoparasites of Increasing Medical
and Veterinary Importance.’’ Dr. Sengbusch discussed various species of
lice and mites that are important ectoparasites of man. Recent studies on
the prevalence of the head louse, Pediculus humanus Linnaeus, in different
groups of children were discussed. Many excellent slides accompanied the
talk.

NOTES AND EXHIBITIONS

A member noted the death of Elmer L. Mayer, an entomologist with the
Department of Agriculture’s Plant Industry Station in Beltsville, Maryland,
for 33 years. Mr. Mayer passed away on February 4, 1982.

Visitors were introduced and the meeting was adjourned at 9:20 PM, after
which refreshments were served.

Thomas E. Wallenmaier, Recording Secretary

886th Regular Meeting—April 1, 1982

The 886th Regular Meeting of the Entomological Society of Washington
was called to order by President Margaret S. Collins at 8:00 PM on April
1, 1982 in the Naturalist Center, National Museum of Natural History. Thir-
ty-six members and twenty-two guests attended.

President-elect Manya Stoetzel reported details of the forthcoming annual
banquet.

Membership Chairman Joyce Utmar read the names of the following ap-
plicants for membership:

R. M. Hendrickson, Jr., Beneficial Insects Research Laboratory, USDA-
ARS, 501 South Chapel Street, Newark, Del. 19713.

H. G. Larew, Florist and Nursery Crops Laboratory, BARC-East, Belts-
ville, Md. 20705.

W. P. Mackay, 1300 Elmhurst Drive, El Paso, Tex. 79925.

M. J. Raupp, Department of Entomology, University of Maryland, Col-
lege Park, Md. 20742.

G. Stephens, Route 2, Box 29, Laramie, Wyoming 82070.

G. M. Stonedahl, Department of Entomology, Oregon State University,
Corvallis, Oregon 97331.

E. A. Thomas, 537 Wilson Bridge Drive, Apt. C2, Oxon Hill, Md. 20745.

870 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Minutes of the previous meeting were read and approved.

The speaker for the evening was Donald R. Davis, a Smithsonian Insti-
tution scientist. His talk was entitled ‘‘Entomological Research on Taiwan.”
Dr. Davis began his talk with a discussion of the biogeography of Taiwan.
He then discussed the details of his 1980 visit to Taiwan where he visited
several entomology research institutions. He also collected more than 21,000
Lepidoptera specimens and described the details of his collecting. Dr. Davis
also collected one specimen of a rare family of Lepidoptera, the Neopseus-
tidae, and he described the unique morphology of these small moths.

NOTES AND EXHIBITIONS

Dr. Barris exhibited local termite specimens.

Dr. Anderson displayed the newly published *‘Bark and Ambrosia Beetles
of North and Central America’ by Stephen L. Wood. This monumental
work represents the most complete treatment of this group of beetles; its
1300 pages cover more than 1400 species.

Visitors were introduced and the meeting was adjourned at 9:30 PM, after
which refreshments were served.

Thomas E. Wallenmaier, Recording Secretary

887th Regular Meeting—May 6, 1982

The 887th Regular Meeting of the Entomological Society of Washington
was called to order by President Margaret S. Collins at 8:00 PM on May 6,
1982 in the Naturalist Center, National Museum of Natural History. Twenty-
nine members and sixteen guests attended.

Membership Chairman Joyce Utmar read the names of the following ap-
plicants for membership:

Nathan Bacon, 14282 Molly Pitcher Hwy., Greencastle, Pa. 17225.

Tohko Y. Kaufmann, 1211 Crown Point Ave., Apt. 107, Norman, Okla.
73069.

Juan M. Mathieu Veillard, Hildago No. 1328 Pte., Col. del Valle, Cd
Obregon, Sonora, Mexico.

Michel Padiou, 8 Rue du Cdt. L. Bouchet, Cressely—Nagny les Ma-
meaux, 78470 Saint Remy Les Chevreuse, France.

George Butler Wilson, Entomology and Plant Pathology, University of
Tennessee, P.O. Box 1071, Knoxville, Tenn. 37901.

Richard Zack, Department of Entomology, Washington State University,
Pullman, Wash. 99164.

The speaker for the evening was Dr. Donald H. Messersmith of the De-
partment of Entomology, University of Maryland, whose talk was entitled

ee

VOLUME 84, NUMBER 4 871

‘Insect Collecting in Belize.’’ Dr. Messersmith led a group which camped
at the Desmond B. Slattery Field Biology Station on Blue Creek in Belize.
More than 25,000 insect specimens were collected. Dr. Messersmith de-
scribed the day to day living of the collecting expedition with many inter-
esting slides and lively anecdotes. He also showed slides of some of the
insects found as well as other forms of wildlife. At the conclusion of his
talk he introduced Warren Steiner and Joseph Anderson who accompanied
him on the trip and who spoke briefly on their collecting. Dr. Margaret
Collins also accompanied Dr. Messersmith on the expedition and she con-
cluded by describing her success collecting termites, especially members of
the genus Armitermes.

NOTES AND EXHIBITIONS

T. J. Spilman showed a new book entitled ‘‘The Gypsy Moth: Research
Toward Integrated Pest Management”’ edited by Charles C. Doane and Mi-
chael L. McManus. It is USDA Technical Bulletin 1584, 757 pages, pub-
lished in 1981 and sells for $30.00. This book contains a wealth of infor-
mation on every aspect of this forest pest.

G. Steyskal showed a new book entitled ‘*A Revisionary Study of Leaf-
mining Flies (Agromyzidae) of California,’’ published in 1981.

T. J. Spilman regretfully brought to our attention the recent death of J.
L. Gressitt, age 67, and his wife in a plane crash in China. Gressitt—a great
figure in entomology—was most noted for his works on cerambycid and
chrysomelid beetles. It was also noted that Maurice T. James, a Diptera
specialist, died recently in Pullman, Washington.

Visitors were introduced and the meeting was adjourned at 9:45 PM, after
which refreshments were served.

Thomas E. Wallenmaier, Recording Secretary

888th Regular Meeting—June 3, 1982

The Entomological Society of Washington and the Plant Pest Society of
Washington Joint Annual Banquet was held on June 3, 1982, in the Ches-
apeake Room on the College Park campus of the University of Maryland.
Manya B. Stoetzel and Albert B. DeMilo were banquet chairmen and Eu-
gene F. Wood was master of ceremonies. After the social hour and dinner
Dr. Allen L. Steinhauer, Chairman of the University of Maryland Depart-
ment of Entomology spoke on the subject ‘‘A Look at IPM in the Third
World.”

The banquet was attended by 105 persons. After the talk by Dr. Stein-
hauer drawings were conducted for door prizes. Door prizes were donated
by: William Bickley, Chevron Chemical Company, CIBA-GEIGY Corpo-

872 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

ration, John A. Davidson, John W. Kennedy Consultants, Inc., Mansanto
Agricultural Products Company, National Agricultural Chemicals Associa-
tion, and Kjell Sandved, Smithsonian Institution.

Table centerpieces were provided by Dr. John W. Neal, Jr., and Mr.

Bobby Taylor, Beltsville Agricultural Research Center, U.S. Department of
Agriculture.

Thomas E. Wallenmaier, Recording Secretary

PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON

MISCELLANEOUS PUBLICATIONS
Cynipid Galls of the Eastern United States, by Lewis H. Weld -_-...-._
Cynipid Galls of the Southwest, by Lewis H. Weld_—-........__._.-_
Bothspapers: on cynipid galls= 2 ee eee
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman -.....-

Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides,
DY day oR: clraver 22 oie Fe a oe ee ee ae

A Short History of the Entomological Society of Washington, by Ashley B.

Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by
<scorpe: C2 Steyskal 2a oe ee ee ee eee ee

Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae),
By: George: @.;Stey sical) st ee es Se ee

MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse.
USO a FA an ae ee re ee eee

No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by
Adam 'G. Boving. (907 .0- he ee ee

No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by
Paul Wilsont@man’ 1949 22 oe ee a eee

DE a a es ee ee ee

No. 5. A Classification of the Siphonaptera of South America, by Phyllis T.
JOHMSO US LO 7 a ee eee eee

No.6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P.
Murdoch and Hirost Takahasi: 1969) 2 Se

No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette
hide cy} (oF oad 87 Re ai. eo SEN USER evn tee ee a SEPA SES RWS a ie sais ES

No. 8. The North American Predaceous Midges of the Genus Pa/pomyia Mei-
gen (Diptera: Ceratopogonidae), by W. L, Grogan, Jr. and W. W. Wirth.
| NO 19 fo eS cee Rees mete 2 1 Ce RNa EAS oc ee Bort esi Coe Snel

No.9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Chris-
Lian PHONTPSON dO 6 Usa ee eee

No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume
in Honor of Curtis W. Sabrosky. Edited by Wayne N. Mathis and F.
Christian: Thompsons V98? 42... ee

1.00

1.00

2.00

$15.00

(out of

print)

15.00

15.00

15.00

15.00

11.00

12.00

10.00

Back issues of the Proceedings of the Entomological Society of Washington are available
at $25.00 per volume to non-members and $13.00 per volume to members of the Society.

Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are
allowed a discount of 10 per cent on all items, including annual subscriptions, that are
paid in advance. All orders should be placed with the Custodian, Entomological Society
of Washington, c/o Department of Entomology, NHB 168, Smithsonian Institution,

Washington, D.C. 20560.

CONTENTS

(Continued from front cover)
LAVIGNE, R. J.—Notes on the ethology of Neoscleropogon elongatus (Diptera:
Asilidae) in South Australia
MARSH, P. M.—Two new species of Heterospilus (Hymenoptera: Braconidae) from
Mexico being introduced against the cotton boll weevil, Anthonomus grandis
(Coleoptera: Curculionidae)

ORTH, R. E. and T. W. FISHER—A new species of Tetanocera Dumeéril from
Colorado (Diptera: Sciomyzidae)

SELANDER, R. B.—Larval development in blister beetles of the genus Linsleya
(Coleoptera: Meloidae)

SELANDER, R. B.—Sexual behavior, bionomics, and first-instar larvae of the Lauta
and Diversicornis groups of Epicauta (Coleoptera: Meloidae)

STARY, P. and P. M. MARSH—A new species of Trioxys (Hymenoptera: Aphidiidae)
parasitic on a pecan aphid

STARK, B. P. and K. W. STEWART—Oconoperla, a new genus of North American
Perlodinae (Plecoptera: Perlodidae)

VALLEY, K. R.—A new Liriomyza mining leaflets of black locust (Diptera: Agro-
myzidae)

WHEELER, A. G., JR. and E. R. HOEBEKE—Psallus variabilis (Fallén) and P.
albipennis (Fallén), two European plant bugs established in North America, with
notes on taxonomic changes (Hemiptera: Heteroptera: Miridae)

NOTES:
KISSINGER, D. G.—Insect label production using a personal computer
STAINES, C. L., JR.—Distributional records of Platypodidae (Coleoptera) in Mary-

STEYSKAL, G. C.—An Oriental fly found in Trinidad (Diptera: Platystomatidae)

VARSHNEY, R. K.—Ferrisia lobdellae, a new name for Ferrisia setosa (Lobdell,
1930) (Homoptera: Pseudococcidae)

BOOK REVIEWS:

NICKLE, D. A.—The North American Grasshoppers. Volume I, Acrididae: Gompho-
cerinae and Acridinae (Daniel Otte)

NICKLE, D. A.—Faunal Affinities, Systematics and Bionomics of the Orthoptera of
the California Channel Islands (D. C. F. Rentz and D. B. Weissman)

OBITUARY:

LUELLA WALKLEY MUESEBECK (by C. F. W. Muesebeck)
SOCIETY MEETINGS

ANNOUNCEMENT

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